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HH

THE

meOck i DINGS

OF THE

LINNEAN SOCIETY

OF

NEw SOUL VWALES

FOR THE YEAR

1945

VOL. LXXxX.

WITH THIRTEEN PLATES.
215 Text-figures and 2 Maps.

SYDNEY:
PRINTED AND PUBLISHED FOR THE SOCIETY BY

AUSTRALASIAN MEDICAL PUBLISHING CO. LTD.,
Seamer Street, Glebe, Sydney,

and
SOLD BY THE SOCIETY.
1945.

li

CONTENTS OF PROCEEDINGS, 1945.

PARTS I-II (Nos. 317-318).
(Issued 15th May, 1945.)

Pages.
Presidential Address, delivered at the Seventieth Annual General Meeting, 28th
March, 1945, by W. R. Browne, D.Sc. i-xxiv
Elections XXV
Balance Sheets for the Year ending 28th February, py Seen dU et ee 5.6.q1t.0.qn it!
Some Points in the External Morphology of the Pouch Young of the Marsupial,
Thylacinus cynocephalus Harris. By W. Boardman. (Plate i and three
Text-figures.) 1-8
Stenoporids from the Permian of New South Wales and Tasmania. By Joan
Crockford, M.Sc., Linnean Macleay Fellow of the Society in Palaeontology.
(Plates ii—iii and twenty-five Text-figures.) 9-24
The Cranial Nerves of Neoceratodus. By H. Leighton Kesteven, D.Sc., M.D.
(Four Text-figures.) 25-33
Correlation of Some Carboniferous Sections in New South Wales, with Special
Reference to Changes in Facies. By Alan H. Voisey, M.Sc. (Plates iv—v.) 34-40
Evidence of an Eustatic Strand-line Movement of 100 to 150 feet on the Coast of
New South Wales. By Wilson H. Maze, M.Se. (Four Text-figures and one
Map.) 41-46
On Australian Dermestidae. Part iv. Notes and the Description of a New
Genus and Four New Species. By J. W. T. Armstrong. (Three Text-
figures. ) 47-52

CONTENTS.

PARTS III-IV (Nos. 319-320).
(Issued 15th February, 1946.)

Relation of the Orchid Flora of Australia to that of New Zealand, with the
Description of a New Monotypic Genus for New Zealand. By H. M. R.
Rupp, B.A., and EH. D. Hatch

Geographic Variation in the Lizard Hemiergis decresiensis (Fitzinger). By
Stephen J. Copland, B.Sc. (Plate vi and five Text-figures. )

Revision of Australian Lepidoptera. Oecophoridae. xiii. By A. Jefferis
Turner, M.D., F.R.E.S. ..

Studies on Australian Marine Algae. ii. Notes extending the Known
Geographical Range of Certain Species. By Valerie May, M.Sc. ..

A Bryozoan Fauna from the Lake’s Creek Quarry, Rockhampton, Queensland.
By Joan Crockford, M.Sc., Linnean Macleay Fellow of the Society in
Palaeontology. (Twelve Text-figures.) :

Miscellaneous Notes on Australian Diptera. xi. Evolution of Characters in
the Order: Venation of the Nemestrinidae. By G. H. Hardy. (Two Text-
figures. )

Principal Microspore-types in the Permian Coals of New South Wales. By
J. A. Dulhunty, B.Se. (Plate vii and three Text-figures.)

Petrological Studies in the Ordovician of New South Wales. iii. The
Composition and Origin of the Upper Ordovician Graptolite-bearing Slates.
By Germaine A. Joplin, B.Sc., Ph.D., Linnean Macleay Feilow of the
Society in Geology. (Four Text-figures.)

On the Family Smarididae (Acarina). By R. V. Southcott, M.B., B.S. (Four
Text-figures. )

lii

Pages.

53-61

62-92

93-120

121-124

125-134

135-146

147-157

158-172

173-178

iv CONTENTS.

PARTS V-VI (Nos. 321-322).
(Issued 15th July, 1946.)

Pages.
The Hair Tracts in Marsupials. Part ii. Description of Species, continued.

By W. Boardman. (Plate viii and forty-three Text-figures. ) 179-202
Nitrogen Fixation in Leguminous Plants. vi. Further Observations on the

Effect of Molybdenum on Symbiotic Nitrogen Fixation. By H. L. Jensen,

Macleay Bacteriologist to the Society. (Plate ix and one Text-figure.) 203-210
Contributions to a Knowledge of Australian Culicidae. No. viii. By Frank H.

Taylor, F.R.E.S., F.Z.S. (One Text-figure.) 211-212
Notes on Some Fijian Mosses. By William Greenwood .. 213-218
Notes on Australian Mosquitoes. (Diptera, Culicidae). Part vi. The Genus

Tripteroides in the Australasian Region. By David J. Lee, B.Sc. (Plates

xii-xiili and sixty-nine Text-figures.) 219-275
Observations on the Morphology and Biology of the Subspecies of Anopheles

punctulatus Donitz. By A. R. Woodhill. (Three Text-figures and one

Map.) 276-287
Notes on New South Wales Orchids: A New Species and Some New Records.

By the Rev. H. M. R. Rupp, B.A. (One Text-figure.) .. 288-290
Catalogue of Reptiles in the Macleay Museum. Parti. Sphenomorphus pardalis

pardalis (Macleay) and Sphenomorphus nigricaudis nigricaudis (Macleay).

By Stephen J. Copland, B.Sc. (Plate xi and seven Text-figures. ) 291-311
Studies on Trombidiidae (Acarina). Some Observations on the Biology of the

Microtrombidiinae. By R. V. Southcott, M.B., B.S. (Three Text-figures.) 312-316
A Contribution to a Study of the Physiology of Decay in Apples. By Mary

Cash, M.Sc. (Ten Text-figures.) 317-327
The Diptera of the Territory of New Guinea. xiii. Family Tabanidae. Part i.

The Genus Chrysops. By Frank H. Taylor, F.R.E.S., F.Z.S. (Plate x and

two Text-figures. ) 328-332
Studies in the Metabolism of Apples. vi. Preliminary Investigations on the

Respiration of Sliced Apple Tissue. By Frances M. V. Hackney, M.Sc.,

Linnean Macleay Fellow of the Society in Plant Physiology. (Three Text-

figures. ) 333-345
Abstract of Proceedings See LET eG SO Ceara Mac Oa Sor XX1X—XXXil
Tist: Of MeEMbeCLSe ce oes seed) cect cas (eae Stay oe Salm at eres 0 any 2) eget EE RONCNCU TI RERGNG VATU
List of Plates .. XXXVii
List of New Genera and Species XXXVili
General Index .. XXXix—xli

IBRA

ANNUAL GENERAL MEBRTING.

WEDNESDAY, 28th Marcu, 1945.

The Seventieth Annual General Meeting was held in the Society’s Rooms, Science
House, Gloucester Street, Sydney, on Wednesday, 28th March, 1945.

Dr. W. R. Browne, President, in the Chair.

The minutes of the preceding Annual General Meeting (29th March, 1944) were
read and confirmed.

PRESIDENTIAL ADDRESS.

Although the shadow of war still lies dark upon the earth, so that researches in
Natural Science apart from those directed to the immediate purposes of the war-effort
are in partial eclipse, yet the events of the last few months give substantial grounds
for the hope that ere another year has passed the struggle, in Europe at any rate, will be
over, and that vigorous forward movement in various fields of research of the type
for which this Society stands will have been resumed.

It is pleasing to be able to record that, in spite of adverse circumstances, the Society
has succeeded in maintaining its wonted activities during the past year.

Nevertheless it is somewhat disquieting to find that attendances at the general
meetings have shown no signs of increasing, but if anything the contrary. This
tendency seems to be common to all our scientific societies, and various expedients
have been adopted to stimulate the flagging interest of members. It is desirable, of
course, to treat causes rather than symptoms, but in the present instance these
are rather hard to determine. It may be that the war is to blame, and that with. the
return of peace matters will improve, but it is possible that the root of the trouble
lies deeper, and some have suggested that in these days of intensive specialization,
when the old type of scientific man with very wide interests is a well-nigh extinct
species, a general scientific society such as ours is an anachronism. If this be so then
it is up to us to consider whether there are any means whereby our activities can be
modified or extended so as to bring the Society into more direct relation with the
scientific needs and aspirations of its members. This is a matter that should not be
the concern of the Council alone; it should exercise the minds of all who have the
welfare of the Society at heart, and I commend it to the earnest personal attention of
each and every one of you.

The Financial Statements show the General Account debit at 28th February, 1945,
as £620 18s. 8d. This is unusually high, but is accounted for by outstanding amounts
of interest on mortgages, together with the fact that the sum of £15,000 in the Fellowship
Fund was reinvested in June last in the Commonwealth loan which closed on 15th
October, and no interest on this investment was received during the Society’s financial
year ending 28th February, 1945. Practically all of the Society’s income is derived from
investments and consequently the income has shown a gradual reduction over the past
ten or fifteen years. During the year portion of a mortgage was repaid by the mortgagor,
and the reinvestment of this money (£15,000) in Commonwealth loans resulted in a
reduction in income of about £260 per annum. Hach loan conversion, under present
circumstances, results in decreased income. We should be reminded that the compara-
tively sound financial position of the Society is in large measure due to the fact that
during the decade 1920-1930 your Council invested a large portion of the surplus income
received while returns from investment were at a high level.

The concluding part of Volume lxix of the Society’s Proceedings was issued in
December. The complete volume (278 + xxviii pages, ten plates, 166 text-figures and

A

a
*

li PRESIDENTIAL ADDRESS.

1 map) contains twenty-five papers on various branches of Natural History and, in

_ addition, a memorial account of Alexander Greenlaw Hamilton. Though much smaller

than..in pre-war years, the volume is slightly larger than the preceding one, and the
number ‘of /papers submitted in recent months has shown a marked increase. The
number of, exchanges received from scientific societies and institutions during the

‘session. again showed a marked decline, the total amounting to only 664, compared with
'T,200, 749 and 878 for the three preceding years. Throughout the year the Society has

continued its co-operation with the Allied Geographical Section, South-west Pacific Area,
in the preparation of an annotated bibliography of all published material in Australia
dealing with the South-west Pacific.

Since the last Annual Meeting the names of six members have been added to the list,
three membei's have been lost by death, the names of two have been removed on account
of arrears of subscription, and six have resigned.

Charles Anderson, who died at Sydney on 25th October, 1944, was born in the Orkney
Islands on 5th December, 1876. He was educated at Stenness Public School and later at
Kirkwall Burgh School and then followed a brilliant academic career at the University
of Edinburgh. He had charge of the Ben Nevis Observatory when, in July, 1901, he
accepted the position of Mineralogist at the Australian Museum; thereafter for a
considerable period he devoted his attention mainly to mineralogy and crystallography,
and in 1908 was awarded the degree of Doctor of Science by the University of Edinburgh
for researches in this field. On 14th February, 1921, he was appointed Director of the
Australian Museum, a position which he held until his retirement on 31st October, 1940.
In the intervals of administrative duties he found time to pursue the study of vertebrate
palaeontology, one of his early interests. Dr. Anderson possessed a genial personality,
had a wide knowledge of many branches of science, and always took a keen and active
interest in the work of scientific societies. He frequently took part in discussions, and
his contributions, often illustrated by apt anecdote, were marked by shrewd insight no
less than by a whimsical humour.

He joined this Society in 1906, served as a member of Council from June, 1926, until
March, 1942, and was President in 1932-33. He had also been President and one of the
Honorary Secretaries of the Royal Society of New South Wales, President of the Anthro-
pological Society of New South Wales and of the Geographical Society of New South ©
Wales, while the American Museum of Natural History had elected him a corresponding
member.

Harrold Fosbery Consett Davis, who was a life member of the Society, was killed
in an aircraft accident in New Guinea on 12th December, 1944. He was educated at The
King’s School, Parramatta, and entered the University of Sydney in 1931, and after a
brilliant academic career, graduated in Science with First Class Honours in Entomology
in 1934 and Botany in 1935. In March, 1937, he was awarded a Linnean Macleay Fellow-
ship in Zoology, and was reappointed to a Fellowship in 1938 and 1939. However, in
February, 1939, he resigned his Fellowship to take up the position of Lecturer in Biology
in the New England University College at Armidale. As a Fellow Dr. Consett Davis
made a special study of the order Embioptera mainly from the taxonomic viewpoint, but
he also contributed materially to a knowledge of their bionomics. In addition, he devoted
some time to the study of the ecology of the Five Islands and of the plant ecology of
the Bulli district. In 1938 Council granted him leave of absence for three months, and
during this period he visited the United States, England, France and other European
countries where he made a special study of the taxonomy of the order Embioptera from
a world standpoint. He also made a study of the respiration of the aquatic beetles
belonging to the family Dryopidae. Dr. Davis was accepted for the R.A.A.F. in 1941,
later transferred to the A.I.F., and still later, was seconded to the Commonwealth
Scientific Liaison Bureau, for which he was conducting important scientific research at
the time of his death. Throughout the years of his army life, Dr. Davis continued to
carry on his study of the Embioptera, and he was awarded the degree of Doctor of
Science by the University of Sydney in'1943 for his entomological researches. Possessed
of outstanding ability in research, remarkable energy and an original mind, the death
of Dr. Davis at the early age of thirty-one is a very grave loss to the biological sciences.

PRESIDENTIAL ADDRESS. lil

Perey Charles William Shaw died in Sydney on 8th September, 1944, at the age
of 84 years. Mr. Shaw, who was a retired engineer, joined the Society in 1941. In
addition to several books on birds, he bequeathed to the Society an extensive series of
his own excellent paintings of native and exotic flowers.

Towards the close of the year the Council decided to reduce the size of the Society’s
suite of rooms by vacating three rooms and partitioning off two offices from the meeting-
room. This effected a considerable saving in rent without adversely affecting the
comfort and convenience of members or the efficiency of the staff.

It is of interest to record that Science House, of which this Society owns one-third,
is in a flourishing condition. All the rooms in the building are let; it is impossible to
meet the demand for the use of the halls for evening meetings, and already it is
frequently found that the large hall is quite inadequate for the audiences which desire
to use it. The need for expansion is obvious, and a deputation representing the three
owner-bodies has already waited on the responsible Minister to ask that the Government
’ make a grant of the land adjoining Science House to provide for future additions to the
building.

During the past year the Council has taken an active part in a number of matters
connected with the preservation of the native flora and fauna. On the invitation of the
Board of Trustees, this Society, in conjunction with other scientific societies, made
suggestions concerning the setting apart of primitive areas in the Kosciusko State Park,
and discussions with the Trustees are still in progress.

The proclamation protecting certain wild flowers and native plants was renewed for a
further period of three years from ist July, 1944.

On the suggestion of the Chairman of the Sir Joseph Banks Memorial Fund Trust,
this Society has accepted the responsibility of obtaining the co-operation of allied
societies with a view to approaching the Government to press for the erection of a new
National Herbarium as one of the first post-war undertakings. At present the collections
are most inadequately housed and the fire-risk is great.

During the year your Council wrote to the Government advising that, while it was
very concerned at the despoliation of the native fauna, it did not feel that total
prohibition of the sale of flowers would have the desired effect of preventing it. It was
suggested that the position could best be met by issuing licenses for picking only to
people who could show they had legitimate access to cultivated plants or to adequate
supplies of naturally growing plants on private land.

Representatives of your Council also attended a conference called by the Director
of Physical Education to consider the possibility of preparing an ordered classification
of parks and reserves of Crown lands. :

It is a pleasure to be able to record that the Premier of New South Wales has added
a further £2,000 to the fund collected years ago to commemorate the life and work of
Sir Joseph Banks and has directed that the fund be used for the editing and publication
of the Banks papers.

We offer congratulations to Professor EH. Ashby on his appointment as Scientific
Attaché to the Australian Legation at Moscow, to Dr. N. C. W. Beadle on attaining the
degree of Doctor of Science of the University of Sydney, and to Dr. H. L. Kesteven on
his award of the Walter Burfitt Prize by the Royal Society of New South Wales.

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

Dr. H. L. Jensen, Macleay Bacteriologist to the Society, has continued his work on
the influence of reaction of the medium on symbiotic nitrogen fixation in pasture legumes,
especially lucerne and subterranean clover, and a paper on nitrogen fixation by these two
plants in acid and alkaline soil has been published in these Procrerpines. The experi-
mental work has aimed chiefly at determining the nitrogen-fixing efficiency of the root-
nodule tissues in sand medium of different reaction. The results as a whole indicate that
the relative efficiency of the nodules is generally higher at neutral to alkaline than at
acid reaction, although this seems to depend on the experimental conditions, and is
always higher in lucerne than in subterranean clover. Further work on the influence of
molybdenum is nearing completion and supports the earlier view that this element
seems to have a specific influence on the process of nitrogen fixation. Experiments on

vi PRESIDENTIAL ADDRESS.

were world-wide. In Europe the two earlier glacial stages, Giinz and Mindel, covering
together with the intervening interglacial stage about 175,000 years (Zeuner, 1935),
were separated from the two later, Riss and Wiirm (about 230,000 years), by a very
prolonged interglacial interval of the order of 200,000 years, so it is possible to
distinguish Early, Middle and Late Pleistocene epochs. The early and late epochs each
comprised two glacial stages and an interglacial stage. A similar subdivision of the
Pleistocene Period has been established for North America.

It is possible that there was yet another and pre-glacial Pleistocene epoch in
Australia, as there appears to have been elsewhere. Chapman (1920), describing a
collection of marine fossils from Ooldea on the Transcontinental Railway which he
regarded as Lower Pleistocene, pointed out that it contained forms, particularly the
foram Orbitolites complanata, which live to-day only in the warm waters of lower
latitudes. Moreover, crocodilian remains of probable Pleistocene age have been
discovered in latitudes very far south of those frequented by the crocodiles of to-day.
These occurrences, admittedly inconclusive, would suggest that the earliest epoch of
Pleistocene time was marked by a climate warmer than that of the present day.
However, much more evidence will be required before this epoch can be regarded as
definitely established.

A rather important stage of the Late Pleistocene epoch was that occupied by the
melting away of the Wtirm ice-sheets; according to Daly (1935) this may have taken
as much as 16,000 years. The reduction of the world’s glaciers to approximately their
present dimensions marked the end of Pleistocene and the beginning of Recent time,
and is thought to have been accomplished some 9,000 years ago.

There appears to be general acceptance of the view that glacial and interglacial
stages of the Pleistocene were synchronous in the Northern and Southern hemispheres.
The major waxing and waning of the ice-sheets produced eustatic changes in sea-level—
respectively emergence and submergence—all over the world, and these movements, of
some of which we have evidences around our coasts, can serve to some extent as post-
Tertiary time-markers. A relatively slight increase in the extent of the world’s glacier-
ice, which is believed to have taken place some 3,000 or 4,000 years ago, led to a small
emergence of which evidences are numerous around our coasts, and this event serves:
to divide the Recent Period into an earlier and a later epoch.

MARINE SEDIMENTATION.

In Pleistocene time Australia had a coastline differing not very greatly from that of

the present day. There was still in existence, however, a substantial remnant of the
' shallow Murravian Gulf which had in Tertiary time extended from the Murray mouth
inland nearly to Broken Hill, and occupied much of south-eastern South Australia and
north-western Victoria, and the sea still flowed in the Nullarbor Bight at least as far
north as Ooldea on the Transcontinental Railway. On the sites of these embayments,
now raised well above sea-level, traces of Pleistocene marine and littoral deposits have
been disclosed by bores and in outcrops.

In the Tintinara bore in South Australia (Howchin, 1929a) a total thickness of
220 feet of Pleistocene beds, abundantly fossiliferous, was encountered, all of the
determined species except three being of living types, and some 25 miles to the south-west
another bore at Alfred, near the Coorong, is stated to have passed through 348 feet of
marine Pleistocene deposits, all below present sea-level. From the other side of the old
gulf, in the cliffs near the mouth of the Glenelg River (Victoria), thin deposits of
limestone and shell-beds, which Singleton (1941) regards as Lower Pleistocene, appear
at aititudes up to 200 feet above sea-level.

A bore put down at Sorrento, near the entrance to Port Phillip Bay, passed
through a succession of 377 feet of estuarine clays, beach-sands, shallow-water lime-
stones and dune-sands. From the foraminiferal and ostracodan content these were
considered by Chapman (1928) to be Pleistocene. The top is about 100 feet below
sea-level.

From outcrops at Ooldea on the Nullarbor Plain, Chapman (1920) described a
collection of marine fossils which he regarded as Lower Pleistocene.

PRESIDENTIAL ADDRESS. vil

Apart from the occurrences indicated by these records, the Pleistocene marine
deposits—or most of them—are still beneath the sea.

GLACIATION.

Of the four major glacial stages recorded from the Northern Hemisphere, traces
of only two or at the most three have been found in the Commonwealth, in Tasmania
and New South Wales. In 1933 the late Dr. A. N. Lewis, whose untimely death a little
more than a year ago was a great loss to geological science, recognized in Tasmania three
stages or phases; (a) an earlier or ice-cap stage (Malannan) which probably covered
between one-third and one-half of the island, and whose ice-sheets came down almost to
sea-level in the west and south-west; (b) a cirque-cutting phase (Yolandian); and (c)
a mountain-tarn phase (Margaret), which may or may not have been independent of the
Yolandian. These last two had a very much less extent than the Malannan and were
confined to the higher altitudes.

It is a matter of considerable interest that at Gormanstown, near Mt. Lyell, on the
West Coast, have been found the only Pleistocene deposits of glacially varved clays or
shales known in the Commonwealth (Lewis, 1928). There are two occurrences on
different horizons closely associated with till and evidently related to interglacial
intervals of the Malannan stage.

In New South Wales only the highest country, that around Kosciusko, was glaciated.
Recent surveys (Browne, Dulhunty and Maze, 1944) suggest that ice-sheets during the
earlier stage extended about 20 miles northwards along the Main Divide from Mt.
Kosciusko, and that they may have covered a total area of at least 150 square miles.
Traces of only one later glaciation have been recognized with certainty in this region,
during which small glaciers produced cirques, rock-basins and moraine-dammed lakes
(David, 1908).

It seems natural to correlate the ice-cap glaciations of Tasmania and Kosciusko with
one another, but it is hard to tell to which of the European glaciations they should be
referred. The Mindel and Riss stages seem to have been of almost equal intensity, and
were the most severe of all, and with one or other of them it seems most likely that the
Australian ice-caps were synchronous. On physiographic grounds David estimated an
age between 100,000 and 200,000 years for the older Kosciusko glaciation, and if this
estimate has any value then it may have been contemporaneous with the Riss. David
also calculated that the later Kosciusko glaciation might have occurred not more than
15,000 or 20,000 years ago, and it may therefore be assignable to a phase of the Wiirm
stage. The extreme freshness of the Tasmanian moraines of the Yolande and Margaret
- glaciations has often been commented on, and they may perhaps correspond to two
phases of the Wiirm.

LAKES AND LAKE-DEPOSITS.

For most of Australia, apart perhaps from the periglacial zones, where doubtless
the desiccating effect of the glacial anticyclones was felt, the Pleistocene Period was a
time of high rainfall and the continent, even in its central parts, was a land of brim-
ming rivers, spreading swamps and full-bosomed lakes, and of a varied and abundant
fauna and flora. A few of the larger lakes are deserving of special mention. Some time,
probably in the early Pleistocene, an east-west compound earth-fold was upraised athwart
the south-flowing rivers of Central Australia, and in the compensating downward sag
to the north, their waters were impounded (Howchin, 1929b; Fenner, 1931), with the
result that a great inland lake or sea was formed embracing the present Lake Eyre and
possibly also Lakes Blanche, Gregory, Callabonna and Frome (David, 1932). This may
have had one or more overflows to the south, one being by way of the Pleistocene Lake
Torrens to Spencer Gulf. The vast sheet of water, forming what the late Sir Edgeworth
David used to call Lake Dieri, must have covered an area of about 40,000 square miles
and was therefore comparable with the Pleistocene Lakes Bonneville and Lahontan in
North America. Into this lake vast quantities of sediment were transported by the
mighty streams that entered it, and it endured throughout Pleistocene and into earlier
Recent time. The total thickness of sediment laid down in this lake is unknown, but

oe

viii PRESIDENTIAL ADDRESS.
TABLE
A Very Tentative Post-Tertiary
(N.B.—Not all entries are of
chest ei __
|
Lake-, Swamp-
Periods. | Epochs. European Marine Glaciations. _ earl River-
Subdivisions. Deposits. Spring- Deposits. Deposits,
a
|
Present-day deposits, |
| clastic, organic and
chemical, including | Present-day
salt and gypsum in flood-plains,
LATER lakes of 8.A., W.A., |
| RECENT. ete;
| (e. 4,000 years.) also mound-springs. |
MID-RECENT |
a INTERVAL.
Zz
= |_ —
a |
a=] ere
| Rae Deposits in coastal
ens @o0mraized swamps, e.g., Mowbray | :
| Benches Swamp (Tasm.) and Very
; Bouse Swany Wet) | ime
| EARLIER ana. c ypsum eposits of | alluvium
RECENT. | poder ie’ a s.w. N.S.W. and deposited
| (¢c. 5,000 years.) eetiaries Mallee district, Vict. | except by
3: Ipelse silts cael sole | coastal
F eposits wi Ossi Tivers.
| Greed Barner vertebrates, e.g.,
| in part Lake Callabonna, S.A. |
Wiirm
Recession.
|
Yolande and
Wurm Margaret phases H
Glacial. (asm):
2nd Kosciusko
Great Barrier stage (N.S.W.).
“ LATE ea
LEISTOCENE. - - in part,
(c. 280,000 years.) | fi#e Warm and |
g : clastic sediments
beneath it.
Malanna (Tasm.)
and Ist Kosciusko
Riss glaciations |
Glacial. (ice-cap), with |
till, moraines,
outwash gravels, |
etc.
e a
Zz Some extinct
taal mound-springs of
> Gt. Artesian Basin | Deep |
° MIDDLE Mindel-Riss in S.A alluvium in
a PLEISTOCENE. Interglacial. mature and
iS (c. 185,000 years.) Sediments of Lake old river-
a Dieri, the Fortescue | valleys, —
A depression (W.A.), | in places
Au “ ete. | basalt-
covered ; also
alluvial
; terraces
Mindel ste
f above existing
Glacial. Tiver-level.
EARLY | Fossil re
PLEISTOCENE. ————— of vertebrates
(ce. 175,000 years.) Giinz-Mindel oecasio
Interglacial. found.
Giinz
Glacial. |
_
oe a
oe Warburton &.,
PRE-GLACIAL. Tovsiliterous etc.
(? years.) Ooldes Falenele R | with crocodilian
Sorrento Bore, — rena
ete. Some river
terraces.

)
r

ya.

| Chronology for the Commonwealth.
the same degree of probability.)

Coastal
Aeolian
Deposits.

Some older

dunes and

dune-ridges
mostly

vegetated.

Some older

those with
soil-horizons
in s.w. Vict.

| Coastal
| Limestones
| of W.A.

|
|

|

PRESIDENTIAL ADDRESS.

(Sea-level may
have been
ec. 130’ above
present position
and 100-150’
benches and
terraces may
belong here
in part.)

S.A. with
formation of
Lake Dieri.

Milder and more
humid than at
present ;
coastal waters
warmer.

- Inland | Notable Events
Aeolian Volcanic Eustatic Tectonic Climate. in Organic
Deposits. Rocks. Movements. Moventents. World.

Recurrent
movement on
Pleistocene and More or less
Earlier Recent as at present. | Inland spread
faults. 2 Minor oscillations | of surviving
Gentle folding of temperature | plants and
of raised beach- | and rainfall. animals from
| | deposits of n.w. less arid
| W.A. coastal zones.
Emergence of |
15-20’ Onset of cooler |
producing | and wetter
raised beaches, | conditions.
ete. |

Sandridge
deserts, Hot and dry.
chiefly in Basic lavas’ | | Faulting and

W.A., S.A; and tuffs uplift along Arid conditions
and N.T. from | Gt. Aust. except in Extinction of

central vents Bight, s.e. narrow coastal giant fauna,

Sandridges in n.Qd. and | coast of §.A. belts. and of flora

of Burnett R. and | and s. coast of in part.
Mallee district, in w. Vict. and Kangaroo I. General
Vict. ste io As desiccation.
ka | TEES Doses PP en wslilades haere Zeal
| | Final sub- Getting warmer ;
| mergence of glaciers melt in
| about 270’. Tasm. and on Kosciusko.
Arrival of
Emergence. Australoid man
Down-faulting with dingo
of east Generally cold in Australia.
coast of and wet with
Submergence. Australia and frigid conditions
45’ terrace Tasmania. of decreasing intensity
Extensive formed. in Tasm. and on
basic flows, Kosciusko during
chiefly in the glacial stages.
n.Qd. Emergence to Arrival of
| and western | possibly 250’ Tasmanoid man
Vict., below present in the
also in sea-level. Formation of Commonwealth.
S.A. and Tasm. great anticline
(May have con- athwart
= tinued from Lower Murray R.
| late Pliocene with damming of
time.) Lake Nawait.
Deep Probably
submergence, warmer and
producing wetter than
100’ terrace at present. Spread of
in Tasm., W.A. Rain-forest.
and N.S.W.
Faulting in
S. Australian
and
| Emergence to Port Phillip
| possibly | horsts and Rise of
| 250’ below | sunklands. fauna of
present Cold and giant
sea-level. humid generally, marsupials
but snowline and
| Submergence. still above flightless
land-surface. birds.
This continued
Emergence. Upwarping of through
Gawler Range- Pleistocene
Olary ridge in time.

x PRESIDENTIAL ADDRESS.

in Lake Frome Pleistocene and Recent deposits attain a thickness of 400 feet. Lake
Torrens, then much larger than now, was also heavily alluviated.

An extensive area in South Australia around the Murray River above Overland
Corner was the site of another but much smaller Pleistocene lake, possibly 8,000 square
miles in extent, which may be called Lake Nawait, formed by the upwarping of a part
of the Tertiary strata constituting the floor of the Murravian Gulf (Howchin, 1929a).
Sections in the river-cliffs show current-bedded ferruginous clayey sands, grits and
gravel, with limestone containing freshwater gastropods. The lake extended into the
south-west of New South Wales and the Mallee district in north-west Victoria, and in it
sediments accumulated to as much as 300 feet. Relics of this lake are Lake Victoria
in New South Wales, Lake Bonney in South Australia and a host of dried-up gypsum-
pans in Victoria. The barrier that formed the lake rose very gradually, and was
eventually cut through by the Lower Murray, so that the lake was wholly or partially
drained, probably in very late Pleistocene or early Recent time.

A third large lake may be briefly noted here. The shallow Bass Strait had been
formed by sagging or faulting during Tertiary time. It can be seen from Fig. 1 that
a lowering of the present sea-level by 40 fathoms would result in the formation of a
lake with only a narrow exit to the sea. During the most severe of the Pleistocene
glacial phases sea-level was lowered more than 240 feet and, as Noetling (1910) has
pointed out, a lake must have been left either at sea-level or above it, into which flowed
rivers from Tasmania and Victoria. This lake was, of course, transient in character
and Bass Strait was re-established when an interglacial phase supervened. It would
appear as if Tasmania was joined to the mainland at least once and possibly twice
during the Ice Age.

——s 2502

Sams.

144

x
iis

al
Ase

Fig. 1.—Submarine contours, in fathoms, of Bass Strait.

In Western Australia the Upper Fortescue River flows in a huge saline depression
120 miles long and with an average breadth of 25 miles. This is the remnant of a lake
of much greater extent, in which the sediments are known to be more than 100 feet
thick. This and other smaller, deeply alluviated depressions appear to be of Pleistocene
age (Jutson, 1934).

PRESIDENTIAL ADDRESS. xi

SWAMPS AND SWAMP-DEPOSITS.

Apart from the swamps that are in process of formation both inland and along the
eoasts at the present day, there are traces of others which existed in Pleistocene or
earlier Recent time. The site of one old tidal swamp is to be seen at Rocky River, in
the south-west of Kangaroo Island, 3 or 4 miles in from the present shoreline and at a
height of 320 feet above it. Its deposits, which are associated with old dune-limestones,
contain in addition to foraminifera and land-shells remains of Diprotodon and other
extinct marsupials (Tindale, Fenner and Hall, 1935).

Among a number of extensive peaty deposits in the valley-plain near the mouth of
the Duck River, on the north-west coast of Tasmania, is the Mowbray Swamp with a
surface from 25 to 50 feet above sea-level, composed of sandy and muddy peat more than
10 feet thick, overlying marine or estuarine sands (Nye, 1934). The peat has yielded
remains of extinct marsupials and the whole series of deposits is probably of earlier
Recent age. Other similar deposits are the Boneo Swamp, west of Cape Schank
(Victoria) and the Pejark Marsh in western Victoria. :

_ SPRING-DEPOSITS.

Around the margin of the Great Artesian Basin there are numerous mound-springs,
natural outlets for the artesian water. In the course of time these springs have built
up mounds composed of clay, sand and travertine from 10 to 80 feet in height. In South
Australia there are in addition older and larger mounds built up by springs at a time
when the head of water was much greater than at present. Ward (1921) recognizes
two series of these, an older up to 130 feet high, several square miles in area and much
dissected, and a younger, also dissected and intermediate in size between the older
mounds and those of the present day. These extinct springs may have functioned during
periods of high rainfall in Pleistocene time.

RIVERS AND RIVER-DEPOSITS.

The whole of the Pleistocene continent must have been traversed by great rivers,
even those parts which are now arid. There are some evidences, for example, that the
curiously elongated salt-lakes or playas which characterize much of the inland parts of
Western Australia, were once parts of river-systems (Gregory, 1914; Browne, 1934),
and the alluvial deep leads, such as those of Kanowna and Bulong, are probably
Pleistocene, in part at all events, and testify to a time when great rivers brought down
huge volumes of gravel, sand and siit. The coastal streams of the north-west, now
feeble and fitful, were also of some magnitude in Pleistocene time.

The streams emptying into the Lake Eyre lake-system, and those forming the
Murray-Darling system, are characterized in their mature and senile courses by alluvium
which may be some hundreds of feet in thickness. Some of this may be of Tertiary age,
and some is Recent, but undoubtedly much is Pleistocene. The deeper sediments in
the mature valleys of many of the Victorian rivers, which were famous 90 years ago for
their gold content, are in part Pleistocene, and our New South Wales rivers traversing
the Western Slopes and the Western Plains have spread out great sheets of post-Tertiary
alluvium along their courses to form something in the nature of alluvial piedmonts.
The black soil of some of the rivers, like those of the Liverpool Plains and the country
about Warialda, overlies sandy or gravelly alluvium, and a somewhat similar sequence
is found in the Darling Downs in Queensland. The reason of this change in the character
of the river-deposits is yet to seek. The late Tertiary and post-Tertiary alluvium often
appears in the form of terraces along the rivers, indicating probably renewed uplift in
post-Tertiary time, and occasionally Pleistocene gravels are found underlying basalts, as
at Hinasleigh, Herberton and elsewhere in northern Queensland. Some of the basalt-
capped deep leads of western Victoria may belong here (Hills, 1938).

Another series of alluvial fans forming a regular piedmont belt constitutes part of
the Adelaide Plains (Fenner, 1931), brought down by the rivers from the upfaulted Mt.
Lofty Range. An alluvial piedmont also fronts the scarp of the Darling plateau in
Western Australia, and indeed, similar deposits were laid down, chiefly in Pleistocene
time, wherever the surface had been uplifted differentially through faulting or warping

xii PRESIDENTIAL ADDRESS.

during the late Pliocene Kosciusko uplift. The east coastal rivers of Queensland are
entrenched to depths of as much as 75 feet in Pleistocene alluvial plains and terraces,
and the old silts extend far up towards the Main Divide, while downstream they merge
into raised deltas and marine terraces. Alluvial terraces are also prominent and
extensive on some of the eastern rivers of New South Wales, particularly those of the
North Coast. It is perhaps significant that in some rivers here, as in southern Queens-
land, there are gravel terraces at levels of a little over 100 feet above present river-level
and silt-terraces at lower levels. The higher ones may be late Tertiary and the lower
Pleistocene. . Near Sydney we are familiar with the Pleistocene flood-terraces on the
Nepean at Wallacia and on the Hawkesbury between Penrith and Windsor.

Alluvial terraces above present flood-plain level characterize some of the Victorian
south-flowing rivers, as the Yarra and the Werribee, and the same is true of many
Tasmanian rivers.

The antiquity of some of the river-deposits mentioned above is attested by the
discovery in them, at various places, of remains of post-Tertiary fossil vertebrates.

PLEISTOCENE STRANDLINE MOVEMENTS.

The conception that during the Ice Age the waxing and waning of the ice-sheets
in the glacial and interglacial phases were responsible for alternate falling and rising
of sea-level all over the world, has been made familiar to us by the writings of Sollas
(1924), Daly (1935), and others. At the beginning of Pleistocene time there were
probably few, if any, great ice-sheets on the earth, and the shoreline stood higher than
at present by an average height of about 130 feet, according to Daly. During each of
the glacial stages, sea-level was lowered by an amount depending on the severity of
refrigeration, and a new and lower base-level of erosion was imposed on the rivers and
on the waves of the sea. The raising of sea-level in the interglacial intervals resulted
in the formation of terraces of erosion and deposition along the coastline and also along
the lower courses of the coastal rivers. Since the interglacial climate was variable
but in general milder than at present, these terraces appear at various heights above
present sea-level. Similarly, evidences of the glacial stages remain in the shape of
drowned valleys at various depths below present sea-level.

It would appear (Zeuner, 1935) from the European record that each of the four
glacial phases had its maxima of cold with intervening milder intervals—two maxima
each for the first three and three for the Wiirm—and theoretically at all events each of
these intervals should have recorded its existence along the coastline. Around the
Australian coast there are benches and erosional terraces and raised beaches which from
their relative constancy of altitude seem to be eustatic and to indicate intervals of
emergence. Unfortunately, however, for few if any of these have accurate measurements
of height above a constant datum been made.

At various places a 100-foot terrace either of accumulation or of erosion has been
noted. In Tasmania Lewis (1934) recognized it in the north, east and south-east, and
along the Derwent and other estuaries, and Loftus Hills (1914) recorded a raised beach
from Point Hibbs on the west coast. R.M. Johnston’s (1888) Helicidae sandstone from
some of the Bassian islands appears to be in part a raised beach-deposit. In Western
Australia a 100-foot raised beach was found by Dr. Arthur Wade at Shark Bay, and Prof.
E. de Courcy Clarke has informed me of-an erosion-bench at about the same altitude on
the south coast near Hast Mt. Barren.

In a paper to be presented to-night Mr. W. H. Maze, by an analysis of the contours
of the Military topographic maps, shows evidence of a bench or terrace at between 100
and 150 feet above sea-level along a considerable stretch of the New South Wales coast.

Terraces of accumulation at between 40 and 50 feet above sea-level have been
recorded from Tasmania, and from Western Australia near Cape Naturaliste and between
Port Hedland and Broome. So far as I am aware this terrace has not been noted else-
where, unless possibly that described from the Maribyrnong River near Melbourne
represents it (Mahony, 1943b).

Although observations on these terraces are still very few, it is perhaps permissible
to hope that they may be found. eventually all around our coasts. <A correlation is

PRESIDENTIAL ADDRESS. xiii

suggested with the 30-metre and 10-metre terraces of the Mediterranean coast, Great
Britain and North America, which are generally assigned to the Mindel-Riss and Riss-
Wiirm interglacial stages respectively. It seems probable that during the Mindel-Riss
interglacial epoch all ice melted off the earth, hence its shoreline and that of the pre-
glacial epoch may both be represented in the 100-150 foot terrace.

Lewis (1934) recorded exceedingly interesting profile-sections obtained from borings
made across the Derwent estuary in Tasmania. The results were such as to indicate
clearly a submarine valley-in-valley structure. A similar structure is disclosed in some
sections across Port Jackson and George’s River near Sydney previously published by
Smith and Iredale (1924). It seems highly probable that, as Lewis suggested, the
submerged valley-floors are related to times of low sea-level marking intervals of glacia-
tion, though it would be very hard to tell precisely which of the glacial culminations are
indicated, because, inter alia, there is no certainty whether the shallower or the deeper
level is the older. Both Lewis and Edwards (1941) have attempted correlations, the
latter ascribing the deeper valley to the Riss glaciation. However, it is not impossible
that the phenomenon of the valley-in-valley structure is a combined or cumulative effect.
During the phases of each glacial stage the ocean-surface may have come down again
and again to approximately the same level, and this may account for the much greater
maturity of the shallower valley as compared with the deeper one, which was probably
eroded during one severe glacial phase, such as the earlier one of the Riss, or perhaps
during the Mindel and Riss glaciations. It will be of interest to see if similar submerged
valley-in-valley profiles are revealed by borings in other Australian estuaries.

The melting of the Wiirm ice-sheets was the last event of Pleistocene time, and it
resulted in the rise of the strandline to approximately its present level, or more precisely
to about 15 or 20 feet above its present level. Strictly speaking, if the drowning was
eustatic we should find that its vertical extent was everywhere the same around our
coasts, but in actual fact where measurements have been made they show variations.
The greatest depths to bedrock have been recorded at Stockton, near Newcastle, and the
Hawkesbury estuary, 7 miles from its mouth, both about 250 feet; elsewhere the figures
have generally been in the neighbourhood of 100 or 150 feet. The discrepancies are
perhaps to be explained by the varying distances of the positions of measurement from
the original shoreline, since the apparent depth of drowning decreases upriver from the
coast. If allowance is made for the subsequent small emergence of 20 feet in mid-Recent
time, it would appear that drowning amounted to not less than 270 feet; this is in close
agreement with Daly’s estimate of between 250 and 300 feet for a world-wide positive
eustatic movement of the strandline.

The drowning was, aS we have seen, spread over a considerable interval of time,
and it may have been punctuated by stillstands and even retrogressions. Some interesting
information bearing on this matter was obtained in a number of excavations near
Sydney. From Shea’s Creek (Etheridge, David and Grimshaw, 1896), near Botany Bay,
a submerged forest passing in places into a peat-bed was discovered at a depth of 15 feet
below H.W.M. covered by estuarine beds containing marine shells. In a trial shaft put
down in Port Jackson the normal sandy estuarine shell-bearing beds of the harbour were
found to be underlain by very carbonaceous clays with abundant plant-remains at a
depth of about 90 feet below H.W.M. At Narrabeen Lagoon (David and Halligan, 1908)
shallow boring disclosed at about 59 feet below H.W.M. the presence of carbonaceous
clays with roots of trees and gastropod shells of shallow water habit; these were over-
lain by marine sands. Sussmilch (1936) records that a bore put down at Botany Bay
to a depth of 10634 feet (collar-set 26 feet above sea-level) passed through no less than
five layers of peat varying from 6 inches to 12 feet in width below sea-level and inter-
bedded with sand. In a well put down hard by tree-stumps in the position of growth
were found at about 29 feet below sea-level.

The presence of peat-beds in all these sequences, and in others which have been
reported, indicates intervals of stillstand during the progress of submergence.

THE MID-RECENT EMERGENCE.
This small eustatic movement, directly and indirectly one of the important influences
in modifying our coastline, has manifested itself in a great number of ways. Raised

XiV PRESIDENTIAL ADDRESS.

beaches and shellbanks, extensive low-lying coastal plains, subdued and vegetated cliffs
fronting the open sea, tied islands, filled-in embayments, barrier-beaches enclosing
lagoons, wave-cut rock-benches, stacks and sea-woin caves now beyond the reach of wave
and tide, dead coral in situ above high-water mark, coastal corridors in which fossil
mangroves and coral-débris are found—taken collectively these phenomena, abundantly
displayed up and down our coastline and around the coastal islands, attest the actuality
of the movement.

Unfortunately it would appear that but few of the features of emergence have been
accurately referred to a fixed and common datum-level, and since the present position of
deposits or erosion-benches depends on the depth below sea-level at which they were
formed a mere statement of height above sea-level gives no precise indication of the
extent of movement. When all allowances are made, however, there does seem to be
some evidence of a uniform emergence, completed perhaps in more than one stage, and
amounting to about 15 or 20 feet. According to Cotton’s (1928) computation the move-
ment was accomplished between 3,000 and 5,000 years ago. Daly (1935) is of opinion
that evidence of a similar movement of small amount is to be found all over the world,
and he has attributed it to ‘“‘a small drop of general sea-level about midway in post-glacial
time’”’, due to a refrigeration of the earth’s climate, with the locking-up of more water in
the high-altitude and high-latitude glaciers.

Not everywhere around our coasts are the evidences of eustatic emergence incontro-
vertible; some of the phenomena such as bay-head flats, silted estuaries and tied islands
might well be referred to normal processes of coastal deposition, but these would clearly
be facilitated and accelerated by a downward movement of the strandline. - Also some
accumulations of shingle or coral-débris are probably attributable to exceptional storm-
waves o1 hurricane-waves. Again, there is no agreement among physiographers as to
whether the rock-platforms are due to emergence or are being formed by storm-waves
at the present day. Finally, differential movement of the land has occurred in Recent
time in some places, as in the unstable areas of Port Phillip Bay and the South Australian
sunklands.

It is noteworthy that, while the shelly marine fauna of the raised beaches is composed
of existing species, some of the forms are no longer found in the latitude of-the deposits,
but frequent the warmer parts of the coast. For instance, the raised beach at Port
Wakefield on St. Vincent Gulf (Howchin, 1929b) contains an amazing profusion of Arca
trapezia, which is no longer present in the waters of the Gulf, and where the same beach
has been identified in bores near Adelaide at between 9 and 25 feet below H.W.M., it
contains one of the shallow-water foraminifera, Orbitolites complanata, which at present
comes no farther south than Shark Bay on the west and the Great Barrier Reef on the
east: Many of the species found in the upraised Recent shelly sands of the Swan River
estuary are not nowadays known to exist south of Geraldton.

It would appear, therefore, that in earlier Recent time, before the emergence, the
coastal waters were notably warmer than at present.

POST-TERTIARY TECTONIC MOVEMENTS.

As distinct from the eustatic strandline movements just discussed, there appear to
have been in some parts of Australia differential movements of the land in the shape of
faulting, unwarping and monoclinal folding, though for some of them the precise dating
is a matter of difficulty. Andrews (1910) has shown that the movements, chiefly of
uplift, which eastern Australia experienced in Tertiary time, culminated in the Kosciusko
uplift which was accomplished mainly about the end of the Pliocene and certainly before
the coming of the Pleistocene Ice Age. Accumulated evidence makes it clear that the
Kosciusko movements affected in varying degree the whole of the continent, and there
can be little doubt that they continued at intervals through post-Tertiary time and
that in some places minor adjustments are taking place at the present day.

In South Australia the horsts of the Mt. Lofty and Flinders Ranges and the sunk-
lands forming St. Vincent and Spencer Gulfs were brought into existence probably at the
Kosciusko epoch, but Howchin (19290) has given good reasons for the belief that both
downward and upward movement was renewed in Pleistocene and even in Recent time,

PRESIDENTIAL ADDRESS. XV

and the earthquake-shocks such as that experienced in 1911 along the plain fronting the
Mt. Lofty Range suggest that equilibrium has not yet been attained. Likewise the
Mornington earthquake of 1932, which was clearly related to the fault forming the
eastern boundary of Port Phillip sunkland, showed that the sinking in “that region,
begun during Tertiary time and renewed in the Pleistocene and Recent periods, is not
yet complete.

The upwarping of the east-west Gawler Range—Olary ridge in South Australia and
the sagging to the north of it which formed Lake Dieri may have begun in late Pliocene
time, and indeed Fenner considers that they antedated the formation of the Mt. Lofty—
Flinders Range horsts, but the other great low bulge in the south-east of South Australia,
which has been shown to stretch from about Morgan on the Lower Murray in a south-
south-east direction into the Mallee and Wimmera districts of Victoria (Hills, 1939) is of
Pleistocene and Recent date. For in the first place the Pleistocene strata laid down
in the old Murravian Gulf have been involved in the movement, and secondly, on the west
the elevated ground rising to 300 feet above sea-level descends to the present coastline
by a series of upraised shorelines and coastal dunes, some of which, at least, are of
Recent age. It would appear that the upbowing of the surface, whereby the coastline in
the south-east of South Australia has been pushed westward for 50 miles, was saltatory
or periodic in character and may have been accomplished in part by faulting. Somewhat
similar features have been described from the south-west coast of Kangaroo Island by
Tindale, Fenner and Hall (1935), who believe that there was Pleistocene or Recent coastal
uplift amounting to about 300 feet.

If Chapman’s determination of Lower Pleistocene marine fossils at Ooldea, 380 feet
above sea-level and upwards of 90 miles inland, is correct, then it would seem that the
draining of the Nullarbor Bight, which in Tertiary time extended to as much as 200
miles north of the present coastline, was accomplished by vertical uplift in Pleistocene
or Recent time. There is some evidence that this was accompanied or followed by down-
faulting along the coast.

It is exceedingly probable that along the faults bounding the east coast of Australia
movement occurred in post-Tertiary time, perhaps in part as compensation for the
Kosciusko uplift. The faulting along the Queensland coast, which seems to have let
down the submarine platform whereon the Great Barrier Reef was built, must have
persisted into comparatively recent time, since the coastal rivers rejuvenated through
the foundering of a strip of coastline have cut their gorges back for only a short distance,
amounting in the Barron River to no more than a dozen miles. Movement was evidently
more or less continuous or perhaps periodic, and the total extent of the foundering must
be at least 800 feet.

Smith and Iredale (1924) have shown that there has been a downfaulting or down-
warping off the New South Wales coast where, at from 10 to 15 miles offshore, the
presence of consolidated beach-rock with littoral molluscs has been determined along a
stretch of 200 miles at between 70 and 75 fathoms. The molluscan content of the rock
is essentially of Recent type but the forms show closer resemblances to living species
from Victoria and Tasmania than to those of the New South Wales coast, suggesting
that at the time when they lived Tasmania was joined to the continent. It is probable
that the downward movement was of Pleistocene age. In this connexion it is of interest
to note that along the east coast of Tasmania post-Tertiary faulting is suggested by the
asymmetry of the river-systems, the watershed between east- and west-flowing streams
being in places only a few miles inland.

Gentle warping of the raised beaches on the north-west coast of Western Australia
denotes a slight deformative movement of Recent age in that region (Raggatt, 1936).

THE GREAT BARRIER REEF.

(Steers, 1929, 1930, 1932, 1937; Spender, 1930; Richards and Hill, 1942.)

This wonderful structure, unique of its kind, has of recent years been the subject
of somewhat intensive study and description by both geologists and biologists, but its
secrets have been by no means all laid bare. It may be inferred that the Reef rests
ultimately on a rock-platform that has been faulted down or warped down from the

Xvi PRESIDENTIAL ADDRESS.

Queensland coast, but borings at Michaelmas Cay and Heron Island penetrated to depths
of 600 and 732 feet respectively without encountering bedrock, the coral-material, of a
thickness of about 450 feet, resting on some hundreds of feet of sand with shell-fragments,
foraminifera and molluscan shells. Sinking of the rock-platform was evidently accom-
panied or followed by heavy sedimentation. But since reef-building corals cannot live at
a greater depth than 30 or at most 50 fathoms, coral-growth must have begun at a time
when the surface of the sandy sediments was not more than 300 feet below the sea-level
of the time. It may be, therefore, that sinking was temporarily halted, or slowed down
sufficiently for sedimentation to gain on it and form a floor at a relatively shallow depth,
and that later movements gradually carried the submerged shelf down to its present
position of perhaps as much as 800 feet under the sea. Alternatively the water may have
been shallowed during a Pleistocene glacial phase and the coral-growth started then as
noted below; in tropic seas the temperature, even at such a time, might well have been
favourable to the growth and spread of reef-corals. :

Iredale has reported that all the molluscan shells contained in the sands underlying
the coral-material are of existing types, and from this, Richards and Hill have inferred
that the Reef is entirely a Recent growth. It is perhaps permissible to suggest that the
list of molluscan forms is not long enough to rule out the possibility that the assemblage
is Pleistocene. There are some grounds for believing that the Reef had reached the
surface of the sea more than 4,000 years ago, inasmuch as a number of the coral-islands
display raised beaches and erosion-benches slightly above sea-level and certain
fringing reefs have suffered small relative elevation (Steers, 1930). These features
presumably appear as the result of the mid-Recent emergence. The growth of the Reef,
therefore, if Recent, would have had to be compressed into an interval of about 5,000
years, which would necessitate a rate of growth of the order of 1 foot in 10 years. It
seems more likely that coral-growth began towards the end of Pleistocene time, perhaps
during the Wtirm interval of low sea-level, and was continued while the ice-sheets were
melting and the last drowning was taking place, upward growth of the Reef occurring
more or less pari passu with the submergence.

The marked differences which have been noted between the northern and southern
parts of the Reef may indicate that its evolution was by no means simple nor its history
entirely uniform throughout.

THE COASTAL DUNES.

One of the most interesting and significant features of the coastal areas is provided
by an amazingly extensive series of sand-dunes. It has been long recognized that these
may be broadly divided into a younger series of living and moving and growing dunes
and an older series which are essentially fixed. The former, of course, belong to later
Recent time; the latter include dunes of both Pleistocene and earlier Recent formation.
The older dunes are often vegetated, and in many the sand has been converted into
calcareous or ferruginous sand-rock by the solution and redeposition of shell-fragments
or iron compounds, as at Narrabeen and at Red Point, Port Kembla. Again the older
dunes may be higher than the younger and farther inland, and in certain places they
have been found separated from the present shoreline by expanses of low, flat land,
as much as 10 miles wide, reclaimed from the sea by the mid-Recent emergence. Some
of the older dunes have yielded fossil vertebrate remains, and in others in which caves
have been formed the cave-deposits include similar remains.

Of all these dunes, except perhaps the last group, the most we can say is that they
are Pleistocene or earlier Recent. Others, however, are definitely to be classed as
Pleistocene, namely, those which appear where there are now no sandy beaches hard by
to supply material. The explanation of the occurrence of such accumulations is that
they belong to a glacial stage during which sea-level was lowered, exposing sandy beaches
which were later submerged. Probably our nearest example to Sydney is at Five
Islands, off Port Kembla (Davis, Day and Waterhouse, 1938). On Montague Island,
farther south, are patches of sand which must have had a similar origin. From the
Head of the Bight in South Australia, sand accumulations have been described by Tate
(1879) which, on the same kind of evidence, are to be regarded as Pleistocene, and Hills

PRESIDENTIAL ADDRESS. XVii

(1938) has found at Barwon Heads and other places on the Victorian coast, dunes in
which layers of soil alternate with the blown sand, indicating possibly accumulation of
sand during glacial stages and formation of soil during interglacial intervals.

Mention can be made here of only a very few of the more noteworthy dune-
formations. In south-eastern Queensland Moreton, Bribie, Fraser and Stradbroke Islands
are covered with dunes, in part probably Pleistocene, which attain a height of more than
900 feet, and are heavily vegetated (Whitehouse, 1940, p. 69). In Victoria an important
series of old dunes forms part of the Sorrento Peninsula; they rise to an extreme
height of 225 feet above the sea, and the Sorrento bore penetrated them for possibly
480 feet below sea-level, sporadically interbedded with Pleistocene fluviatile and estuarine
sediments.

In the south-east of South Australia and the adjacent parts of Victoria a remarkable
series of incoherent and consolidated calcareous dune-sand ridges runs in a north-north-
west direction parallel to the coastline and extends inland with gradually increasing:
altitude for nearly 60 miles. These are associated with the raised beaches which mark
the pauses in the uplift that formed the low ridge o1 bulge in this region referred to
earlier. The more westerly of these ridges at least are Recent (Fenner, 1931; Crocker,
1941; Ward, 1941). A similar series of dunes has been described from the south-
west coast of Kangaroo Island (Tindale, Fenner and Hall, 1935).

The formation known as the Coastal Limestone, a cemented calcareous aeolian sand-
rock, extends as a series of parallel ridges along the coast of Western Australia for-more
than 1,000 miles altogether (Maitland, 1919). West from Albany it forms cliffs up to
nearly 700 feet above the sea, and from Perth to North-west Cape there is a more or less
continuous belt of the limestone rising to a height of 200 feet. Extensive caves occur in
the limestones, and in one of these near Cape Naturaliste enormous quantities of remains
of fossil vertebrates were obtained.

EARLIER RECENT ARIDITY AND ITS CONSEQUENCES.

There is nowhere in Australia any continuity of associated strata which would
reveal changes of climatic conditions as from Pleistocene to Recent time; nevertheless
it accords best with the known facts to postulate that the cold, moist climate of the
- Pleistocene gave place after the melting of the glaciers to a time of extreme and almost
continent-wide aridity, from which Australia has not yet recovered. The most spectacular
result of the decline of the rainfall is to be seen in the vast expanses of sandridge desert
which cover so much of the interior, and which have been made familiar to us by the
unrelievedly gloomy descriptions of the early explores and in more recent times through
the aerial 1reconnaissances of Dr. Madigan (1938) and Mr. Donald Mackay (Bennett,
109)335))) 5

As indicated by the map (Fig. 2), the sandridge deserts cover an area of approxi-
mately 523,000 square miles between latitudes 18° and 31° S. Some parts of this region
appear never to have been penetrated by aborigines, and white men have seen them only
from the air. The sand on the ridge-crests is loose and liable to be blown about by every
breeze, but this merely serves to emphasize the essential fixity of the ridges as a whole,
and no more striking proof of the prolonged and complete aridity of earlier Recent time
can be imagined than the ordered array of parallel ridges of wind-piled sand, rising to a
height of about 60 feet and stretching in some instances perfectly straight and continuous
for as much as 100 miles.

The vegetation of the ridges indicates that since the time of their formation, when
the prevailing winds blew in the same direction as now but with very much greater
velocity, the climate has experienced some degree of amélioration, whereby the more
hardy plants have been able to creep back to their former habitat. The present rainfall
of the region ranges from 5 to 20 inches a year, and in the drier areas the ridges
support scattered clumps of Spinifer and Tvriodia irritans (poreupine-grass), while
acacias, eucalypts and other types are found where the rainfall is higher. On the margin
of the desert area, in south-west Queensland, Whitehouse (1940, p. 67) made the
interesting discovery a few years ago of dune-ridges east of Cooper’s Creek not merely
fixed and vegetated but also low and gradually being destroyed by weathering, evidently

B

xviii PRESIDENTIAL ADDRESS.

as a result of an increased rainfall in a region which was once arid. Though there is no
direct evidence on the point, it is perhaps not unreasonable to suppose that the increase
occurred about the beginning of later Recent time, when, as the mid-Recent emergence
suggests, the climate became somewhat colder, and presumably more humid.

|
|

—. Alice
eT \ wo

\
zy Ss

= of -“pricorn

:

100} parzoo! = 300) A400 cy

Scale of Miles * = its hee

oO

Fig. 2.—Map of the sandridge areas of Australia, embodying data compiled by Madigan
and the results obtained by the Mackay aerial expeditions. About one ridge in every four is
plotted. ;

The map also shows diagrammatically the sandridges or north-western Victoria.

Probably to the earlier Recent epoch of aridity also belong the east-west dune-ridges
of the Mallee country in north-western Victoria (Hills, 1939) which are likewise
vegetated and flattened. Low, vegetated sandridges, no more than gentle undulations,
are also to be seen athwart the Murray valley east of the Goulburn River and are not
improbably contemporaneous with those just described.

It is to this arid interval that we must assign the drying-up of the great lakes and
rivers of the interior. For instance, Lake Dieri was gradually reduced in size and
eventually split into a series of more or less independent smaller lakes, from which
the water ultimately vanished completely. The rivers that fed them, of course, also
dried up, or at best became very intermittent.

It is probable, too, that the former Lake Nawait on the Lower Murray disappeared
partly through desiccation, and the numerous deposits of gypsum in the far west and

PRESIDENTIAL ADDRESS. PATEK:

south-west of New South Wales (Jones, 1925) mark the sites of Pleistocene inland lakes
which were extinguished in the great drought of earlier Recent time. Then, too, the
system of rivers that had drained the great plateau of Western Australia shrank and
shrivelled and ceased to flow. Even where the effects of the aridity were least, in eastern
Australia, the rivers diminished markedly in volume, and even at the present day we
find many ‘misfits’-—large valleys carved out by the mighty torrents of the Pleistocene
now inadequately traversed by relatively tiny and ineffective streams. Indeed, it is
probably not too much to say that of the stream-erosion accomplished since the Kosciusko
uplift only a small fraction is to be attributed to Recent time.

Another event that may fairly be ascribed to this avid interval is the extinction of
the continent-wide giant fauna to be presently mentioned.

VULCANISM.

Hastern and southern Australia had been subject to repeated volcanic activity in
Tertiary time, but since the close cf the Tertiary this has been confined almost entirely
to the extreme north and south of the volcanic belt. The age of the post-Tertiary flows
is generally determined by the remarkably fresh and unweathered condition of the rocks
and the perfection of the structures shown by them, and in some instances by their
relations to late-Tertiary and post-Tertiary sediments. For the most part the lavas,
usually basic, are notably vesicular, and the original boundaries of the lava-streams can
often be distinctly traced and their relations to centres of eruption demonstrated.

In Tasmania some of the drowned valleys, like that of the Derwent, seem to have
been partially filled with Pleistocene basaltic lava, and in Victoria, chiefly around
Melbourne and Geelong and in the Western district, the Pliocene eruptions were prolonged
into Pleistocene and Recent time (Hills, 1938). The wonderful and beautifully preserved
phenomena of Recent vuleanism in south-western Victoria, described in detail by Skeats
and James (1937), include breached craters, scoria-cones, lava-tunnels, lava-blisters or
domes and ‘stony rises’ or lava-ridges, all of which are almost as fresh as though they
had been formed but yesterday. The region of Recent vulcanicity continues across the
border into South Australia, where several volcanic centres are found in a belt extending
in a north-west—south-east direction for some 30 miles. From their relation to certain
dune-ridges in the area, which on Crockev’s interpretation may be of earlier Recent date,
it appears that the volcanic rocks of the Mt. Burr Range may be Pleistocene while those
of Mt. Gambier and Mt. Schank are essentially Recent.

In Queeensland (Richards, 1926) post-Tertiary lavas appear in the islands of Torres
Strait and cover large areas in the northern part of the State. The Atherton Tableland,
3,000 feet above sea-level, is formed in large measure of basalt-flows, probably Pleisto-
cene, crowned by very perfect Recent craters, including those occupied by Lakes Barrine
and Hacham. Around the headwaters of the Burdekin River is a great basaltic plateau
of 2,000 square miles with an average altitude of 2,000 feet, between the Hinasleigh and
Herbert Rivers. According to Maitland (1891) streams of slaggy and ropy lava have
flowed down some of the existing valleys from numerous centres, large and small, which
rise aS more or less conspicuous hills above the plateau. The Recent age of the out-
pourings is attested by “chaotic heaps of angular blocks of basalt, tossed in every variety
of disorder’, by the existence of such structures as lava-tunnels and subsidence caverns,
and by the general absence of dissection.

Another immense basaltic area stretches from the vicinity of Charters Towers west
for upwards of 150 miles. It forms a plateau up to 2,800 feet above sea-level, composed
very largely of a somewhat vesicular olivine-basalt, whose surface is diversified by
centres of eruption. One of these which I have examined, Mt. Emu, is a well-preserved
breached crater partly of the explosion type. It is possible that the craters are Recent
and that the flows covering the plateau are Pleistocene; at all events they are almost
certainly post-Tertiary, since they have filled canyons and gorges in the plateau eroded
presumably as a result of the Kosciusko uplift.

In addition to these major occurrences there are numbers of small flows of very
scoriaceous lava, derived from central vents, which have filled existing valleys; examples
are found at the head of the Endeavour River and in the valleys of the Mulgrave,

xX PRESIDENTIAL ADDRESS.

Herbert, Tate and other rivers. Possibly the most southerly manifestation of Recent
vulcanicity in Queensland is in the valley of the Burnett River, where there are large
quantities of fresh, slaggy lava, as well as the very perfect cone and craters of Mt.
Le Brun (Bryan, 1938).

In New South Wales no basalts are known that can confidently be classed as post-
Tertiary.

VERTEBRATE FAUNA.
(Howchin, 1930; Anderson, 19338.)

The fossil-record of the terrestrial deposits shows that there was a great expansion
and development among the vertebrates in post-Tertiary times. As shown by the map
(Fig. 3), for which no claim to completeness is made, they had a wide distribution on
the continent, including even those parts of it which are now the most arid. Their
remains have been found:

(1) In old flood-plains and alluvial terraces and in deep leads, both ‘alluvial’
and basalt-covered;

(2) In lake-, swamp- and spring-deposits;

(3) In caves;

(4) In beds of volcanic ash.

MAP OF

AUSTRALIA

Scale of Miles
0 50 100 200 300

Proved et the Deparcaet of Leste Sytaey NSW, 1040. o> oe 7 |

Fig. 3.—Distribution of known remains of extinct post-Tertiary vertebrates in the Commonwealth. i

PRESIDENTIAL ADDRESS. xxi

Most of the forms found fossil are now extinct, but some belong to existing genera,
and frequently the fossils are much larger than their living representatives.

The most remarkable feature of the fauna is the great variety of herbivorous
marsupials. Of the Macropodidae the largest known member, Palorchestes azael, had a
skull about 16 inches long, and other genera also attained a large size. Many species
were closely related to, but much larger than, the living kangaroo, wallaby and wallaroo.
The largest known marsupial, Diprotodon, allied to the wombats, was as big as a
rhinoceros, and roamed very widely over the continent. Thylacoleo carnifex, the
‘marsupial lion’, of which only the skull is definitely known, seems to have been as
large as a small lion and was possessed of a remarkable dentition which points to
carnivorous habits.

Of the lizards the largest example was Megalania prisca, described from a number
of places in eastern Australia, which was from 14 to 20 feet in length.

Crocodilian remains have been described from the Condamine River and elsewhere
in Queensland, from the Warburton River (South Australia) and from as far south as
the neighbourhood of Port Augusta, South Australia.

Of the large flightless birds the most important genera were Genyornis and Dromornis,
both with affinities to the emu and the cassowary. The genus Dromaeus, to which the
emu belongs, had a wide distribution.

The Tasmanian Wolf (Thylacinus cynocephalus) and the Tasmanian Devil
(Sarcophilus ursinus), now living only in the island-state, have fossil representatives on
the mainland, in both instances larger than the existing species, and fossii remains of
the ‘native cat’, and of marsupial mice and bandicoots, mostly living species, are known
from Wellington Caves and elsewhere.

It was customary in the past to assign a late Pliocene to Pleistocene age to this
fauna, mainly, it would appear, because so much of it is now extinct and because some
examples were found in auriferous leads believed to be late Tertiary. Some of the
deposits, such as the Canadian Lead at Gulgong, New South Wales, in which remains of
the chelonian Meiolania were found, may perhaps be late Tertiary, and the oldest known
Australian marsupial, Wynyardia bassiana, was found in Miocene beds, but the majority
of the deposits are certainly younger than Tertiary. It is reasonable to suppose that the
giant herbivores, which were able to roam all over the continent and were able to attain
a very great size, lived in the Pleistocene Period when rainfall was high and food was
plentiful. But though this was probably the time of their greatest development, reasons
have been adduced for believing that they survived till Recent time, such as the
following:

(1) In a number of places their remains are associated with those of living
marsupial species;

(2) The bones of Diprotodon and other extinct forms have been found with the
remains of .the dingo, which expert opinion nowadays regards as an
immigrant brought to Australia by man in Recent or very late Pleistocene
time;

(3) A number of extinct forms has been found in Mowbray Swamp (Tasmania),
which is in physiographic equilibrium with present-day sea-level, and is
therefore hardly likely to have existed before Recent time.

Various reasons have been advanced to explain the eventual extinction of the giant
fauna, such as their own gigantism, the increasing aridity of earlier Recent time, the
advent of man with his hunting weapons and the introduction of the carnivorous dingo.
Of these possible causes the first two were probably the most important, and in support of
the second it has been pointed out that the fossil skeletons have frequently been found in
old lake- and swamp-deposits, as if the animals had been bogged in trying to reach the
shrinking waters. As against the last two suggested reasons it may be urged that since
the present indigenous animals are to a large extent descended from the fossil forms, it is
reasonable to suppose that during the earlier Recent arid epoch they survived chiefly in
the less arid parts of the continent, e.g., in the coastal belts of the east and south, which
are just the parts that man and his domesticated dingo would chiefly frequent.

XXii PRESIDENTIAL ADDRESS.

POST-TERTIARY FLORA.

As to the nature of the prevailing Pleistocene flora we have little, if any, direct
evidence, but it certainly did not differ much from that of the present day, though its
geographical distribution was vastly different.

Even as far back as early and middle Tertiary time, to judge from the evidence of
the deep leads, a flora existed with leaves resembling very closely indeed those of some
elements of our rain-forest flora, though of course identity is not thereby proven. Certain
younger leads of western Victoria, considered to be late Pliocene or early Pleistocene,
have yielded logs, leaves and seed-vessels, some of which are indistinguishable from
species of Banksia and Hucalyptus now growing in the surrounding country. There is
a suggestion, therefore, that early in Pleistocene time, probably before the onset of
glacial conditions, the local physiography, rainfall and vegetation did not differ very
greatly from those of the present day. We may infer that at a later date, when the
effective rainfall became much greater, there was a big spread of the rain-forest flora
all over the continent, with marked expansion of tropical jungle during the long Middle
Pleistocene interglacial epoch and as marked a shrinkage on the return of colder
conditions. This was the heyday of the giant herbivores. The coming of the earlier
Recent arid epoch heralded a widespread and most destructive change whereby, save
in the coastal belt and in Tasmania, the prevailing flora was wiped out except for a
few small colonies that managed to survive in specially favoured areas. Some of these
colonies, or their descendants, still exist, but without the power of venturing beyond
the bounds of their original refuge. It is in some such way that we must explain, for
instance, the little colony of palms (Livistona Mariae) and cyeads in Palm Creek in
the Macdonnell Ranges (Howchin, 1930), and the solitary remnant of Pandanus palms
in a gorge in the Carr Boyd Range, East Kimberley, Western Australia, recently seen
by Mr. W. H. Maze. Probably, too, the outliers of rain-forest flora on the basalt residuals
of Mts. Tomah, Wilson and Irvine, on the Blue Mountain plateau, are similar relics
preserved from extinction through being rooted in soil of higher fertility and greater
water-holding capacity than that of the surrounding sandstone country.

Another curious survival is the solitary patch of Rhododendrons on the top of the
Bellenden Ker Range in north Queensland. According to David (1932) these probably
migrated from New Guinea during a Pleistocene glaciation, apparently while low tempera-
tures prevailed and a land-bridge to Australia existed.

THE COMING OF MAN.

The most recent and comprehensive summary of data bearing on the problem of the
antiquity of man in the Commonwealth is that compiled by the late D. J. Mahony (1934a).
There seems to be a measure of agreement among anthropologists that the earliest
immigrants into the Commonwealth were of the type of the now extinct Tasmanian
aborigines, and that one or more waves of Australoid type followed much later and
peopled the mainland, displacing or absorbing the previous occupants. If we concur in
the now generally accepted view that the dingo was brought to Australia by man, then,
since it never reached Tasmania, a reasonable inference is that the very primitive
Tasmanians arrived and spread to Tasmania at a time when Bass Strait either did not
exist or was much less extensive than at present, i.e., during a glacial phase, and that
the incursion of the Australian aborigines with their dingoes took place after a
submergence, and probably in late Pleistocene or Recent time.

How far back in Pleistocene time the first migration took place it is difficult to say.
As already mentioned, a continuous land-bridge with Tasmania was most probably in
existence only during the Mindel and/or the Riss glacial stage, but, on the other hand,
during other less severe glacial maxima, the strait may well have been dotted with
islands which were in sight of each other, and it has been argued that the Tasmanian
aborigines in such circumstances could have made their way across by sea from the
mainland.

Some years ago an obvious artefact was found in some high-level gravels, considered
to be outwash glacial deposits and estimated to be possibly 100,000 years old, at Gladstone
in north-east Tasmania, and was accepted by the late Sir Edgeworth David (19238) as

PRESIDENTIAL ADDRESS. Xxiil

evidence of the existence of man in Tasmania in Pleistocene time. The finders of the
flake were quite satisfied that it was embedded in the gravel, and the late Mr. W. H.
Twelvetrees, Government Geologist of Tasmania and a most careful investigator, was
convinced that it could not have been washed down from the surface. Emphatic and
downright disbelief was subsequently expressed (Meston, 1936) that the flake could
have been in gravels of such age in view of its fresh and unworn condition and the
relatively high degree of culture indicated by its workmanship. Since the flake was
never actually seen by a geologist in situ it would appear that the case must be regarded
by geologists as not proven.

However, there are other somewhat more positive geological facts bearing on the
question of the advent of man. The not infrequent association of remains of the dingo
with those of Diprotodon in cave-deposits suggests that the dog and its Australoid masters
were on the mainland in early Recent time, while the finding of stone tomahawks and the
bones of a dugong hacked by a blunt instrument in the Shea’s Creek excavation at
between 3 and 15 feet below H.W.M. indicates human occupation of the site of Sydney
before the Pleistocene Period had quite come to an end. If, as is contended, the river-
terrace on the Maribyrnong River in which the Keilor skull was found is to be referred
to the Riss-Witirm interglacial stage (Mahony, 19430), this would push the advent of
Australoid man back somewhat farther into Pleistocene time, perhaps into the Riss.

It is in their relation to early man that geology and anthropology find common
ground, and in Hurope the subdivision of post-Tertiary time has been to some extent
based on the evidences of successive human cultures. In this country, however, such
evidences have proved rather scanty. Hale and Tindale (1928), and Tindale (1941), by
‘careful investigation of implement-cultures and of layered deposits in caves or rock-
shelters, have established a sequence of five or more cultural stages; there does not
seem, however, to be any definite proof that these go back beyond Recent time, and the
latest stage was apparently that which existed at the time of the advent of the white man.

EPILOGUE.

A little reflection will make it clear that the happenings of the post-Tertiary periods
have played no small part in moulding the character and way of life and in shaping the
destiny of the white inhabitants of Australia. Though the glacial deposits of the Ice
Age have no agricultural or other economic importance here as they have in Hurope and
North America, the post-Tertiary lava-fields and the Pleistocene flood-plains have provided
some of our richest and most productive soils. To the late Pleistocene submergence we
owe many of our most useful harbours and our most picturesque coastal waterways, and
to the mid-Recent emergence the great expanses of level coastal plain and, indirectly,
the magnificent stretches of sandy beach that are such a feature of our coasts.

The event most fraught with serious consequences was the change from the generally
cool to cold and humid climate of the Pleistocene to the stark and unrelieved aridity of
the succeeding epoch. True there have since been climatic oscillations and some
amelioration, but the permanent drought which still grips so much of the continent has
set the hardest and most impregnable barrier to its full population and development.

In the organic world the post-Tertiary periods have been noteworthy for the rapid
rise and as rapid extinction of a gigantic indigenous fauna, and, more important still,
for the arrival of waves of human migrants upon Australian shores. Of many of the
happenings to which I have alluded, aboriginal man was a bewildered spectator and a
victim. In Tasmania he suffered the rigours of an Ice Age, and later he witnessed the
slow drowning of the coasts and the gradual severance of Tasmania and New Guinea
from Australia. No doubt he gazed with astonishment and fear upon the expiring
convulsions of post-Tertiary vuleanism, and, terror-struck, felt the earthquake-shocks
that accompanied the faultings and warpings of the earth’s crust. In the inland he
experienced the full effects of the economic depression resulting from the shrinking of
rivers, the drying-up of lakes and water-holes, and the gradual disappearance of the
animals that may have formed an important part of his food-supply; and through the
mid-Recent emergence he found some of his coastal camping-grounds left miles away
from the sea. Lastly, in historic time, he viewed with well-founded apprehension and

XXiV PRESIDENTIAL ADDRESS.
+

foreboding the invasion of his land by the strange white men who were so quickly to
dispossess and extirpate him.

References.

ANDERSON, C., 1933.—PrRoc. LINN. Soc. N.S.W., 58: ix (with bibliography).
ANDREWS, E. C., 1910.—J. Roy. Soc. N.S.W., 44: 420.
BENNETT, H. T., 1935.—Aust. Geographer, 2 (8): 3 (with map).
BROWNE, W. R., 1934.—Ibid., 2 (4): 13.
————, DuLHuNTY, J. A., and Mazz, W. H., 1944.—Proc. Linn. Soc. N.S.W., 69: 238.
Bryan, W. H., 19388.—Proc. Roy. Soc. Qd., 49: 106.
CHAPMAN, F., 1915.—Rec. Geol. Surv. Vict., 3 (4).

—, 1920.—Proc. Roy. Soc, Vict., 82 (2): 225.
————,, 1928.—Rec. Geol. Surv. Vict., 5 (1).
CorTron, L. A., 1928.—Proc. Third Pan-Pacif. Sci. Congr., Tokyo, 1926: 1777.
Crockgsr, R. L., 1941.—Trans. Roy. Soc. S. Aust., 65: 108.
Daty, R. A., 1935.—The Changing World of the Ice Age.
Davip, T. W. E., 1908.—PrRoc. LINN. Soc. N.S.W., 33: 657.
, 1923.—Proc. Roy. Soc. Tasm., p. 109.
————, 1932.—Explanatory Notes to accompany a New Geological Map of the Commonwealth.
————, and HALLIGAN, G. H., 1908.—J. Roy. Soc. N.S.W., 42: 229.
Davis, C., Day, M. F., and WATERHOUSE, D. F., 1938.—PrRoc. LINN. Soc. N.S.W., 63: 357.
EipWwarps, A. B., 1941.—Proc, Roy. Soc. Vict., 53 (2).: 2338.
ETHERIDGE, R., Junr., Davip, T. W. H., and GrRiMSHAW, J. W., 1896.—Proc. Roy. Soc. N.S.W.,

30: 158.
FENNER, C., 1931.—South Australia: A Geographical Study.
GREGORY, J. W., 1914.—Geogr. J., 43: 656.
HALE, H. M., and TINDALG, N. B., 1928.—Ree. S. Awst. Mus., 4 (1); 145.
Hiuus, #. S., 1938.—Proc. Roy. Soc. Vict:, 51 (1): 112.
————, 1939.—Ibid., 51 (2): 297.
Hiuus, Lorrus, 1914.—Tasm. Geol. Surv., Bull. 18.
HowcHINn, W., 1929a.—Trans. Roy. Soc. S. Aust., 53: 167.
————, 1929b.— Geology of South Australia, 2nd Ed.
, 1930.—The Building of Australia. Part iii, p. 645. Brit. Sci. Guild (S.A.) Handb.

JOHNSTON, R. M., 1888.—The Geology of Tasmania.
Jonss, L. J., 1925—N.S.W. Geol. Surv. Min. Res., 33.
JuTSON, J. T., 1934.—W. Aust. Geol. Surv., Bull. 95.
Lewis, A. N., 1933.—Proc. Roy. Soc. Tasm., p. 67.
————, 1934.—Ibid., p. 75.
, 19385.—Aust. Geographer, 2 (8): 14 and 30.
MADIGAN, C. T., 1938.—J. Roy. Soc. N.S.W., 71 (for 1937-38): 503 (with references).
Manony, D. J., 19438a.—Mem. Nat. Mus. Vict., 13: 7 (with bibliography).
—, D. J., 1943b.—Ibid., 13: 79.
MAITLAND, A. GIBB, 1891.—Qd. Geol. Surv., Publ. 71.
———,, 1919.—- W. Aust. Geol. Surv., Mem. 1: 51.
Mauston, A. L., 1936.—Proc. Roy. Soc. Tasm., p. 85.
NoETLING, F., 1910.—Ibid., p. 231.
NYE, P. B., 1934.—Tasm. Geol. Surv., Bull. 41.
Raeeatr, H. G., 1936.—J. Roy. Soc. N.S.W., 70: 162.
RICHARDS, H. C., 1926.—Rep. Aust. Ass. Adv. Sci., Adelaide, 1924, 17: 275.
, and Hiuu, DorotHy, 1942.—Rep. Gt. Barrier Reef Comm., 5.
SINGLETON, F. A., 1941.—Proc. Roy. Soc. Vict., 53 (1): 1.
SKEATS, EH. W., and JAmzEs, A. V. G., 1937.—Ibid., 49 (2): 245.
SMITH, T. H., and IREDALE, T., 1924.—J. Roy. Soc. N.S.W., 58: 157.
SoLutas, W. J., 1924.—Ancient Hunters, 38rd Ed. Chapter 1.
SPENDER, M. A., 1930.—Geogr. J., 76.
STEERS, J. A., 1929.—Ibid., 74: 341.
————, 1930.—Rep. Gt. Barrier Reef Exped., 1928-29, Sci. Rep., 3, No. 1.
————.,, 1932.—-The Unstable Earth.
——, 1937.—Geogr. J., 89: 1 and 119.
SUSSMILCH, C. A., 1936.—J. Roy. Soc. N.S.W., 70: 265.
TaTE, R., 1879.—Rep. Phil. Soc. Adelaide, 2: 114.
TINDALE, N. B., 1941.—Avust. J. Sci., 3: 144.
, FENNER, F. J., and Haut, F. J., 1935.—-Trans. Roy. Soc. S. Auwst., 59: 103.
Warp, L. K., 1921.—Rep. Third Interstate Conf. on Artesian Water, Adelaide: 7.
, 1941.—S. Aust. Geol. Surv., Bull. 19.
WHITEHOUSE, F. W., 1940.—Univ. Qd. Pap. Dept. Geol., 2, No. 1.
ZEUNER, F. B., 1935.—Geol. Mag. Lond., 72: 350.

PRESIDENTIAL ADDRESS. xXxXV

The Honorary Treasurer, Dr. A. B. Walkom, presented the Balance Sheets for the
year ended 28th February, 1945, duly signed by the Auditor, Mr. S. J. Rayment, A.C.A.
(Aust.); 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 year to be duly made:

President: Ida A. Brown, D.Sc.

Members of Council: Ida A. Brown, D.Sc., Lilian Fraser, D.Sc., Professor J.
Macdonald Holmes, B.Sc., Ph.D., F.R.G.S., F.R.S.G.S., G. D. Osborne, D.Se., Ph.D.,
T. C. Roughley, B.Sc., F.R.Z.S., E. Le G. Troughton, C.M.Z.S., F.R.Z.S.

Auditor: S. J Rayment, A.C.A. (Aust.).

A cordial vote of thanks to the retiring President was carried by acclamation.

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xXX1X

ABSTRACT OF PROCEEDINGS.

SPECIAL GENERAL MEETING.
28th Marcu, 1945.

Dr. W. R. Browne, President, in the Chair.
The following alterations to the Rules of the Society were unanimously adopted:
Rule iii, for lines 7-10 read—‘‘The name of every Candidate for election as an Ordinary
Member shall be exhibited in the rooms of the Society for the four weeks next
following the receipt of the nomination.”
Rule iv, replace the first sentence of the existing rule by the following:
“Hlection to Ordinary Membership shall be by ballot at an Ordinary Meeting
following the exhibition of the Candidate’s name in accordance with Rule iii.”
Rule 1, replace existing rule by the following:
“The Ordinary Meetings of the Society shall be held at such place and time as
shall be decided by the Council.”
Rule lii, line 5, for ten read seven.
Rule liii, replace Section 3 by the following:
“Candidates whose names have been exhibited in accordance with Rule iii shall
be balloted for.”

ORDINARY MONTHLY MEETING.
28th Marcu, 1945.

Dr. Ida A. Brown, President, in the Chair.

The Donations and Exchanges received since the previous Monthly Meeting
(29th November, 1944), amounting to 12 Volumes, 204 Parts or Numbers, 7 Bulletins,
7 Reports and 3 Pamphlets, received from 67 Societies and Institutions and 3 private
donors, were laid upon the table.

PAPERS READ.

1. On Australian Dermestidae. Part iv. Notes and the Description of a New
Genus and Four New Species. By J. W. T. Armstrong.

2. Evidence of an Eustatic Strand-line Movement of 100 to 150 Feet on the Coast
of New South Wales. By Wilson H. Maze, M.Sc.

38. The Cranial Nerves of Neoceratodus. By H. Leighton Kesteven, D.Sc., M.D.

SPECIAL GENERAL MEETING.
18th Apri, 1945.
Dr. Ida A. Brown, President, in the Chair.

It was unanimously resolved that the alterations to Rules approved at the Special
General Meeting of 28th March, 1945, be confirmed.

ORDINARY MONTHLY MEETING.
18th Aprin, 1945.

Dr. Ida A. Brown, President, in the Chair.

The President announced that the Council had elected Dr. A. B. Walkom to be
Honorary Treasurer for the Session 1945-46.

The President also announced that the Council had elected Mr. R. H. Anderson,
Mr. F. H. Taylor, Mr. E. Le G. Troughton and Dr. W. R. Browne to be Vice-Presidents
for the Session 1945-46.

GG

Xxx ABSTRACT OF PROCEEDINGS.

Major H. B. Burgh, Mosman, Mrs. Gwenda L. Davis, B.Sc., Armidale, Major A. N.
Johnston, B.Sc.Agr., Burwood, Private David Mackerras, Cowra, and Mr. Jack Roper,
M.1.H.S., Broken Hill, were elected Ordinary Members of the Society.

The President referred to the death, on 12th April, 1945, of Dr. E. S. Stokes, who
had been a member of the Society since 1905.

The Donations and Exchanges received since the previous Monthly Meeting
(28th March, 1945), amounting to 1 Volume, 21 Parts or Numbers and 4 Bulletins,
received from 21 Societies and Institutions and 1 private donor, were laid upon the table.

PAPERS READ.

1. Stenoporids from the Permian of New South Wales and Tasmania. By Joan
Crockford, M.Sc., Linnean Macleay Fellow of the Society in Palaeontology.

2. Some Points in the External Morphology of the Pouch Young of the Marsupial,
Thylacinus cynocephalus Harris. By W. Boardman.

3. Correlation of some Carboniferous Sections in New South Wales. By Alan H.
Voisey, M.Sc.

WILLIAM FARRER CENTENARY.

The attention of the meeting was directed to the fact that on the 3rd April, 1945,
Australia celebrated the 100th anniversary of the birth of one who has been described —
as its greatest benefactor—William James Farrer.

He was a pioncer investigator in wheat breeding research and the phenomenal
increase in the yield per acre and on the total yields for wheat produced in Australia
for several decades from 1910 onwards was a direct result of Farrer’s work. Farrer’s
wheats over a period of years averaged up to six bushels an acre more than the old
varieties and he gave to this country wheats which resulted in an extension of the
wheat belt, and wheats which permitted development of an export trade on the world’s
markets.

ORDINARY MONTHLY MEETING.
27th JUNE, 1945.*

Dr. Ida A. Brown, President, in the Chair.

Captain R. V. Southcott, M.B., B.S., Adelaide, South Australia, was elected an
Ordinary Member of the Society.

The President, on behalf of members, offered congratulations to Dr. Alan H. Voisey,
on attaining the degree of Doctor of Science of the University of Sydney.

The Donations and Exchanges received since the previous Monthly Meeting
(18th April, 1945), amounting to 5 Volumes, 93 Parts or Numbers, 1 Bulletin, 4 Reports
and 13 Pamphlets, received from 45 Societies and Institutions and 2 private donors, were
laid upon the table.

PAPERS READ.

1. Relations of the Orchid Flora of Australia to that of New Zealand: With the
Description of a New Monotypic Genus for New Zealand. By Rev. H. M. R. Rupp, B.A.,
and E. D. Hatch.

2. Geographic Variation in the Lizard, Hemiergis decresiensis (Fitzinger). By
Stephen J. Copland, B.Sc.

3. A Bryozoan Fauna from the Lake’s Creek Quarry, Rockhampton, Queensland.
By Joan Crockford, M.Se., Linnean Macleay Fellow of the Society in Palaeontology.

ORDINARY MONTHLY MERTING.
25th Jury, 1945.
Dr. Ida A. Brown, President, in the Chair.
Mr. D. F. Ross, Wahroonga, was elected an Ordinary Member of the Society.
The Donations and Exchanges received since the previous Monthly Meeting
(27th June, 1945), amounting to 10 Volumes, 65 Parts or Numbers, 3 Reports and 1
Pamphlet, received from 42 Societies and Institutions, were laid upon the table.

* No meeting of the Society was held in May, 1945.

ABSTRACT OF PROCEEDINGS. TOC

PAPERS READ.

1. On the Family Smarididae (Acarina). By R. V. Southcott, M.B., B.S.

2. Principal Microspore-Types in the Permian Coals of New South Wales. By J. A.
Dulhunty, B.Sc.

3. Miscellaneous Notes on Australian Diptera. xi. Evolution of Characters in the
Order: Venation of the Nemestrinidae. By G. H. Hardy.

4. Studies on Australian Marine Algae. ii. Notes extending the Known
Geographical Range of Certain Species. By Valerie May, M.Sc.

5. Revision of Australian Lepidoptera. Oecophoridae. xiii. By A. Jefferis Turner,
M.D., F.R.E.S.

6. Petrological Studies in the Ordovician of New South Wales. iii. The Composition
and Origin of the Upper Ordovician Graptolite-bearing Slates. By Germaine A. Joplin,
B.Se., Ph.D., Linnean Macleay Fellow of the Society in Geology.

ORDINARY MONTHLY MEETING.
26th SEPTEMBER, 1945.*

Dr. Ida A. Brown, President, in the Chair.

The President, on behalf of members, offered congratulations to Lieutenant G. A. V.
Stanley, R.A.N.V.R., on the award of the Distinguished Service Cross in recognition of
sustained courage and endurance and “skill of a standard beyond the ordinary course of
duty under most hazardous conditions in the Aitape-Wewak area’’.

The President referred to the release of Dr. C. E. M. Gunther from a prisoner-of-war
camp in Singapore.

The President also referred to the death on 2nd September, 1945, of Professor J. T.
Wilson, who had been an Ordinary Member of the Society from 1892 to 1923, an
Honorary Member since 19238, a former Councillor, and President, 1897-1899; and to the
death on 14th August, 1945, of Archdeacon F. EH. Haviland, who had been an Ordinary
Member of the Society from 1911 to 1943 and a Corresponding Member since 1943.
Reference was also made to the death on 22nd September, 1945, of Mr. A. F. Basset Hull,
who had been an Ordinary Member of the Society from 1907 to 1940, a member of
Council for approximately twenty-five years of that period, and President, 1923-1924.

The President announced that the Council is prepared to receive applications for
four Linnean Macleay Fellowships tenable for one year from ist March, 1946, from
qualified candidates. Applications should be lodged with the Secretary, who will afford
all necessary information to intending candidates, not later than Wednesday, 7th
November, 1945.

The Donations and Exchanges received since the previous Monthly Meeting
(25th July, 1945), amounting to 6 Volumes, 70 Parts or Numbers, 2 Bulletins and 1
Report, received from 33 Societies and Institutions and 1 private donor, were laid upon

the table.

PAPERS READ.
1. Observations on the Morphology and Biology of the Subspecies of Anopheles
punctulatus Donitz. By A. R. Woodhill.
2. The Hair Tracts in Marsupials. Part ii. Description of Species, continued. By
W. Boardman.
3. Notes on some Fijian Mosses. By William Greenwood.

LECTURE.
An illustrated lecture entitled ‘In Search of Orchids’”’ was delivered by Rev. H. M. R.
Rupp, B.A.
* No meeting of the Society was held in August, 1945,

xxxii ABSTRACT OF PROCEEDINGS.

ORDINARY MONTHLY MEETING.
31st OcroBer, 1945.
Dr. Ida A. Brown, President, in the Chair.

The President reminded candidates for Linnean Macleay Fellowships, 1946-47, that
Wednesday, 7th November, 1945, is the last day for receiving applications.

The Donations and Exchanges received since the previous Monthly Meeting
(26th September, 1945), amounting to 2 Volumes, 108 Parts or Numbers and 1 Bulletin,
received from 22 Societies and Institutions, were laid upon the table.

PAPERS READ.

1. Notes on Australian Mosquitoes (Diptera, Culicidae). Part vi. The Genus
Tripteroides in the Australasian Region. By David J. Lee, B.Sc.

2. Notes on New South Wales Orchids: A New Species and some New Records. By
Rev. H. M. R. Rupp, B.A.

3. Nitrogen Fixation in Leguminous Plants. vi. Further Observations on the Effect
of Molybdenum on Symbiotic Nitrogen Fixation. By H. L. Jensen, Macleay Bacteriologist —
to the Society.

NOTES AND EXHIBITS.

Mr. E. Cheel exhibited fresh flowering specimens of Bottlebrush, cultivated in his
garden at Ashfield, as follows: (1) Callistemon linearis DC.; (2) C. hortensis Cheel;
(3) C. acuminatus Cheel; (4) Callistemon sp. citrinus (lanceolatus) x acuminatus;
(5) C. linearifolius DC.; (6) C. lilacinus Cheel. When describing this as a distinct
species, he suggested that its origin was probably as a result of hybridism. The
specimen exhibited is from one of the numerous seedlings he has raised from C. hortensis.
Specimens of Leptospermum emarginatum Wendl. were also exhibited.

ORDINARY MONTHLY MEBRTING.
28th Novemserr, 1945.
Dr. Ida A. Brown, President, in the Chair.

Miss Pauline G. Larcombe, Burwood, and Mr. James J. Lawrence, B.Sc., Clovelly,
were elected Ordinary Members of the Society.

The President announced that the Council had reappointed Miss Frances M. V.
Hackney, M.Sc., and appointed Miss June Lascelles, B.Sc., to Linnean Macleay Fellow-
ships in Plant Physiology and Biochemistry respectively for one year from 1st March,
1946.

The Donations and Exchanges received since the previous Monthly Meeting
(31st October, 1945), amounting to 14 Volumes and 12 Parts or Numbers, received from
14 Societies and Institutions, were laid upon the table.

PAPERS READ.
1. Studies on Trombidiidae. By R. V. Southcott, M.B., B.S.

2. A Contribution to a Study of the Physiology of Decay in Apples. By Mary
Cash, M.Sc.

3. Catalogue of Reptiles in the Macleay Museum. Part i. Sphenomorphus pardalis
pardalis (Macleay) and Sphenomorphus nigricaudis nigricaudis (Macleay). By Stephen
J. Copland, B.Sc.

4. Studies in the Metabolism of Apples. vi. Preliminary Investigations on the
Respiration of Sliced Apple Tissue. By Frances M. V. Hackney, M.Sc., Linnean Macleay
Fellow of the Society in Plant Physiology.

5. Contributions to a Knowledge of Australian Culicidae. No. viii. By Frank H.
Taylor, F.R.E.S., F.Z.S. :

6. The Diptera of the Territory of New Guinea. xiii. Family Tabanidae. Part i.
The Genus Chrysops. By Frank H. Taylor, F.R.E.S., F.Z.S.

1940

1927
1905
1940

1922
1899

1927
1938
1912

1919
1940
1935
1940
1907

1941
1929

1923
1924
1941
if yiLa
1943
1931

1945
1920

1901
1927

1930
1934

1905

1945
1936
1899
1932
1901

1942

1931
1942
1908

1928

XXX1il

LIST OF MEMBERS.
(15th December, 1945.) 7+

ORDINARY MEMBERS.

Abbie, Andrew Arthur, M.D., B.S., B.Se., Ph.D., c.o. University of Adelaide, North Terrace,
Adelaide, South Australia.

* Albert, Michel Francois, ‘‘Boomerang”’, 42 Billyard Avenue, Elizabeth Bay, Sydney.

tAlien, Edmund, ‘‘Daintree’’, Curlew Street, Toowong, Brisbane, Queensland.

Allman, Stuart Leo, B.Se.Agr., M.Sc., Entomological Branch, Department of Agriculture,
Farrer Place, Sydney.

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

Andrews, Ernest Clayton, B.A., F.R.S.N.Z., No. 4, ‘““Kuring-gai’’, 241 Old South Head Road,
Bondi, N.S.W.

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

Ashby, Professor Eric, D.Se., D.I.C., F.L.S., Botany School, Sydney University.

Aurousseau, Marcel, B.Sc., c.o. Mr. G. H. Aurousseau, 16 Woodland Street, Balgowlah,
N.S.W.

Barnett, Marcus Stanley, “The Hill’, Victoria Street, Mount Victoria, N.S.W.

Basnett, Miss Elizabeth Marie, M.Sc., New England University College, Armidale, N.S.W.

*Beadle, Noel Charles William, D.Sc., Botany School, Sydney University.

Beattie, Mrs. Joan Marion, M.Sc. (née Crockford), 219 Victoria Road, Gladesville, N.S.W.

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

Blake, Stanley Thatcher, M.Sc., Botanic Gardens, Brisbane, Queensland.

Boardman, William, M.Sec., Department of Biology, University of Queensland, Brisbane,
Queensland.

Brough, Patrick, M.A., D.Sc., B.Se.Agr., Botany School, Sydney University.

Brown, Miss Ida Alison, D.Se., Department of Geology, Sydney University.

Browne, Miss Helen Rowan, 51 Nelson Street, Gordon, N.S.W.

Browne, William Rowan, D.Se., Department of Geology, Sydney University.

Bryan, Clement, B.A., Central School, Boorowa, N.S.W.

*Burges, Norman Alan, M.Sc., Ph.D., Botany School, University of Cambridge, Cambridge,
England.

Burgh, Henry Bertram, 4 Rose Crescent, Mosman, N.S.W.

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

Campbell, John Honeyford, O.B.E., 1.8.0., 336 Chapel Street, Ottawa, Canada.
Campbell, Thomas Graham, Council for Scientific and Industrial Research, Box 109,
Canberra, A.C.T.

Carey, Miss Gladys, M.Sc., 32 Rawson Street, Epping, N.S.W.

*Carey, Samuel Warren, D.Sc., Government Geologist, Department of Mines, Hobart,
Tasmania.

Carne, Walter Mervyn, c.o. Department of Commerce, 337 Collins Street, Melbourne,
Victoria.

Cash, Miss Mary Phyllis, M.Sc., 28 Lavender Street, Lavender Bay, North Sydney, N.S.W.

*Chadwick, Clarence Earl, B.Sc., Royal Hotel, Keira Street, Wollongong, N.S.W.

Cheel, Edwin, 40 Queen Street, Ashfield, N.S.W.

Churchward, John Gordon, B.Sc.Agr., Ph.D., 1 Hunter Street, Woolwich, N.S.W.

Cleland, Professor John Burton, M.D., Ch.M., University of Adelaide, Adelaide, South
Australia.

Cleland, Kenneth Wollaston, M.B., Resident Medical Officer, Goulburn District Hospital,
Goulburn, N.S.W.

Colefax, Allen Neville, B.Se., Department of Zoology, Sydney University.

Copland, Stephen John, B.Sc., 7 Creewood Street, North Strathfield, N.S.W.

Cotton, Professor Leo Arthur, M.A., D.Sc., Department of Geology, Sydney University.

Craft, Frank Alfred, B.Sc., 91 High Street, Taree, N.S.W.

Addresses and degrees as at 31st May, 1946.

* Life Member.
£ Since deceased.

XXXIV LIST OF MEMBERS.

1929 Dakin, Professor William John, D.Sc., Department of Zoology, Sydney University.

1945 Davis, Mrs. Gwenda Louise, B.Se., 143 Mossman Street, Armidale, N.S.W.

1936 Day, Maxwell Frank, Ph.D., B.Sc., Australian Scientific Liaison Office, Room 614 Dupont
Buildings, Washington 6, D.C., U.S.A.

1934 Day, William Eric, 23 Gelling Avenue, Strathfield, N.S.W.

1925 de Beuzeville, Wilfred Alexander Watt, J.P., ‘“Melamere”, Welham Street, Beecroft, N.S.W.

1937 Deuquet, Camille, B.Com., 126 Hurstville Road, Oatley, N.S.W.

1927 *Dixson, Sir William, ‘‘Merridong’’, 586 Gordon Road, Killara, N.S.W.

1937 Dulhunty, John Allan, D.Se., Department of Geology, Sydney University.

1926 Dumigan, Edward Jarrett, State School, Toowoomba East, Queensland.

1941 Edwards, Eric Thomas, Ph.D., M.Sc.Agr., National Press Pty. Ltd., 126-130 Phillip Street,
Sydney.

1932 *Ellis, Ralph, 12, Administration Building, University of Kansas, Lawrence, Kansas, U.S.A.

1943 Ellison, Miss Dorothy Jean, M.Sc., Abbotsleigh College, Wahroonga, N.S.W.

1930 English, Miss Kathleen Mary Isabel, B.Se., 7 Dudley Road, Rose Bay, N.S.W.

1914 Enright, Walter John, B.A., P.O. Box 14, West Maitland, N.S.W.

1930 Fraser, Miss Lilian Ross, D.Sc., ‘““Hopetoun’’, 25 Bellamy Street, Pennant Hills, N.S.W.

1935 *Garretty, Michael Duhan, M.Sc., 477 St. Kilda Road, Melbourne, S.C. 2, Victoria.

1938 Gibbs, William James, M.Sc. (S./L.), R.A.A.F.. Meteorological Services, Box 1289K, G.P.O.,
Melbourne, Victoria. :

19386 Gilmour, Darcy, M.Se., 18 Myuna Flats, Braddon, Canberra, A.C.T.

1944 Greenwood, William Frederick Neville, c.o., Colonial Sugar Refining Co. Ltd., Lautoka,
Fiji.

1910 Griffiths, Edward, B.Se., Department of Agriculture, Farrer Place, Sydney.

1936 Griffiths, Mervyn Edward, M.Sc., 38a Mona Road, Edgecliff, N.S.W.

1939 Gunther, Carl Ernest Mitchelmore, M.B., B.S., D.T.M., Bulolo, New Guinea.

1939 Hackney, Miss Frances Marie Veda, M.Sc., 40 Smith Street, Summer Hill, N.S.W.

1925 Hale, Herbert Matthew, South Australian Museum, Adelaide, South Australia.

1928 Hamilton, Edgar Alexander, 16 Hercules Street, Chatswood, N.S.W.

1917 Hardy, George Huddleston Hurlstone, ‘‘Waldheim’’, Waldheim Street, Annerley, Brisbane,
S.3, Queensland.

1932 Harris, Miss Thistle Yolette, B.Se., 14 Pacific Street, Watson’s Bay, N.S.W.

1930 Heydon, George Aloysius Makinson, M.B., Ch.M., School of Public Health and Tropical
Medicine, Sydney University.

1938 Hill, Miss Dorothy, M.Se., Ph.D., Department of Geology, University of Queensland,

Brisbane, Queensland.
3 Hindmarsh, Miss Mary Maclean, B.Sec., 78 Dover Road, Rose Bay, N.S.W.
30 Holmes, Professor James Macdonald, Ph.D., B.Sc., F.R.G.S., F.R.S.G.S., Department of
Geography, Sydney University.

1943 Horowitz, Benzoin, Eng.Agr.S., Dr.Agr.Se. (Cracow, Poland), Botany School, Sydney
University. ;

1982 Hossfeld, Paul Samuel, M.Se., 132 Fisher Street, Fullarton, South Australia.

1942 Humphrey, George Frederick, M.Sc., Department of Biochemistry, Sydney University.

1987 Hurst, Mrs. Evelyn Anne, B.Sc.Agr. (née Mercer), ‘““‘The Mount’, Wyong Creek, Wyong,
N.S.W.

1938 Ingram, Cyril Keith, c.o. Mrs. C. K. Ingram, Macleay Street, Frederickton, Macleay River,
N.S.W.

1917 Jacobs, Ernest Godfried, ‘‘Cambria’’, 106 Bland Street, Ashfield, N.S.W.
1938 Jacobs, Maxwell Ralph, D.Ing., M.Se., Dip.For., Commonwealth Forestry Bureau, Canberra,
INO,
1930 Jensen, Hans Laurits, D.Se.Agr. (Copenhagen), Department of Bacteriology, Sydney
University.
1945 Johnston, Arthur Nelson, B.Se.Agr., Hawkesbury Agricultural College, Richmond, N.S.W.
1907 Johnston, Professor Thomas Harvey, M.A., D.Sc., F.L.S., University of Adelaide, Adelaide,
South Australia.
1937 Jones, Mrs. Valerie Margaret Beresford, M.Sc. (née May), Botanic Gardens, Sydney.
30 Joplin, Miss Germaine Anne, B.Sc., Ph.D., ‘Huyton’, 18 Wentworth Street, Hastwood,
N.S.W.
Judge, Leslie Arthur, 36 Romsey Street, Hornsby, N.S.W.

2)
930 Julius, Sir George Alfred, B.Sc., B.E., M.I.Mech.E., M.I.E.Aust., 67 Castlereagh Street,
Sydney.

* Life Member.

LIST OF MEMBERS. XXXV

Kendall, Mrs. May Marston, M.Se. (née Williams), 71 Victoria Road, Drummoyne, N.S.W.

Kesteven, Geoffrey Leighton, B.Sc., Fisheries Section, C.S.I.R., Marine Biological Labora-
tory, Cronulla, N.S.W.

Kesteven, Hereward Leighton, D.Sc., M.D., 23 The Crescent, Vaucluse, N.S.W.

Kinghorn, James Roy, C.M.Z.S., Australian Museum, College Street, Sydney.

Langford-Smith, Trevor, Ministry of Post-war Reconstruction, Canberra, A.C.T.
Lascelles, Miss June, M.Sc., 28 Jackson Street, Balgowlah, N.S.W.

Lawson, Albert Augustus, 9 Wilmot Street, Sydney.

Lee, David Joseph, B.Se., c.o. Department of Zoology, Sydney University.

Lee, Mrs. Alma Theodora, M.Sc. (née Melvaine), Botanic Gardens, Sydney.

Liddell, Miss Jean, Department of Biology, University of Adelaide, Adelaide, South
Australia.

Lothian, Thomas Robert Noel, 68 Hewitts Road, Merivale, Christchurch, N.W. 1, New
Zealand.

Mackerras, David, 3 Aust. Recruit Trg. Bn., Cowra, N.S.W.

Mackerras, lan Murray, M.B., Ch.M., B.Se., Veterinary Research Station, Yeerongpilly,
Queensland.

*Mair, Herbert Knowles Charles, B.Sc., Flat 4, ‘““Wellesley’’, 443 Glenferrie Road, Hawthorn,
E.2, Victoria.

Marshall, Alan John, ‘‘Talofa’’, Moran Street, Mosman, N.S.W.

Martin, Donald, B.Sc., Box 17, Huonville, Tasmania.

Mawson, Sir Douglas, D.Se., B.E., F.R.S., University of Adelaide, Adelaide, South
Australia.

Maze, Wilson Harold, M.Sec., Department of Geography, Sydney University.

McCulloch, Robert Nicholson, B.Se.Agr., B.Sc., School of Public Health and Tropical
Medicine, Sydney University.

Mckie, Rev. Ernest Norman, B.A., The Manse, Guyra, N.S.W.

Mercer, Frank Verdun, B.Sc., St. Andrew’s College, Newtown, N.S.W.

Middleton, Bertram Lindsay, B.A., M.D., Bridge House, Murrurundi, N.S.W.

Miller, David, Ph.D., M.Sc., F.R.S.N.Z., F.R.E.S., Cawthron Institute, Nelson, New Zealand.

Milthorpe, Frederick Leon, B.Sc.Agr., Botany School, Sydney University.

Moye, Daniel George, B.Sc., Dip.Ed., ‘“Whiporie’’, Holland Street, Cronulla, N.S.W.

Moye, Mrs. Joan, B.Se. (née Johnston), ‘““‘Whiporie’, Holland Street, Cronulla, N.S.W.

Munch-Petersen, Hrik, Ph.B., M.Sc. (Haunensis), M.I.F., Animal Health Research Labora-
tory, Parkville, N.2, Victoria.

Mungomery, Reginald William, c.o. Meringa Sugar Experiment Station, Box 146, Gordon-
vale, North Queensland.

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

Naylor, George Francis King, M.A., M.Se., Dip.Ed., University of Queensland, Brisbane,
Queensland.

Newman, Ivor Vickery, M.Se., Ph.D., F.R.M.S., F.L.S., Department of Botany, Victoria
University College, Wellington, New Zealand.

Nicholson, Alexander John, D.Sc., F.R.E.S., Council for Scientific and Industrial Research,
Box 109, Canberra, A.C.T.

*Noble, Norman Scott, D.Se.Agr., M.Se., D.I.C., Science House, Gloucester and Hssex
Streets, Sydney.

Noble, Robert Jackson, B.Sc.Agr., Ph.D., 324 Middle Harbour Road, Lindfield, N.S.W.

North, David Sutherland, 42 Chelmsford Avenue, Lindfield, N.S.W.

O’Brien, Brian Robert Alexander, c.o. Post Office, Palm Beach, N.S.W.

Oke, Charles George, 34 Bourke Street, Melbourne, C.1, Victoria.

Osborn, Professor Theodore George Bentley, D.Sc., F.L.S., Professor of Botany, University
of Oxford, Oxford, England.

Osborne, George Davenport, D.Se., Ph.D., Department of Geology, Sydney University.

Pasfield, Gordon, B.Sec.Agr., 60 Whiting Street, Artarmon, N.S.W.

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

Pincombe, Torrington Hawke, B.A., Box 1652, G.P.O., Sydney.

Plomley, Kenneth Francis.

Pope, Miss Elizabeth Carington, M.Sc., Australian Museum, College Street, Sydney.

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

Pryor, Lindsay Dixon, M.Sce., Dip.For., c.o. Department of the Interior, Canberra, A.C.T.

* Life Member.

XXXvVi LIST OF MEMBERS.

1929
1936
1932
1945
1945
1925

1932
1919
1928
1930
1937
1916
1943
1945

1942
1937

1926

1898
1935
1911
1904
1944

1930
1940

1944
1943
1921
1902
1904

1917

1930
1940
1934

1909
1930
1911

1936

1928
1927

1941
1911
1941
1926
1934

1932

1936

Raggatt, Harold George, D.Se., Census Building, Canberra, A.C.T.

Roberts, Noel Lee, 43 Hanna Street, Beecroft, N.S.W.

Robertson, Rutherford Ness, B.Se., Ph.D., Botany School, Sydney University.

Roper, Jack, M.1.H.S., 651 Williams Street, Broken Hill, N.S.W.

Ross, Donald Ford, 31 Braeside Street, Wahroonga, N.S.W.

Roughley, Theodore Cleveland, B.Sc., F.R.Z.S., Chief Secretary’s Department, Box 304.
G.P.O., Sydney.

Salter, Keith Eric Wellesley, B.Se., ‘““Hawthorn’’, 48 Abbotsford Road, Homebush, N.S.W.

«Scammell, George Vance, B.Se., 7 David Street, Clifton Gardens, N.S.W.

Selby, Miss Doris Adeline, M.Sce., M.B., 11 Locksley Street, Killara, N.S.W.

Sherrard, Mrs. Kathleen Margaret, M.Sc., 43 Robertson Road, Centennial Park, Sydney.

Simpson, Arthur Cecil.

Smith, Miss Vera Irwin, B.Sc., F.L.S., ‘“Loana’’, Mt. Morris Street, Woolwich, N.S.W.

Smith-White, Spencer, B.Sc.Agr., 7 Merriwa Street, Gordon, N.S.W.

Southcott, Ronald Vernon, M.B., B.S., 12 Avenue Road, Unley Park, Adelaide, South
Australia.

Spencer, Terence Edward, 16 Attunga Street, Woollahra, N.S.W.

Spencer, Mrs. Dora Margaret, M.Sc. (née Cumpston), 16 Attunga Street, Woollahra,
N.S.W.

Stanley, George Arthur Vickers, B.Sc., c.o. Messrs. Robison, Maxwell and Allen, 19 Bligh
Street, Sydney.

Stead, David G., ‘‘Boongarre’’, 14 Pacific Street, Watson’s Bay, N.S.W.

Still, Jack Leslie, B.Sc., Ph.D., Department of Biochemistry, Sydney University.

*Sulman, Miss Florence, ‘“Burrangong’’, McMahon’s Point, N.S.W.

Sussmilch, Carl Adolph, F.G.S., 11 Appian Way, Burwood, N.S.W.

Sykes, Stephen Myles, B.Sc.Agr., 209 Johnston Street, Annandale, N.S.W.

=tTaylor, Frank Henry, F.R.E.S., F.Z.S., School of Public Health and Tropical Medicine,
Sydney University.

Taylor, Keith Lind, B.Sec.Agr., Forestry Commission, Division of Wood Technology, 96
Harrington Street, Sydney.

Thorpe, Ellis William Ray, B.Sc., Department of Geography, Sydney University.

Tindale, Miss Mary Douglas, M.Se., 60 Spruson Street, Neutral Bay, N.S.W.

*Troughton, Ellis Le Geyt, C.M.Z.S., Australian Museum, College Street, Sydney.

Turner, A. Jefferis, M.D., F.R.E.S., Dauphin Terrace, Brisbane, Queensland.

Turner, Rowland E., F.Z.S., F.R.E.S., c.o. Standard Bank of South Africa, Adderley Street,
Cape Town, South Africa.

Veitch, Robert, B.Sc., F.R.E.S., Department of Agriculture and Stock, William Street,
Brisbane, Queensland.

Vickery, Miss Joyce Winifred, M.Sc., Botanic Gardens, Sydney. .
Vincent, James Matthew, B.Sc.Agr., Dip.Bact., Faculty of Agriculture, Sydney University.
Voisey, Alan Heywood, D.Sc., New England University College, Armidale, N.S.W.

Walkom, Arthur Bache, D.Se., Australian Museum, College Street, Sydney.

Ward, Melbourne, Pasadena Flats, Cross Street, Double Bay, Sydney.

Wardlaw, Henry Sloane Halcro, D.Se., F.A.C.I., Department of Physiology, Sydney
University.

Waterhouse, Douglas Frew, M.Sc., Council for Scientific and Industrial Research, Box
109, Canberra, A.C.T.

Waterhouse, Lionel Lawry, B.E., “Rarotonga’’, 42 Archer Street, Chatswood, N.S.W.

Waterhouse, Walter Lawry, D.Sc.Agr., M.C., D.I.C., Faculty of Agriculture, Sydney
University. :

Watson, Irvine Armstrong, Ph.D., B.Sc.Agr., Faculty of Agriculture, Sydney University.

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

Wearne, Walter Loutit, “Telarah’’, 6 Collingwood Street, Drummoyne, N.S.W.

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

Womersley, Herbert, F.R.E.S., A.L.S., South Australian Museum, Adelaide, South
Australia.

Woodhill, Anthony Reeve, B.Sc.Agr., Department of Zoology, Sydney University.

Zeck, Emil Herman, 694 Victoria Road, Ryde, N.S.W.

* Life Member.

Since deceased.

1923

1902
1942
1943

LIST OF MEMBERS; LIST OF PLATES. XXXVii

HONORARY MEMBER.

Hill, Professor James Peter, D.Se., F.R.S.Z., F.Z.S., F.R.S., Royal College of Surgeons,
Lincoln’s Inn Fields, London, W.C. 2, England.

CORRESPONDING MEMBERS.

Broom, Robert, M.D., D.Sc., F.R.S., Transvaal Museum, Pretoria, Transvaal, South Africa.

Rupp, Rev. Herman Montague Rucker, B.A., 24 Kameruka Road, Northbridge, N.S.W.

Waterhouse, Gustavus Athol, D.Sc., B.E., F.R.E.S., c.o. Australian Museum, College Street,
Sydney.

LIST OF PLATES.

PROCEEDINGS, 1945.

i—Morphology of the Pouch Young of Thylacinus cynocephalus.

ii—iii— Stenoporids from the Permian of New South Wales and Tasmania.
iv—v.—Correlation of Carboniferous Sections in New South Wales.
vi—Geographic Variation in Hemiergis decresiensis (Fitzinger).
vii.—Microspore-Types in the Permian of New South Wales.

viii—The Hair Tracts in Marsupials.

ix.—Nitrogen Fixation in Leguminous Plants.

x.—Diptera of the Territory of New Guinea.

xi.—Reptiles in the Macleay Museum.

xii—xiii— Wings of Tripteroides spp.

XXXVIil

LIST OF GENERA AND SPECIES DESCRIBED AS NEW

IN THIS VOLUME.

(1945.)
Genera.
Page. Page.
Ancistrodes (Lepidoptera: Oecophori- Macrophara (Lepidoptera: Oecophori-
dae) .. ee ee dae) Hat FiG Ee isis eels:
Anthocoma (Lepidoptera: Oecophori- Myrmeanthrenus (Coleoptera: Der-
dae) .. ws 97 mestidae ) 49
Aporostylis (Orchidaceae) . ee 8160 Platyphylla (Lepidoptera: Oecophori-
Colpoloma (Lepidoptera: Oecophori- dae) .. 98
dae) .. Pes tare Teak fl Ke Ptyoptila (Lepidoptera: Oecophori-
Diaphorodes (Lepidoptera: Oecophori- dae) .. Bip oa er evicieer JUKE
dae) .. ose Ie ue) Pyricausta (Lepidoptera: Oecophori-
Epicharactis (Lepidoptera: Oecophori- dae) Ly cei eet) ee eC),
dae) .. st ee OS Stenodiscus (Bryozoa: Batostomel-
Gymnoceros (Lepidoptera: Oecophori- lidae) 21
dae) SEL see OS.
Species.

Page. Page.
alboscutellata (Tripteroides) .. .. 235 micromita (Machimia) eel Ord
albula (Machimia) be tees ers Nace alli miltosticha (Machimia) so IL
aneureta (Macrophara) Os miniatula (Callithauma) . 101
atypa (Lepidotarsa) bo, JMOXD) minuta (Polypora) 50 118383
australiensis (Fessonia) ao aly® mollis (Machimia) : : 5 lite}
baliosticha (Machimia) 5 so JLale moniliformis (Stenodiscus) 5
bisquamata (Tripteroides) .. 56 CD nephospila (Machimia) pealtly
brevicilia (Machimia) .. BK) nicetes (Lepidotarsa) 99
callubriensis (Psacus) .. .. 48 nigronitidum (Trogoder ma) 49
clintoni (Aédes) We Ao. alll nipholeuca (Platyphylla) oo whe
coccoscela (Machimia) signe JEN nissanensis (Tripteroides) .. 559 eu)
collessi (Tripteroides) .. bo 24Uab notoporphyra (Machimia) a LL:
concinna (Tripteroides) o. -260 ochra (Machimia) 56 NOY)
confusa (Tvripteroides ) 55. AGL oncospila (Machimia) 50 1K)
conspersa (Machimia) .. . 106 pallida (Tripteroides) 2 255
contigua (Stenopora) 14 pallidula (Gymnoceros) 98
crocopepla (Tortricopsis) 50. YH parallela (Stenopora) sly)
eryptorrhoda (Machimia) ao daly} paroospila (Zonopetala) -2 Oz
cuphosema (Machimia) ea. JLab phanerozona (Huphiltra) ae . 102
eylicotypa (Machinia) Pay stud phryganophanes (Ancistrodes) .. 98
cyphopleura (Machimia) seg) alalyy picinum (Trogoderma) Saree: 3)
delosticta (Machimia) . 9 ale picturata (Machimia) Bees ta la
dictyosema (Habroscopa) 5.0 OM) pilosa (Tripteroides) 55 CAD
didymosticha (Zonopetaia) ae alos playporphyra (Machimia) seed
dystheata (Machimia) . Bo bales propria (Zonopetala) so JOB
ebenosticta (Machimia) Arey Weeks ee od LG purpurata (Thelymitra) . 288
erythropepla (Tortricopsis ) Bees yt )) pustulosa (Stenopora) .. 5 EAMG
etheridgei (Stenopora) SP aakses ley) pyrrhopasta (Machimia) Bical eal)
eubrocha (Machimia) > lal! rhaphiducha (Machimia) eer Le
euryscia (Diaphorodes) 99 rhodochila (Machimia) epel Sy
euterpnes (Anthocoma) Meroe rhodopechys (Lepidotarsa) . 100
expansa (Fenestrellina) 5.6 Lay) rhoecozona (Machimia) : ae JS
exquisita (Tisobarica) .. og KY rockhamptonensis ee ellina) ~ 128
fraxinea (Colpoloma) ; De ee lO rufescens (Machimia) . . 108
frontalis (Myr meanthrenus) Ran bart) rugosa (Stenopora) Rep mld AS
fuliginosa (Tripteroides) ene O AS. similis (Machimia) 3 ao abil)
fuscipleura (Tripteroides) .. .. 246 simulatrix (Fenestrellina) . Sie JVI
habroschema (Machimia) 59 OY) sparsa (Fenestrellina) Zs
haploceros (Machimia) oo JOY) sparsinodata (Fenestrellina) . 128
hebes (Machimia) .. 5 Jl! spiculata (Stenopora) Meets:
hirsuta (Stenopora) Paces ef spinifera (Fenestrellina) 5 128)
holochra (Machimia) eelelyg splendens (Tripteroides) so 2483
homopolia (Machimia) so JG stenomorpha (Machimia) Bal ea et
interjecta (Machimia) oo JLG stenorrhoda (Machimia) so OT
kingi (Tripteroides) . 256 stygnodes (Machimia) Sell
lechriozona (Wingia) so 1 OB styphlodes (Hoplomorpha) so iily)
lera (Machimia) : tt Seelalts submissa (Machimia) FeelOG
longipalpata (Tripter oides) so) CEM) subobscura (Tripteroides ) 507 FAS
loxomita (Machimia) .. 108 thaumastica (Machimia) SeelOG
metagypsa (Machimia) So Jaks umbratica (Machimia) aa . 108
metaxantha (Machimia) elalto zophosphena (Platyphylla) 98

XXXKIX

GENERAL INDEX.
(1945.)

Address, Presidential, i.

Algae, Australian Marine, Studies on, 121.

Anderson, C., obituary notice, ii.

Anderson, R. H., elected a Vice-President,
OND

Anopheles punctulatus Donitz, Observations
on the Morphology and Biology of the
Subspecies of, 276.

Apples, Contribution to a Study of the
Physiology of Decay in, 317—Studies in
the Metabolism of, 333.

Armstrong, J. W. T., Australian Dermes-
tidae. iv. Notes and the Description of
a New Genus and Four New Species, 47.

Ashby, Prof. E., congratulations to, iii.

Attempted Post-Tertiary Chronology for
Australia. v.

Australia, Attempted Post-Tertiary Chron-
ology for, v.

Australian Culicidae, Contributions to a
Knowledge of, 211—Dermestidae, 47—
Diptera, Miscellaneous Notes on, 135—
Lepidoptera, Revision of, Oecophoridae,
93—Marine Algae, Studies on, 121—Mos-
quitoes (Diptera, Culicidae), Notes on,
219.

Balance Sheets for the Year ending 28th
February, 1945, xxvi-xxviii.

Beadle, N. C. W., congratulations to, iii.

Boardman, W., Hair Tracts in Marsupials,
179—Some Points in the External Mor-
phology of the Pouch Young of the

Marsupial, Thylacinus cynocephalius
Harris, 1.
Browne, W. R., Attempted Post-Tertiary

Chronology for Australia, v—elected a
Vice-President, xxix.
Bryozoan Fauna from the Lake’s Creek
Quarry, Rockhampton, Queensland, 125.
Burgh, H. B., elected a member, xxx.

Carboniferous Sections in New South Wales,
Correlation of some, with Special Refer-
ence to Changes in Facies, 34.

Cash, Mary, Contribution to a Study of the
Physiology of Decay in Apples, 317.
Catalogue of Reptiles in the Macleay
Museum. i. Sphenomorphus pardalis
pardalis (Macleay) and Sphenomorphus
nigricaudis nigricaudis (Macleay), 291.

Cheel, E., see Exhibits.

Chronology, Attempted Post-Tertiary, for
Australia, v.

Coast of New South Wales, Evidence of an
Eustatic Strand-line Movement of 100
to 150 Feet on the, 41.

Contribution to a Study of the Physiology
of Decay in Apples, 317.

Contributions to a Knowledge of Australian
Culicidae, 211.

Copland, S. J., Catalogue of Reptiles in the
Macleay Museum, 291—Geographie Vari-
ation in the Lizard Hemiergis decresi-
ensis (Fitzinger), 62.

Correlation of some Carboniferous Sections
in New South Wales, with Special
Reference to Changes in Facies, 34.

Cranial Nerves of Neoceratodus, 25.

Crockford, Joan M., Linnean Macleay Fellow
in Palaeontology, summary of year’s
work, iv—reappointed, 1945-1946, v—
Bryozoan Fauna from the Lake’s Creek
Quarry, Rockhampton, Queensland, 125—
Stenoporids from the Permian of New
South Wales and Tasmania, 9.

Culicidae, Australian, Contributions
Knowledge of, 211.

(Ko) G2)

Davis, Mrs. Gwenda L., elected a member,
XEXOKG

Davis, H. F., Consett, obituary notice, ii.

Dermestidae, Australian, iv. Notes and the
Description of a New Genus and Four
New Species, 47.

Diptera, Australian, Miscellaneous Notes on,
135.

Diptera of the Territory of New Guinea, xiii.
Family Tabanidae, i. The Genus
Chrysops, 328.

Donations and Exchanges, xxix—xxxii.

Dulhunty, J. A., Linnean Macleay Fellow in
Geology, Principal Microspore-types in
the Permian Coals of New South Wales,
147—resignation from Fellowship, iv—
summary of work, iv.

Elections, xxv, XXx, XXxil.

Evidence of an Eustatic Movement of 100
to 150 Feet on the Coast of New South
Wales, 41.

Exchange relations, ii.

Exhibits:

Cheel, E., Fresh flowering specimens of
Bottlebrush, xxxii—Specimens of Lep-
tospermum emarginatum Wendl., xxxii.

Family Smarididae (Acarina), 173.
Farrer, William, reference to centenary, Xxx.
Fijian Mosses, Notes on some, 213.

Geographic Variation in the Lizard Hemier-
gis decresiensis (Fitzinger), 62.

Greenwood, W., Notes on some Fijian Mosses,
213.

Gunther, C. EH. M.,
a_ prisoner-of-war
EXENEXCI/s

reference to release from
camp in Singapore,

Hackney, Frances M. V., Linnean Macleay
Fellow in Plant Physiology, summary of
year’s work, iv—reappointed, 1945-46,
v—reappointed, 1946-47, xxxii—Studies
in the Metabolism of Apples, 333.

Hair Tracts in Marsupials. ii. Description
of Species, continued, 179.

Hardy, G. H., Miscellaneous Notes on Aus-
tralian Diptera, 135.

Hatch, E. D., see Rupp, H. M. R., and E. D.
Hatch.

x] GENERAL INDEX.

Haviland, Archdeacon F. E., reference to
death, xxxi.

Hemiergis decresiensis (Fitzinger),
graphic Variation in the Lizard, 62.

Hull, A. F. B., reference to death, xxxi.

Geo-

Jensen, H. L., Macleay Bacteriologist, sum-
mary of year’s work, ii—Nitrogen Fixa-
tion in Leguminous Plants, 203.

Johnston, A. N., elected a member, xxx.

Joplin, Germaine A., Linnean Macleay
Fellow in Geology, summary of year’s
work, iv—reappointed, 1945-46, v—Pet-
rological Studies in the Ordovician of
New South Wales, 158.

Kesteven, H. L., congratulations to, iii—

Cranial Nerves of Neoceratodus, 25.

Kosciusko State Park, re preservation of
native flora and fauna of, iii.

Lake’s Creek Quarry, Rockhampton, Queens-
land, Bryozoan Fauna from the, 125.
Larcombe, Pauline G., elected a member,

XSXOXHE

Lascelles, June, appointed Linnean Macleay
Fellow in Biochemistry, 1946-47, xxxii.

Lawrence, J. J., elected a member, xxxii.

Lecture entitled “In Search of Orchids”
delivered by Rev. H. M. R. Rupp, xxxi.

Lee, D. J., Notes on Australian Mosquitoes
(Diptera, Culicidae), 219.

Leguminous Plants, Nitrogen Fixation in,
9X08}.

Lepidoptera, Australian, Revision of, Oeco-
phoridae, 93.

Linnean Macleay Fellowships, 1945-46, reap-
pointments, v—1946-47, applications
invited, xxxi—xxxii—reappointment and
appointment, xxxii.

Mackerras, D., elected a member, xxx.

Macleay Museum, Catalogue of Reptiles in
the, 291.

Marsupials, Hair Tracts in, 179.

May, Valerie, Studies on Australian Marine
Algae, 121.

Maze, W. H., Evidence of an Eustatic
Strand-line Movement of 100 to 150 Feet
on the Coast of New South Wales, 41.

Miscellaneous Notes on Australian Diptera,
135.

Mosquitoes (Diptera, Culicidae), Australian,
Notes on, 219.

Mosses, Fijian, Notes on some, 213.

Neoceratodus, Cranial Nerves of, 25.

New Guinea, Diptera of the Territory of, 328.

New South Wales, Permian of, and Tas-
mania, Stenoporids from the, 9.

Nitrogen Fixation in Leguminous Plants, 203.

Notes on Australian Mosquitoes (Diptera,
Culicidae), vi. The Genus Tripteroides
in the Australasian Region, 219.

Notes on New South Wales Orchids: A New
Species and some New Records, 288.

Notes on some Fijian Mosses, 213.

Observations on the Morphology and Biology
of the Subspecies of Anopheles punctu-
latus Donitz, 276.

Orchid Flora of Australia, relation of, to
that of New Zealand; with the Descrip-
tion of a New Monotypic Genus for New
Zealand, 53.

Orchids, New South Wales, Notes on, 288.

Ordovician of New South Wales, Petrological
Studies in the, 158.

Permian Coals of New South Wales, Prin-
cipal Microspore-types in the, 147.

Permian of New South Wales and Tasmania,
Stenoporids from the, 9.

Petrological Studies in the Ordovician of
New South Wales, iii. The Composition
and Origin of the Upper Ordovician
Graptolite-bearing Slates, 158.

Presidential Address, i.

Principal Microspore-types in the Permian
Coals of New South Wales, 147.

Relation of the Orchid Flora of Australia to
that of New Zealand; with the Descrip-
tion of a New Monotypic Genus for New

Zealand, 53.
Reptiles in the Macleay Museum, Catalogue
of, 291.

Revision of Australian Lepidoptera. Oeco-
phoridae, 93. Y

Roper, J., elected a member, xxx.

Ross, D. F., elected a member, xxx.

Rules of the Society, alterations to, adopted,
xxix—confirmed, xxix.

Rupp, H. M. R., Notes on New South Wales
Orchids: A New Species and some New
Records, 288—lecture entitled “In Search
of Orchids” delivered by, xxxi.

Rupp, H. M. R., and EH. D. Hatch, Relation
of the Orchid Flora of Australia to that
of New Zealand; with the Description
of a New Monotypic Genus for New
Zealand, 53.

Shaw, P. C. W., obituary notice, iii.
Smarididae (Acarina), on the Family, 173.

Some Points in the External Morphology of
the Pouch Young of the Marsupial,
Thylacinus cynocephalus Harris, 1.

Southcott, R. V., On the Family Smarididae
(Acarina), 173—Studies on Trombidiidae
(Acarina), 312—elected a member, xxx.

Stanley, G. A. V., congratulations to, xxxi.

Stenoporids from the Permian of New South
Wales and Tasmania, 9.

Stokes, E. S., reference to death, xxx.

Studies in the Metabolism of Apples, vi.
Preliminary Investigations on the Res-

_piration of Sliced Apple Tissue, 333.

Studies on Australian Marine Algae, 121.

Studies on Trombidiidae (Acarina). Some
Observations on the Biology of the Micro-
trombidiinae, 312.

Tasmania, Stenoporids from the Permian of
New South Wales and, 9.

GENERAL INDEX. xii

Taylor, F. H., Contributions to a Knowledge
of Australian Culicidae, 211—Diptera of
the Territory of New Guinea, 328—
elected a Vice-President, xxix.

Thylacinus cynocephalus Harris, Some Points
in the External Morphology of the Pouch
Young of the Marsupial, 1.

Trombidiidae (Acarina), Studies on, 312.

Troughton, H. Le G., elected a Vice-President,
SRSXGIDXG:

Turner, A. J., Revision of Australian Lepi-
doptera. Oecophoridae, 93.

Voisey, A. H., Correlation of some Carbon-
iferous Sections in New South Wales,
with Special Reference to Changes in
Facies, 34—congratulations to, xxx.

Walkom, A. B., elected Hon. Treasurer, 1945-—
46, xxix.

Wilson, Prof. J. T., reference to death, xxxi.

Woodhill, A. R., Observations on the Mor-
phology and Biology of the Subspecies
of Anopheles punctulatus Donitz, 276.

ia
-
‘
‘
.
.

SOME POINTS IN THE EXTERNAL MORPHOLOGY OF THE POUCH YOUNG
OF THE MARSUPIAL, THYLACINUS CYNOCEPHALUS HARRIS.

By W. BOARDMAN.
(Australian Institute of Anatomy, Canberra.)

(Plate i; three Text-figures. )

[Read 18th April, 1945.]

The Tasmanian Wolf (Thylacinus cynocephalus Harris) seems in danger of being
added to the growing list of extinct marsupials. As long ago as 1842 Owen felt
constrained to comment that its “term of existence seems fast waning to its close’. The
animal, unique in many respects among marsupials, is a more than usually interesting
component of the Australian fauna, but after a history in literature which commenced
in 1808 there are still many gaps in our knowledge concerning it.

The present contribution deals with the external features of the pouch young of
which little of moment has hitherto been published. An account, gathered from adult
material, of the hair tracts in the groin of the male is added for comparison.

Acknowledgments.

My best thanks for the loan of specimens are due to Dr. A. B. Walkom, Director
of the Australian Museum, Sydney, and to the late Mr. D. J. Mahony, formerly Director
of the National Museum, Melbourne.

Material.

The specimens on which these observations were made are in three groups from
three different collections as follows:

(a). Australian Institute of Anatomy.—The formalin-preserved carcase of a mature
male (length of head and body 1170 mm., tail 480 mm.) skinned except over the groin,
tail, parts of the face, and the distal ends of the limbs. In addition, there are two
mounted wet preparations of the scrotum and its investing pouch. This material will
be referred to as the ‘A.J.A. series”.

(b). Australian Museuwm.—Register number 762—a very young furred female
(length of head and body 288 mm., tail 119 mm.) probably still at a stage when using
the shelter of the pouch and being suckled.* It will be recorded as the ‘‘A.M. female”’.

(c). National Museuwm.—Register numbers R.3025-8—four almost naked litter mates
(crown-rump length approximately 75 mm.) comprising two males and two females.
They will be referred to as the ‘“‘N.M. litter’.

So tar as can be ascertained there are no juvenile stages housed in Australian
collections other than those listed above.

Hair.

N.M. litter.—These four individuals are naked except for the sensory vibrissae of
the face and some hair, very pale brown in colour, on parts of the head. The develop-
ment of hair on the head has not proceeded at the same rate in all parts, and this,
combined with its distribution, has in the stage under consideration produced a pattern
which is constant throughout the litter. The hairs are furthest advanced on the face
between the ear and the eye above a sharply defined line joining the lateral angle of the

* Owen (1868), writing of the pouch in Thylacinus, records that “In a female which carried
there three young, each 1 foot in length from the snout ta the end of the tail, the length of the
pouch was 8 inches”’.

D

2 THE POUCH YOUNG OF THYLACINUS CYNOCEPHALUS,

eye to the lower limit of the base of the ear. The crown of the head is also haired; the
hairs of this area are continuous laterally with those in front of the ear as described
above, extend forward to the rhinarium, and finish on an arc, convex cranially, which
runs transversely between the upper limit of the bases of the ears. A few scattered
hairs have pierced the skin of the head more caudally. As is usual in marsupial
embryos there is a growth of fine hairs on the regions where tactile vibrissae are
situated. There are also hairs on the lower eyelid and in its vicinity.

A.M. female.—The body is invested by a close rather coarse fur which shows even
at this early age the crispness found in the adult. The hairs are longest on the face
immediately in front of the ear; they are relatively pale and sparse on the ventral
surface from and including the neck back to the root of the tail, and on the medial
aspects of the limbs. The pelage consists of longer and stouter guard hairs overlying
a more delicate under-fur.

Pigmentation.

N.M. litter—Transverse markings on the back in the caudal half of the body are
not apparent. There is present, however, in parts of the skin, a grey pigmentation
which follows a fairly consistent pattern in each of the four litter mates. The pigmenta-
tion is most marked on the head somewhat above the level of the definitive lip line. It
is relatively concentrated beneath the eyes, and on the muzzle cranial of a transverse
line which runs across the head from slightly caudal of the medial canthus on each side;
further back the pigment intensity is weak and dorsally and laterally it is negligible
behind the level of the vertex. The lateral surface of the ear shows pigmentation in
front of the meatus comparable in intensity with that on the muzzle; there is a weaker
irregular distribution on the auricle generally. A pigmented circular area on the chin
has ill-defined boundaries and a diameter rather larger than that of the mouth aperture.
In addition, the tail is pigmented (weakly in the distal half) except for three or four
millimetres at its root, and there is a tendency for similar colour to develop on the
dorsum of the fore- and hind-feet. .

A.M. female.—The characteristic transverse bands of the hinder half of the body are
clearly laid down (Plate i, fig. 1) and are indistinguishable from those found in the
adult condition.

Hair Tracts.

A.M. female—tThe hair tracts of this individual have been described elsewhere
(Boardman, 1943b). The account given of the groin was incomplete for the reason stated,
and provision of a figure was deferred in the hope of securing further material. There
appears to be little chance of this hope being realized, so that I am here supplementing
the previous description by a figure (Fig. 1) which is as comprehensive as the specimen
will permit.

Pocock (1926), in describing an adult female, records that “The hairs on the abdomen
surrounding the pouch were directed towards it, as in the case of the male; and between
the pouch and the cloacal orifice the points of the hairs were directed inwards towards
the middle line, leaving a narrow strip of naked skin forming an ill-defined passage
between that orifice and the mammary area (Text-fig. 42, p. 1066)”. It will readily be
seen by reference to the accompanying figure and previous description based on data
provided by the young A.M. female, that the two accounts of the hair disposition are not
reconcilable. The difference is similar to that found when comparing Pocock’s account
of hair arrangement in the vicinity of the scrotal pouch of the male (v. infra) and,
in view of the fact that the crisp nature and shortness of the adult hair seems admir-
able for the preservation of the gross features of the hair tracts at all ages, the
discrepancy is difficult to understand. Moreover, Pocock’s description would imply the
presence on the postaxial border of the thigh of a divergent line in place of the hair-
ridge which has been recorded in that situation (Boardman, 1943)D).

A.I.A. male.—Fortunately, on this specimen the skin has been left over the groin
and distal parts of the limbs, and, despite the fact that it is adult, charting of the tracts
presented no difficulty.

BY W. BOARDMAN. 3

Hair radiates from the scrotal pouch in all directions—forwards on to the abdomen,
laterally over the medial aspect of the thigh and towards its postaxial margin (along
this margin it encounters the current which apparently sweeps round over the buttocks
as in the female and forms with it a hair-ridge), and caudally towards the cloaca
(Fig. 2). The general pattern of hair in this region is determined largely by the
presence of the convergent ridge formed postaxially along the thigh as mentioned and
a divergent centre situated in the midventral line at the front of the base of the large
hillock which has at its summit the external opening of the cloaca. On the hair-ridge
running along the postaxial margin of the thigh, about opposite a point midway between
the scrotum and the cloaca, there is a convergent interval which is caused by the

LINE
mN

)

SANS

Figs. 1-2.—Thylacinus cynocephalus. 1. Hair tracts of the groin (A.M. female). The hair
could not be charted on the area left blank. 2. Hair tracts of the groin (A.J.A. male). The
skin had been removed from the regions beyond the zone marked with arrows.

diversion of the opposing currents proximally and distally, respectively. From the
midventral divergent centre, hairs stream forward to meet the current flowing caudally
from the pouch and form with it at the point of impact a divergent interval. Laterally
the deflected hairs from this divergent interval pursue a recurved course outwards and
caudalwards and merge with the current comprising the proximal end of the hair-ridge on
the postaxial margin of the thigh. The current thus augmented describes an are round
the cloacal hillock behind which it divides, the cranial division recurving towards the
base of the hillock while the caudal part proceeds along the ventral surface of the tail
towards its tip; in consequence a midventral convergent interval is formed at the root
of the tail.

Beddard (1891) has figured the scrotum and its pouch in Thylacinus and subse-
quently Pocock (1926) pointed out the inaccuracies in the figure and replaced it with
one of his own. The hair tracts in the vicinity of the scrotum as described and figured
by Pocock are not in accord with the condition observed and outlined above for the
A.J.A. male. Pocock records that “the tips of the hairs all round the pouch were directed
towards it, those in front pointing backwards, those at the side inwards, and those behind
forward”. In the three specimens before me, that is, in the A.J.A. male and two wet
mounted preparations of the scrotum and its pouch (Plate i), the opposite is the case.

The currents on the surface of the scrotum originate on the stalk, from which they
radiate round the sides of the sac to converge at the central point on its ventral aspect.
The figures (Plate i) would seem to indicate that the convergent point is situated more
caudally, but after examination of the scrotum in situ in the A.J.A. male, and considering
the relationships of stalk and sac, I am inclined to regard this as an artefact.

4 THE POUCH YOUNG OF THYLACINUS CYNOCEPHALUS,

Nothing further can be added to the account of the distal portion of the limbs
previously described for the female (Boardman, 1943b); it is of interest that the
convergent interval recorded as occurring laterally on the forearm is equally well
delineated in the same position in this mature male.

Facial Vibrissae.*

In 1914, Pocock, in a paper on the distribution of the various groups of facial
vibrissae in the orders of mammals, took systematists to task for their neglect in
comparative studies of these commonly occurring structures. As a result of his
investigations he was able to state that the specimens he had examined were ‘sufficient
to establish certain general principles as to the constancy or inconstancy of the occur-
rence of the tufts of tactile facial vibrissae within the limits of major groups”. Pocock
was preoccupied with a consideration of the vibrissae group as a whole and no attempt
was made by him to count the actual number of vibrissae present in any particular
situation. Later, Danforth (1925) took up the question of the constancy in occurrence
of the individual bristles in a given situation, choosing the mouse as a type animal and
considering particularly a selected number of vibrissae in the mystacial region. He
found that ‘Individual vibrissae show a remarkable constancy in their one-to-one corres-
pondence with the vibrissae of other animals of the same and even of different species”.
The theoretical significance of this interesting enunciation is not here relevant, but the
suggestion to the taxonomist is pointed. For this reason I have set down in some
detail the numbers, so far as they could be ascertained, of the facial vibrissae in their
various groups in Thylacinus. Such a limited set of figures is, of course, of little or no
statistical value, especially as most of them were secured from litter mates of the same
age. Even so, the tables are of interest in that they imply that equality of number from
specimen to specimen though constant in some situations is not, apparently, constant in
others. For instance, the numerical constancy of rows IV and V of the mystacial group
is, in the material examined, not found in the supraorbitals.

Mystacials.—The mystacial vibrissae are arranged in five principal rows. Above
them there are what may be regarded as three further rows consisting each of only a
single vibrissa placed towards the caudal margin of the mystacial zone; these three
single vibrissae which appear to be constant in their occurrence will not be considered
further. The occurrence of single supernumerary vibrissae between the rows and
alternating with adjacent follicles is not infrequent. In recording, supernumeraries
have been associated with the row to which they are nearest or in cases of doubt with
the row above; their number is indicated separately in brackets (Table 1). The five
rows considered are designated IV to VIII from above downwards. Counting on the
A.M. female was made difficult by the density of the surrounding fur and the possibility
of vibrissae having broken off; this specimen has a further row of weaker bristles with
smaller follicles along the margin of the upper lip inferior to row VIII. No count could
be made on the A./J.A. male.

TABLE 1.
The Distribution of the Mystacial Vibrissae.

R.30259. R.30268. R.30279. R.3028¢. AMS.

Row.
Right. Left. Right. Left. Right. Left. Right. Left. Right. Left.

IV 4 4 4 4 4 4 4 4 5 4
Vv 5 5 5 5 5 5 5 5 5 5
VI 6 6 6 (1) 6 6 (1) 6 6 (1) 5 6 7
Vil 6 6 6 (1) 6 6 (2) 5 6 (1) 6 6 7

VIII 6 6 5 (1) 5 (1) 5 (1) 5 (1) 5 6 5 (1) 52

* No vibrissae other than the various facial groups could be detected on the body at any
stage. }

BY W. BOARDMAN. - 5

The most notable feature presented by these figures is the numerical constancy of
rows IV and V which, in only a single instance, differs from four and five vibrissae
respectively. In the remainder of the series the extent of the agreement (leaving obvious
supernumeraries out of consideration) in homologous rows is very marked.

Supraorbitals.—The supraorbital papilla is not very strongly developed; it is a low
oval structure placed above the margin of the orbit somewhat behind the medial canthus.
In the five specimens considered, the same number of vibrissae occurs on both the right
and left sides except in R.3027 where there is a discrepancy of two, but the number, it
will be observed, shows no constancy from specimen to specimen.

TABLE 2.
The Distribution of the Supraorbital Vibrissae.

Side. R.30259. R.30268. R.30279. R.3028¢. A.M.Q.
Right 7 5 5 6 5
Left 7 5 7 6 5

Genals.—The genal papillae are low, of indeterminate extent and occupy the usual
situation on the face; they are placed beneath the lateral canthus but above the level
of the produced lip line. Figures for the number of vibrissae are available only for
the N.M. series; the vibrissae are arranged on the papilla in two rows which run
obliquely from above downwards and forwards; each row most generally contains a
linear series of four vibrissae, but frequently vibrissae are absent or not developed.

Submentals.—The submental vibrissae occur in three rows which are as well defined
and are as numerically constant as those of rows IV and V of the mystacial region.
Figures could be determined only for the N.M. series. The rows are designated I-III,
I being the lateralmost.

TABLE 3.
The Distribution of the Vibrissae in the Submental Area.
R.30259. R.3026¢. R.30272. R.3028¢.
Row.
Right. Left. Right. Left. Right. Left. Right. Left.
ae ¢
I 3 3 3 3 3 4 3 3
II 4 4 (1) 4 4 4 4 (1) 4 4
IIL 4 4 4 4 a i 4 4 4

Interramals.—The interramal papilla on the N.M. group is, like the other facial
papillae, not very prominent. Of the four specimens two have nine, two ten vibrissae
issuing from the area covered by the papillar structure. It was not possible to count
these vibrissae in the A.M. female.

Lips and Oral Fissure.

The lips of the A.M. female are fully formed. In the N.M. litter, however, they are
sealed laterally in the manner characteristic of the marsupial mammary foetus, a
circular aperture being left beneath the rhinarium for the teat to pass into the mouth
cavity. The definitive lip line is indicated by a groove for most of its length.

Rhinarium.

Pocock (1926) has described and figured the adult rhinarium. The A.M. female
differs in some minor points from his account. In the case of the N.M. group, the
contours are quite different and can be correlated with the adult form only by inference.

N.M. litter—tThe rhinarium is divided throughout into right and left halves by a
deep median sulcus; the depth of the sulcus is such that, when viewed from above, the

6 THE POUCH YOUNG OF THYLACINUS CYNOCEPHALUS,

rhinarium is deeply emarginate cranially. Below, just above the level of the lower
limit of the nostril, the sulcus bifurcates and the two grooves thus formed run to the
lateral margin of the rhinarium beneath the nares and so enclose a triangular area of
skin which forms the upper part of the flesh surrounding the circular teat orifice. This
triangular zone is equivalent to the philtrum of the adult animal; its length is shortened
by modification consequent upon the requirements for the admission of the teat. The
rhinarium is covered with a fine mosaic pattern, the units of which are outlined by
shallow grooves pigmented (dark brown) except immediately above the opening of the
nares. The mosaic area is cut off behind by a fairly clearly demarcated transverse line
which also forms the diameter of an adjoining semi-circular naked zone beyond which
hairs occur. Whether or no this naked part is to be regarded as part of the rhinarium
proper could probably only be determined by sections; the presence or absence of hair
follicles would, in all likelihood, be diagnostic in this case.* ;

The rhinarium extends beyond the margin of the upper lip, as described by Pocock
for the adult, so that when viewed in profile its anterior face recedes downwards and
backwards.

The nostrils are oval in outline with the long axes vertical; the opening is directed
laterally rather than laterally and forwards as in the adult. There is a fairly deep
infranarial area on each side. The philtrum, as recorded above, is shortened consider-
ably by the adaptation of the mouth for sucking.

The rhinarium generally is darkly pigmented.

A.M. female.—tiIn its broad features the rhinarium of the A.M. female does not differ
widely from the description given by Pocock (1926) of the adult condition. It is naked
and its separation from the surrounding haired skin is sharply defined. The philtrum
is very wide and is interrupted below by a low rectangular excision from the corner of
which a pair of grooves arises approximately parallel to each other. These two grooves
and the median sulcus have the same relationship as described by Pocock. The fine
mosaic pattern recorded (v. supra) in the pouch young is present in this larger female,
but there are no signs of pigment deposition.

Eye.

In the N.M. specimens the eyes are unopened but the eyelashes of both the upper
and lower lids have appeared and have pierced the investing epitrichium. The A.M.
female has the eyes open and the upper and lower lids are covered with dense fur in
common with the rest of the head.

mi
r)

External Har.

N.M. litter—These four specimens show the ear at an interesting early level of
development which enables some suggestion to be made with reference to exact nomen-
clature of the parts. The pinna is different in shape from that of the adult, being
bluntly pointed above and to the rear; it is thick and fleshy and is recurved so as to be
adpressed against the skull. The definitive line of the anterior border of the external
opening of the auditory meatus is difficult to determine; this point is considered below
in connection with the A.M. female. Pocock’s ridges a, b and c (1926, Text-fig. 29) are
all clearly developed; c has the relationships of a tragoid projection with a pair of
bulges opposite, which constitute, in all likelihood, a double antitragus (at, Fig. 3 A).

A.M. female.—Pocock (1926) has described at length and figured the adult ear. The
architecture of this structure as found in the A.M. female follows in general Pocock’s
account with only minor variation in the size of the component parts. However, it
seems advisable to provide a figure of this interesting specimen for comparative purposes
and most of the differences will readily be apparent by comparing it with that given
by Pocock. Pocock’s ridge b will be seen to extend further forward in the A.M. female
and there is no continuity in front with the ridge a as would seem to be indicated by
his figure. Ridge c does not appear to have the function of “strengthening the border

*In a description of the rhinarium of Perameles nasuta (Boardman, 1943a) a similar area
was noticed which in that species was, after comparison with the adult, regarded as belonging
to the rhinarium proper. : a Bt : aj ote . -- -

BY W. BOARDMAN. 7

Fig. 3.—Thylacinus cynocephalus. The external ear. A, at the level of development seen
in the N.M. litter; B, in the A.M. female. For explanation of letters see text.

of the inferior passage or cleft in front of the auditory orifice’ since there is in this
young stage very little that could be referred to as a border. It would seem possible
that, in development, the cranial portion of the rim of the wall of the external meatus
has been depressed or folded in two places giving rise to Pocock’s “anterior passage”
between ridges a and 6 and a further passage between ridges b and ¢, so that ridge c has
the relationships of an isolated tragoid projection. Pocock has given a name to the
upper passage, viz., “anterior passage’. It seems that the lower passage between 0b and c
is equally worthy of a name so that if it be desirable to retain the term ‘anterior
passage”, the two could be referred to as the “dorsal anterior passage” (ap, Fig. 3) and
the “ventral anterior passage” (ap’, Fig. 3) respectively.

Feet.

The N.M. litter is too shrunken to be of use for examination in this connection. In
all four specimens the claws are black-tipped.

The A.M. female shows only minor differences from the adult.in so far as the
manus is concerned. The discrete constituents of the plantar pad of the pes show to
better advantage than in older specimens; their arrangement is as recorded by Pocock.

Female Pouch.

The structure of the female pouch is well known. In the earlier phases of its
development as exemplified by the A.M. female and the still earlier N.M. litter there is
little or no indication that the definitive pouch will open backwards. In fact, were the
adult structure not known, it would be quite reasonable to assume that this pouch would
develop to open forwards.

N.M. litter.—Two of these are females. The V-shaped appearance of the early pouch
is present in a more precise form than in the A.M. female; the pouch depression is open
in front, but closed behind. The nipples, two on each side, lie in a groove which is sunk
beneath and parallel with the corresponding lateral lip.

A.M. female.—The forming pouch has at this stage thick lips laterally which converge
and almost meet caudally and are slightly inflected cranially. The pouch thus formed
is excavated for a short distance beneath the lips and also shallowly under the caudal
margin; it is open in front, closed behind. The four nipples, arranged in two pairs, have
the cranial pair spaced further apart than the caudal pair. This pouch shows none of
the cranial overhang such as one finds, for instance, in Dasyurus quoll of an
approximately comparable stage of development.

Literature Cited.
BEDDARD, F. E., 1891.—On the Pouch and Brain of the Male Thylacine. Pyvoc. Zool. Soc. Lond.,
1891: 138-145.
BOARDMAN, W., 1943a.—On the External Characters of the Pouch Young of Some Australian
Marsupials. Aust. Zool., 10: 138-160.

oo

THE POUCH YOUNG OF THYLACINUS CYNOCEPHALUS.

BoARDMAN, W., 1943b.—The Hair Tracts in Marsupials. Part i. Description of Species. Proc.
Linn. Soc. N.S.W., 68: 95-113.

CUNNINGHAM, D. J., 1882.—Report on Some Points in the Anatomy of the Thylacine (Thylacinus
cynocephalus), etc. Rep. Sci. Res. H.M.S. Challenger, Zool., v: 1-192.

DANFORTH, C. H., 1925.—Hair in its Relation to Questions of Homology and Phylogeny. Amer.
J. Anat., 36: 47-68.

OWEN, R., 1868.—On the Anatomy of Vertebrates. Vol. iii. Mammals.

Pocock, R. I., 1914.—On the Facial Vibrissae of Mammalia. Proc. Zool. Soc. Lond., 1914:
889-912.

, 1926.—The External Characters of Thylacinus, Sarcophilus, and Some Related

Marsupials. Ibid., 1926: 10387-1084.

EXPLANATION OF PLATE I.
Thylacinus cynocephalus.
1.—The A.M. female. Note the transverse stripes on the lower back.
2.—A female (R.3027) from the N.M. litter.
3-4.—The two wet preparations of the scrotum and its investing. pouch (A.J.A. series).
Both are from adult or nearly adult animals.

STENOPORIDS FROM THE PERMIAN OF NEW SOUTH WALES AND TASMANIA.

By Joan Crocxrorp, M.Sc., Linnean Macleay Fellow of the Society in Palaeontology.

[Read 18th April, 1945.]

INTRODUCTION.

By far the greatest development of Bryozoa in the Palaeozoic of Australia occurs
in the Permian. Great thicknesses of marine deposits of Permian age occur in Western
Australia, Northern Territory, Queensland,.New South Wales and Tasmania, and in all
of these Bryozoa are extremely abundant, forming in many places thick bryozoal lime-
stones and shales. The bryozoan faunas fall into two provinces, the Eastern Australian,
comprising the faunas of Queensland, New South Wales and Tasmania, and the Western
Australian, which includes also those of the Northern Territory. Although the marine
beds in which they occur are believed to represent a similar range of time during the
Permian, and although several individual species are common to Eastern and Western
Australia, the general type of fauna in these two provinces is widely dissimilar.

From Western Australia, species belonging to twenty genera of the families
Fistuliporidae, Batostomellidae, Fenestrellinidae, Acanthocladiidae, Rhabdomesontidae,
and Sulcoreteporidae have so far been described, and a number of other genera is
known to occur in the fauna; with the exception of the Acanthocladiidae, which are,
however, fairly common, representatives of all of these families are extremely abundant.

In Eastern Australia, the total number of genera so far known to occur in the
Permian is twelve, belonging to four families: Batostomellidae (Batostomella,
Dyscritelia, Stenopora, and Stenodiscus, n. gen.), and Fenestrellinidae (fFenestrellina,
_Polypora, Protoretepora, Minilya), which are extremely abundant; and Rhabdomesontidae
(Rhombopora; and also Streblotrypa, which occurs very rarely and has not so far been
recorded) and Acanthocladiidae (Ptilopora; and Penniretepora, which is also unrecorded),
both of these families being very sparsely represented. Since the collections which have
already been made are, in the case of those from New South Wales and Tasmania at
least, very extensive, it is improbable that the number of genera will be greatly increased
by further collecting, or that abundant representatives of the Fistuliporidae and
Sulcoreteporidae, characteristic of the Western Australian Permian, will be found to
occur here. In spite of this it is hoped that as work progresses on the faunas of both
areas, more species of Bryozoa will be added to the few already known to be common
to the two areas.

The faunas throughout the Upper and Lower Marine Series of New South Wales
and of the Permian sequences in Tasmania have been considered extremely uniform.
Many species of Bryozoa common in these deposits are, however, both widespread
geographically and of restricted geological range, and distinct faunas are found where
similar facies are developed at different horizons; for example, Fenestrellinidae are
especially abundant in the Hunter River district of New South Wales on two horizons,
one just above the Hurydesma Horizon in the Allandale Stage of the Lower Marine
Series, and the other in the Fenestella Shales of the Branxton Stage of the Upper Marine
Series, but although there are species which occur on both horizons, other species are
abundant and widespread on one or other of these horizons, but are not common to
both; and other examples occur of similar facies (characterized by an abundance of
Fenestrellinidae or of ramose or massive Batostomellids) containing distinct faunas
when they are developed on different horizons. Because of this, it is hoped that when
the Bryozoa from the Hobart district, where the Permian sequence is very much faulted,
are better known, they may be useful in correlating these outcrops with the New South
Wales sections.

E

10 STENOPORIDS FROM THE PERMIAN OF N.S.W. AND TASMANTA,

Of the Batostomellidae occurring in the Permian of Eastern Australia, the most
abundant genus is Stenopora, Many species of which occur abundantly at different
localities and on different horizons throughout the sequence. Massive and ramose forms
make up the greater number of those occurring here. Some of these forms should be
of great value stratigraphically, as they appear to be very short ranged, and are in some
cases of wide geographical distribution. Before reliance can be placed upon them in
correlation, identification of species of this genus must, however, be made from an
examination of their internal as well as their external structure, although once a
species has been described, additional specimens can in some cases be identified from
casts of the surface, provided the-cast is well preserved and shows clearly the size and
shape of the zooecial apertures, the number and arrangement of the acanthopores and
mesopores, the occurrence of monticules or maculae and the differences in zooecial
structure associated with them; many records have been made of the occurrence of
described species of Stenopora, so that published records indicate that individual species
occurring in Eastern Australia are long-ranged geologically; these records have
frequently been based on an examination of the general form of the zoarium alone, and
where details of the external structure and the internal structure have been examined, a
wide variety of internal structure has usually been ascribed to the one species; these
records have, therefore, in general no value beyond recognition of the occurrence of
fine ramose (as ‘“Stenopora tasmaniensis”’), coarse ramose (as ‘“Stenopora ovata’) or
massive (as “Stenopora crinita’) species of the genus at different localities. Upon a
number of occasions, these species have also been recorded from the Upper Palaeozoic
of India; these records also have no more significance than recognition of the occur-
rence of similar growth-forms of the same genus; in some cases they have been based
merely upon the occurrence of an impression of the surface of either a ramose or a
massive bryozoan, but in spite of this, their occurrence has been quoted by different
workers in support of correlations between the Indian and Australian sequences.
Stenopora ovata has also been recorded from the Upper Carboniferous of Russia
(Nikiforova, 1933, 8), but this record also apparently refers only to a species of similar
form.

The two previously recorded species which are revised in this paper both occur in
Tasmania as well as in New South Wales. In New South Wales, Stenopora crinita
occurs typically in the highest beds of the Upper Marine Series, the Mulbring Stage
of the Hunter River district and the Crinoidal Shales of the South Coast; small zoaria
of this species occur in the Muree Stage, which underlies the Mulbring Stage in the
Hunter River district, and in beds of similar age in other localities; this species occurs.
in Tasmania at Haglehawk Neck, in the highest marine beds of the Permian sequence.
One of a collection of specimens recorded as this species from the Dilly Stage in the
Springsure district of Queensland has been examined and found to belong to a distinct
species. Stenopora johnstoni Etheridge, 1891, described by Etheridge and later revised
by Hummel (1915) from material from Porter’s Bay, near Hobart, and from beds near
the base of the marine sequence on Maria Island, has been found to occur also at a
number of localities in the Allandale Stage of the Lower Marine Series in New South
Wales.

In addition to the revisions of these two species, a discussion is given here of the
characters of Stenopora ovata Lonsdale, 1844, which was originally described from the
Permian of Tasmania, and has since been frequently recorded from many different
localities.

DESCRIPTION OF SPECIES.
Order TrEposToMATA Ulrich.
Family BATOSTOMELLIDAE Ulrich.
Genus STENOPORA Lonsdale, 1844.
STENOPORA CRINITA Lonsdale, 1845. Text-figs. 3, 4.

Stenopora crinita Lonsdale, 1845, 265, Pl. viii, fig. 5, 5a; Chaetetes crinitus
(Lonsdale), Dana, 1849, 711, T. xi, fig. 7; Stenopora crinita Lonsdale, Nicholson and
Etheridge, 1886, 182, Pl. iv, figs. 1-3, [?] 4, 5, [?] text-fig. 2 A, B; Etheridge, 1891, 49,
Pls. ii, iii, iv, fig. 2, v, figs. 1-4, vi, figs. 3-6, vii, fig. 1, [?] fig. 2.

BY JOAN CROCKIFORD. 11

Lectotype (selected by Nicholson and Htheridge, 1886, 183): is in the British
Museum; the specimen was collected by Strzelecki from “Illawarra, New South Wales”.

Occurrences: Stenopora crinita first appears in the Muree Stage of the Upper Marine
Series at Abbey Green, near Singleton, and in rocks of similar age at Bundanoon, in
the South-western Coalfield, and at each of these localities the zoaria, though fairly
common, are small, not exceeding 2 inches in height; the species is abundant in the
Mulbring Stage in the Hunter River district, and occurs at the northern end of the
Singleton Railway Bridge, in railway cuttings about 1 mile towards Wittingham from
Minimbah Platform, and at Bylong; in the South Coast (Illawarra) district it occurs
at Wollongong and at Broughton village, near Berry, in the Crinoidal Stage (equivalent
to the Mulbring Stage of the Hunter River district), and in the Westley Park Tuffs of
the Crinoidal Stage at Black Head, near Gerringong, and in numerous outcrops
of these tuffs on the coast between Gerringong and Kiama (specimens F 381,
F 12267, F19872-3, F 35541, F 37070, Australian Museum Collection, and 4380, 5445-9,
Sydney University Collection). In Tasmania it occurs in the highest marine beds of
the Permian sequence at Haglehawk Neck (5441, Sydney University Collection) and at
Fitzgerald (Tasmanian Geological Survey specimens). It has been recorded (Whitehouse,
1930, 156) from the Dilly Stage in the Springsure district of Queensland; one of these
specimens has been available for examination; it is a massive species distinguished
from S. crinita by the possession of very numerous acanthopores and other differences
in internal structure.

Large, massive Stenopora, with polygonal zooecia, thin-walled and with fine distant
moniiae; surface with regularly placed monticules; mesopores rare, except in the
monticules; diaphragms practically absent; acanthopores small, not numerous, occurring
at the angles of the zooecial tubes.

The zoarium is massive, very variable in shape, the upper surface being smooth to
strongly lobate; zoaria up to at least 1 ft. in diameter are frequently developed; the
base is usually attached to a shell or a small pebble. It is not often possible to detach
the upper surface of the colony from the matrix, but when exposed it shows raised
monticules, from 3 to 4 mm. in diameter, which are placed with their centres from
§ to 13 mm. apart in specimens from Gerringong and with slightly closer average
spacing in specimens from Singleton; these monticules are raised about 1 mm. above
the general level of the surtace, and are differentiated from the remainder of the
surface by their usually larger zooecia and by the greater number of mesopores they
contain. Typically, the zooecia extend continuously from the base of the zoarium to
the periphery, and so may be 6 in. or more long, but in some specimens the zoarium is
built up of several discontinuous layers of zooecia of variable width, usually at least
1-5 cm. wide. The presence of fine monilae about 2 to 4 mm. apart in the zooecial walls
gives broken colonies the stratified appearance described by Lonsdale, Nicholson, and
Etheridge. The tubes are polygonal (generally six-sided) and angular, and are usually
from 0:35 to 0-55 mm. in diameter, and the number usually occurring in 7 sq. mm. is
from 28 to 50; in the monticules the zooecia are larger than over the rest of the surface,
being up to 0-9 mm. in diameter; some monticules can be distinguished in sections
only by the size of the zooecia, but at times these larger zooecia are accompanied by
aggregations of mesopores, as shown in Text-fig. 4; up to about 17 mesopores may occur
in each monticule.

In sections, the zooecial walls are normally very thin, but increase to 0-13 mm. in
width at the level of the monilae, which are pyriform, with their greatest diameter near
the upper end; individual monilae are up to 0:43 mm. long; they are usually fairly
evenly spaced, between 2 and 4 mm. apart, but are much more crowded in the outer
5 mm. of the colony (Text-fig. 3). The thickened walls at the level of the monilae may,
when cut obliquely in vertical sections, give the appearance of a broad diaphragm,
and were figured as such by Etheridge (1891, Pl. vii, fig. 1) and by Nicholson and
Etheridge (1886, Pl. iv, fig. 5, and Text-fig. 2B), who state (p. 184) that: “Tabulae are
developed from these nodal points, but vary much in their numbers.” True complete
diaphragms, in the form of very thin, slightly concave plates, are developed at extremely
distant intervals, and are so rare that they are virtually absent from the tubes; they

yy) STENOPORIDS FROM THE PERMIAN OF N.S.W. AND TASMANIA,

are not necessarily placed at the level of the monilae, nor are they developed at the
same level in adjacent zooecia. Young zooecial tubes cut in transverse section show as
angular tubes much smaller than the average; the monticules may be marked by large
numbers of mesopores. The epitheca covering the surface between successive layers of
zooecia in specimens in which several layers of zooecia are developed, and which covers
the base of the colony, is a thin undulating plate; the zooecia are horizontal for a short
distance immediately above this epitheca, which is composed of the horizontal portion
of the wall on the lower surface of the tube, then they bend gradually to become vertical
within 8 mm. A single acanthopore frequently occurs at the angles of the zooecia; they
are shown best at the level of the monilae, but are often difficult to distinguish,

Text-figs. 1-2.—Stenopora spiculata, mn. sp. Thin sections of the holotype, x 10.
1. Tangential section, passing through a monticule. 2. Longitudinal section, passing from the
surface, near which the monilae are closely spaced, in towards the central part of the zoarium.

Text-figs. 3-4.—Stenopora crinita Lonsdale. Thin sections of specimens from Gerringong
(6400, 6401, Sydney University Collection), x 10. 38. Longitudinal section, showing the small
monilae, closely spaced near the surface, and becoming widely spaced towards the central part
of the zoarium. 4. Tangential section; the larger zooecia and abundant mesopores in the lower
part of this diagram mark the position of a monticule.

All the text-figures were drawn with the aid of a camera lucida.

ee ee

BY JOAN CROCKFORD. 13

especially in the thin-walled portions of the tubes; as Etheridge has noted (1891, 41, 53),
their distribution is variable, and they are sometimes fairly abundant in one part of a
section, but not common in other parts of the same slide.

Nicholson and Etheridge (1886, 182) figure sections of the holotype of this species
(Pl. iv, figs. 1-3), which is in the British Museum Collection. They described as
“tabulae” the wide bands seen when the monilae are cut obliquely in longitudinal
sections, and state that:

“The growth of the corallum was periodic, and the entire mass... is stratified,
the polygonal corallites showing a slight transverse wrinkling as they approach the
upper surface of each successive stratum. It seems almost certain, however, that
the type-Specimen is only the central portion of a large corallum of which none of
the outer portion is preserved; and there is therefore no reason to doubt that the
corallites in the peripheral region of the corallum would exhibit the characteristic
annulations of the genus”

—which they state that they had observed in other specimens of this species; fine monilae
of the type characteristically found in the central part of zoaria of this form are,
however, well shown in the sections of the holotype which they figure. Etheridge
(1891) has described and figured a large number of specimens and sections of this
species; the infilling of the tubes, which he describes, is frequently developed in
specimens from all the localities. The external form of the colony varies very consider-
ably from massive to coarsely lobate (not ramose), but this variation is not accom-
panied by any differences in internal structure, and is not, therefore, of taxonomic
significance.

Stenopora informis Lonsdale, 1845, is, according to Nicholson and Etheridge,
differentiated from S. crinita by possessing smaller, cylindrical zooecial tubes.

STENOPORA SPICULATA, nh. Sp. Text-figs. 1, 2.

Holotype: 5422, Sydney University Collection.

Horizon and locality: Allandale Stage, Lower Marine Series, Por. 34, Par. Middle-
hope, near Helah Rd. crossing of North Coast Railway.

Massive Stenopora, zooecial walls with prominent monilae, especially in the
peripheral region; acanthopores large, numerous, up to 16 around each tube; mesopores
not numerous, except in the monticules; diaphragms absent.

The zoarium is massive, the lower surface of the holotype is infolded, and the upper
surface is very irregular and is thrown up into a number of folds. The colony is about
7 ecm. in diameter and up to 5 em. high. The upper surface could not be freed from the
matrix, so that the development of monticules is not shown, but their occurrence is
indicated in sections by the presence of small areas, their centres 5 to 7 mm. apart, in
which mesopores are more numerous.

The zooecial tubes are angular where they are thin-walled and oval at the level of
the monilae; their diameter is between 0-4 x 0-36 and 0-87 x 0:6 mm. between, and 0-34
to 0:5 x 0-3 to 0:43 mm. at the level of the monilae, being rather larger in the monticules
than between them. The monilae are more crowded at the periphery, where there are
about twelve rows in the outer 4 mm. of the zoarium, than they are in the central part,
where single rows of monilae spaced 1 to 3 mm. apart usually occur: rarely a zone of
four or five rows of closely spaced monilae occur in this part of the zoarium. The walls
are up to 0:22 mm. thick at the level of the monilae. No diaphragms occur. Mesopores
are not numerous, except in the monticules, where they are slightly more numerous than
the zooecial tubes; they do not at any time completely separate the zooecial tubes. In
7 sq. mm. there are 20 zooecia, with 26 mesopores, where a monticule is included in the
field, and 33 to 36, with 1 to 5 mesopores, between the monticules. Large acanthopores
occur in a single row in the tube-walls, and at the surface up to 16, but usually 12 or
less, occur around each aperture.

This species resembles, in the form of its zoarium, the common Stenopora crinita of
the higher part of the Upper Marine Series; it is readily distinguished, however, by its
internal structure, and on weathered surfaces the spacing of the rows of monilae in the
central part of the colony is closer than in S. crinita.

14 STENOPORIDS FROM THE PERMIAN OF N.S.W. AND TASMANIA,

STENOPORA RUGOSA, n. Sp. Pl. ii, fig. 1; Text-figs. 5-7.

Holotype: 414, Sydney University Collection.

Horizon and locality: Fenestella Shales, Branxton Stage, Upper Marine Series, rail-
way cutting 1 mile west of Branxton.

Discoid Stenopora, with prominent regularly placed monticules; monilae strongly
developed; large acanthopores fairly abundant; mesopores not numerous.

The zoarium is discoid, the largest specimen examined being 10-5 em. long and 9 cm.
wide, and about 2:3 cm. high at its thickest part, tapering gradually towards the
periphery. The base is attached to a shell or pebble. The upper surface is convex, and
shows prominent, very regularly placed monticules, each about 3 mm. wide and raised,
where the surface is well preserved, about 2 mm. above the surface, but they are rather
rapidly removed by weathering; the distance between the centres of adjacent monticules
is 7 to 9 mm. The zooecia in the monticules are thicker-walled than those between
them, and the acanthopores in the monticules are greatly thickened. Mesopores are
more numerous in the monticules than over the rest of the surface, but they are never
abundant. The interspaces between adjacent zooecia are usually rather sharply curved,
but are broad and flat where the walls are more strongly thickened; about six acantho-
pores surround each aperture, being most frequently developed at the angles of the
zooecia; they form short, rather blunt spines, and are greatly thickened in the
monticules. :

The zooecia are tubular; between the monilae they are angular and are from 0-36
in their shorter to 0-51 mm. in their longer diameter; they are rounded and corres-
pondingly smaller at the level of the monilae, where the zooecial walls are up to 0:25,
but usually 0-1 to 0-15 mm., thick. The monilae are short and usually almost confluent.
Mesopores are not abundant; they are rounded at the level of, and angular between, the
monilae, and are up to 0-22 mm. in their longer diameter. No diaphragms occur. In a
field of 7 sq. mm. there are about 19 to 23 zooecia and 14 to 19 mesopores where the area
measured includes a monticule, and 33 to 37 zooecia and 6 or 7 mesopores measured
between the monticules.

STENOPORA CONTIGUA, Nn. sp. Text-figs. 8-10.

Holotype: 5431, Sydney University Collection.

Horizon and locality: Muree Stage, Upper Marine Series, base of cliff section of this
Stage in Loder’s Ck., north of road leading to house, Abbey Green, near Singleton.

Discoid Stenopora, with small, closely spaced monticules; monilae strongly developed,
not confluent; mesopores rare, except in the monticules; acanthopores large, developed
at the angles of the tubes.

The zoarium is discoid, the holotype is 5-4 em. in diameter and is rather less than
1-5 cm. in height at the centre, tapering towards the periphery. Regularly placed small
monticules, each about 2 to 2-5 mm. in diameter, and with their centres 4:5 to 6:5 mm.
apart, occur on the upper surface; these monticules are raised about 1 mm. above the
surface, and are differentiated from the rest of the surface by their thicker-walled zooecia
and by their greatly thickened acanthopores, and also by containing more abundant
mesopores than the rest of the surface. :

The zooecia are tubular, and are angular between and rounded at the level of the
monilae, where they are from 0:3 to 0-43 x 0:33 to 0-45 mm. in diameter; -the walls are
from 0:06 to 0-13 mm. thick at this level, so that the tubes are correspondingly slightly
larger in the thin-walled zones between the monilae. The mesopores are usually small,
from 0-03 up to 0-11 x 0-22 mm. in diameter. Acanthopores are developed usually at the
angles of the tubes, and up to five or six surround each aperture; the acanthopores in
the monticules are strongly thickened. No diaphragms occur. The monilae are usually
separated by a short thin-walled zone and are not confluent. In 7 sq. mm. there are
usually 45 to 50 zooecia and 1 to 6 mesopores; where a monticule is included in the
field there are about 38 to 40 zooecia and 10 to 18 mesopores in the same area.

This species is differentiated from Stenopora rugosa, n. sp., which occurs on a lower
horizon in the Upper Marine Series, by its smaller and more closely spaced monticules,

BY JOAN CROCKEFORD. 15

in which the mesopores are rather move abundant, and by the more distant monilae

in the zooecial tubes, and by its smaller, more closely spaced zooecia.

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Text-figs. 5-7.—Stenopora rugosa, n. sp.

5. Part of the surface of the holotype, including
a monticule, x 10.

6. Tangential section of the holotype, x 10, passing through a monticule in
the upper part of the section. 7. Longitudinal section of the holotype, x 10; the section passes
through the thick-walled tubes of a monticule near the right-hand side.

Text-figs. 8-10.—Stenopora contigua, n. sp. 8. Longitudinal section of the holotype, x 10.
9. Part of the surface of the holotype, x 10, with a monticule shown in the centre of the
diagram. 10. Tangential section of the holotype, x 10, passing through parts of two monticules.

STENOPORA OvATA Lonsdale, 1844.

Stenopora ovata Lonsdale, 1844, 163; Stenopora ovata Lonsdale, Lonsdale, 1845, 263,
Pl. viii, figs. 3-3B; Nicholson and Etheridge, 1886, 173, Pl. iii, figs. 1-4; [non] Stenopora
ovata Lonsdale, Auctt.

Lonsdale, in 1844, briefly described this species from material collected by Darwin
from the southern part of Tasmania. In the following year, he gave a much longer
description, accompanied by three figures, of this species, based on additional specimens
collected by Strzelecki from Mt. Wellington, Mt. Dromedary and Norfolk Plains in
Tasmania. The original specimen collected by Darwin is now lost; Strzelecki’s figured
specimen was placed in the British Museum Collection, and thin sections of this
specimen have been figured by Nicholson and Etheridge (1886), who (pp. 174, 175)
considered this specimen the type, as did Etheridge (1891, 56).

16 STENOPORIDS FROM THE PERMIAN OF N.S.W. AND TASMANIA,

The original descriptions do not form an adequate basis for any records of the
occurrence of this form, nor are the figures of thin-sections given by Nicholson and
Etheridge good enough to show definitely which of several ramose species occurring in
the Permian of the Hobart area of Tasmania should be identified with this species.
From the published descriptions, the type specimen is a ramose zoarium, 1/6 in. to 5/12 in.
in diameter, in which the mature zone, with crowded monilae, occupied about one-half
of the radius, and the central axial zone with thin-walled zooecia is crossed by rather
distant arcuate rows of monilae; mesopores were not abundant; large acanthopores
occur at the angles of the tubes; the monilae in the mature region were closely spaced
and rather elongate; diaphragms were (apparently) absent; the diameter of the zooecial
tubes (according to measurements on figures given by Nicholson and Etheridge) is
about 0-33 to 0-54 mm.

Later authors have referred many different species of coarse ramose Bryozoa to
Stenopora ovata, relying almost entirely on the external form of the zoarium for their
identifications; as a result of this, records of the occurrence of this species, which has
been recorded from India and Russia as well as from Australia, are numerous, and on
several occasions its occurrence has been quoted in support of correlations between
Australian and Indian sequences; but without a revision of this species, based on the
specimen figured by Lonsdale in 1845, it would be impossible to recognize this form with
any certainty. Because of this, none of the coarse ramose species from the Permian of
Tasmania described in this paper is referred to Stenopora ovata.

STENOPORA PUSTULOSA, n. sp. Pl. ii, figs. 2, 3; Text-figs. 22, 23.

Holotype: 5436, Sydney University Collection.

Horizon and locality: Berriedale Limestone, Rathbone’s Quarry 2 miles upstream
from Granton, near Hobart (holotype); same horizon, Collinsvale Quarry, near Hobart
(5438, Sydney University Collection); Telosa Rd., Glenorchy, near Hobart (5439, Sydney
University Collection); Newtown, near Hobart (5440, Sydney University Collection) ;
and Huon Rd., Mt. Wellington, 1,000 ft. above sea-level (2426, Sydney University
Collection ).

Coarse ramose Stenopora, surface when well preserved with prominent monticules ;
zooecial tubes without diaphragms, walls with numerous closely-spaced monilae in the
peripheral zone and a few arcuate zones of monilae crossing the axial zone; acantho-
pores large, placed usually at the angles of the tubes; mesopores almost restricted to the
monticules, in which they are abundant.

The zoarium is ramose, arising from an encrusting base and branching irregularly;
the branches are typically cylindrical, but are readily crushed and distorted; they are
usually 2 to 2-7 mm. in diameter, but are not always of very constant diameter, and the
main branch of the holotype gives off two lateral branches firmly imbedded in the
matrix, which were probably not more than 1:5 em. wide. The surface of the branches
shows prominent monticules, usually about 3 mm. in diameter and with their centres
spaced 4:5 to 7 mm. apart; these monticules are raised about 1-5 mm. above the general
level of the surface and in them the zooecia are thicker-walled and sometimes larger,
and the acanthopores, and especially the mesopores, are more abundant. These
monticules are very noticeable when the matrix has been weathered completely away from
the surface; they are, of course, less obvious when any of the matrix still adheres to the
hollowed parts of the surface between them, and they are also readily removed by
weathering of the surface; but they are always prominent in sections, and the difference
in the numbers of acanthopores and mesopores and in the thickness of the zooecial walls
can still be seen on close examination of the surface of weathered specimens; the
monticules, and the difference in zooecial characters which occurs in them, can also be
distinguished in well-preserved casts of the surface.

The zooecia are tubular, rounded in cross-section at the level of the monilae, where
they are usually from 0-27 to 0:36 mm. in their shorter and from 0-35 to 0-43 mm. in
their longer diameter, although a few zooecia up to 0°48 x 0°55 mm. occur; these larger
zooecia usually occur in the monticules, but on the whole, the zooecia in the monticules
are very little larger than those elsewhere, but they are much thicker-walled than usual.

BY JOAN CROCKFORD. ily

The thickness of the walls at the level of the monilae is usually about 0:11 mm., but
may be up to 0:2 mm. The zooecia are angular in section between the monilae, where
they are very thin-walled and the diameter of the tubes is correspondingly increased.
The axial zone comprises approximately one-half of the radius of the zoarium, and here
the zooecia are thin-walled; arcuate zones of single monilae cross this zone at intervals
of about 3:5 to 11 mm., being usually about 7 mm. apart. The tubes bend rather
gradually from the axial to the mature zone, where small, closely spaced, and distinct
monilae are developed in the walls of the zooecia; in the outer 3 mm. of the zoarium
there are about 11 to 14 rows of these constrictions. Large acanthopores are developed
at the angles of the zooecia and occasionally in other parts of the walls: they frequently
indent the zooecial tubes, and from 5 to 10 surround each tube; as well as these, small
acanthopores are occasionally developed. No diaphragms occur. Small mesopores are
fairly abundant in the monticules, but occur only occasionally over the remainder of
the surface. In 7 sq. mm. there are normally about 35 to 43 zooecia and 4 to 8
mesopores, but from 30 to 41 zooecia and 15 to 35 mesopores occur where a monticule
is included in the field. ©

This large ramose species closely resembles the original illustrations of Stenopora
ovata Lonsdale in its general appearance. It is of such similar size, and fractured
surfaces so closely resemble Lonsdale’s figure of S. ovata that it is possible that it is
the same species. For two reasons, however, it is here considered preferable to give this
form a new name until its identity with Stenopora ovata can be confirmed definitely.
Firstly, coarse ramose Stenoporids are abundant in the Permian strata of the Hobart
district, and there are other species of similar size and general appearance which could
equally well be compared with S. ovata: Nicholson and Etheridge (1886, 174-5) have
selected the specimen figured by Lonsdale as the neotype of S. ovata, and comparison of
the specimens here described with the neotype would be necessary to definitely identify
this form with S. ovata; although both Lonsdale (1845, 263) and Nicholson and Etheridge
(1886, 173) have given long descriptions of the specimen figured by Lonsdale, none of
these authors mentions the monticules which are so prominent on the surface and in thin
sections of S. pustulosa, and the figures of thin sections given by Nicholson and
Etheridge are very small and do not adequately illustrate the characters of the species;
if their magnifications are correct, the tubes in the type of S. ovata are larger than in
these specimens (unless the tubes in Nicholson and Etheridge’s figure are drawn from
a monticule, in which case the mesopores would probably be more numerous than they
are shown), and also the monilae are very much longer in the figures of S. ovata—2 of
the monilae in S. ovata being equivalent to 5 or 6 in S. pustulosa. Secondly, the name
“Stenopora ovata’ has been so indiscriminately applied to such a wide variety of forms
in Hastern Australia, India and Russia that, until this characteristic form from the
Berriedale Limestone can be directly compared with the neotype, it is better to give it
an unambiguous new name which will permit its use in local stratigraphy without the
confusion which the name “Stenopora ovata” at present implies.

STENOPORA HIRSUTA, n. sp. PI. ii, fig. 6; Text-figs. 11-13.

Holotype: F 3200, Commonwealth Palaeontological Collection.

Horizon and locality: Berriedale Limestone, Granton, opposite Bridgewater, near
Hobart (holotype); same horizon, Collinsvale Quarry, near Hobart (5442, Sydney
University Collection); same horizon, Rathbone’s Quarry, 2 miles upstream from
Granton (6419, Sydney University Collection).

Coarse ramose Stenopora, surface.with occasional small maculae: zooecial tubes
without diaphragms; walls with numerous, almost confluent monilae in the peripheral
zone, which makes wp one-third to one-half of the radius, thin except where crossed by
occasional arcuate rows of monilae in the axial zone; acanthopores very abundant, in
two or three rows between the apertures; mesopores rarely developed.

The zoarium is ramose, composed of cylindrical branches 6-5 to 8 mm. in diameter,
and branches at an acute angle at distant intervals. The surface is smooth, with
occasional spot-like or elongate maculae in which the zooecia are thicker-walled and the
mesopores and acanthopores more abundant. The zooecial apertures are oval, with their

18 STENOPORIDS FROM THE PERMIAN OF N.S.W. AND TASMANIA,

C

longer axis parallel to the direction of growth, and are 0-32 to 0-41 mm. long and 0-21
to 0:3 mm. wide. The interspaces between the apertures are flattened and bear numerous
acanthopores; these occur in one, more usually two or three, rows on each interspace,
and about 20 surround each aperture; the acanthopores are of two sizes, the larger ones
being more abundant than the smaller. The apertures are irregularly arranged, there
are about 10 to 12 in 5 mm. longitudinally, and about 38 to 42 in 7 sq. mm., with 1 or 2
mesopores in the same area; the mesopores are slightly more numerous in the maculae.
The zooecial tubes are thin-walled in the axial zone, which comprises one-third to
one-half of the radius; in the mature zone the walls are up to 0-22, but usually about
0:15 mm., thick between two apertures transversely, and up to 0:3 mm., but usually about
0-2 mm., thick longitudinally; the monilae are long and are almost confluent. The
zooecia bend rather sharply from the axial to the mature zone. No diaphragms occur.

Text-figs. 11-13.—Stenopora hirsuta, n. sp. Thin sections of the holotype, x 10. ills
Longitudinal section. 12. Transverse section. 13. Tangential section.

Text-figs. 14-16.—Stenopora etheridgei. n. sp. Thin sections of topotypes (6402 A-C,
Sydney University Collection), x 10. 14. Longitudinal section. 15. Tangential section. 16.
Transverse section.

Text-figs. 17-19.—Stenopora parallela, n. sp. Thin sections from specimens in the same
piece of shale as the holotype, x 10. 17. Tangential section. 18. Transverse section. 19.
Longitudinal section.

BY JOAN CROCKEORD. 19

This species has finer branches than those of Stenopora tasmaniensis Lonsdale 1844,
and is also distinguished from Lonsdale’s specimens and from specimens described as
S. tasmaniensis by Bassler (1941, 173) by its far more abundant acanthopores, and by its
proportionately much wider mature zone with more numerous rows of monilae.

STENOPORA PARALLELA, n. sp. Text-figs. 17-19.

Holotype: 2437, Sydney University Collection.

Locality: Huon Rd., Mt. Wellington, Tasmania, 1,000 ft. above sea-level (holotype) ;
same horizon, “Mt. Wellington” of Strzelecki, Strickland Ave. track, 1 mile west of
Cascade (2438, Sydney University Collection). ;

Ramose Stenopora; walls strongly thickened, with weakly-developed monilae, in the
peripheral zone, and thin except where crossed by arcuate rows of monilae in the axial
zone; zooecial tubes without diaphragms; mesopores rare; acanthopores of two sizes,
fairly abundant.

The zoarium is ramose, with cylindrical branches about 4:5 mm. in diameter, which
bifurcate at distant intervals. The surface of the zoarium is smooth, neither monticules
nor maculae being developed. The zooecial apertures are oval, with their longer axes
parallel to the length of the branches, and are from 0:32 to 0-46 mm. long and from
0-17 to 0:24 mm. wide. The interspaces between adjacent apertures are rounded, and
acanthopores are not often developed. Mesopores are rare; they are oval with their
long axes parallel to the length of the branches, and are up to 0-18 mm. long and 0-1 mm.
wide. The apertures are arranged in longitudinal (sometimes rather oblique) and
poorly defined diagonal rows; about 9 apertures occur longitudinally in 10 mm., and
in a field of 7 sq. mm. there are 34 to 40 zooecia and about 5 mesopores. Small solid
maculae occasionally occur.

The peripheral zone has a radius of 0-75 to 1:0 mm., and the bend from the axial
to the peripheral region is at an angle of about 45°. In the axial zone the tubes are
very thin-walled, and are angular in cross-section; this zone is crossed by an occasional
arcuate row of monilae. In the peripheral zone the walls are 0-08 to 0:28 mm. thick;
monilae are rather weakly developed. No diaphragms occur. The acanthopores are of
two sizes; up to five, but generally fewer, large acanthopores occur around each
aperture; the smaller acanthopores are more abundant, but are irregularly developed.

STENOPORA ETHERIDGEI, nh. sp. PI. ii, fig. 4; Text-figs. 14-16.

Holotype: 5434, Sydney University Collection.

Horizon and iocality: Allandale Stage, Lower Marine Series, Jackson’s Hill, Por. 132,
Par. Pokolbin; same stage, Por. 7, Par. Allandale, where road from Allandale to Rothbury
crosses Black Ck.—horizon a of Walkom, 1913, 122 and Pl. ix (5435, Sydney University
Collection).

Fine ramose Stenopora, with oval apertures arranged in rough longitudinal and
diagonal rows; no maculae or monticules; mesopores rare; acanthopores numerous, in
a single row on the interspaces; peripheral zone narrow, with moniliform walls.

The zoarium is ramose; the branches are cylindrical, but because of the great
relative width of the axial zone they are readily crushed, and usually appear flattened.
Uncrushed specimens are about 3-5 to 4.5 mm. in width; the width of flattened specimens
is increased up to 5-5 mm. The zooecial apertures are oval, usually from 0-31 to 0-4 mm.
long, although apertures up to 0-54 mm. long occur, and they are 0-13 to 0:24 mm. wide.
They are not regularly arranged, although they may in places form rough longitudinal
or diagonal rows. Small mesopores occasionally occur. There are no monticules nor
maculae. In 7 sq. mm. there are about 31 to 36 zooecia, and 0 to 5 mesopores. Blunt
acanthopores of two sizes, of which the larger are very much the more numerous, occur
in a single row on the interspaces between the apertures; there are usually from 12 to 15
around each aperture at the surface.

The zooecial walls are strongly thickened in the mature zone, which is from 0-47 to
0-8 mm. wide, and in this distance there are usually three to five rows of almost
confluent monilae; occasional arcuate rows of single monilae cross the axial zone, in
which the zooecial tubes are usually very thin-walled. The tubes bend gradually from
the axial to the mature zone. No diaphragms occur.

20 STENOPORIDS FROM THE PERMIAN OF N.S.W. AND TASMANIA,

Text-figs. 20-21.—Stenodiscus moniliformis, n. sp. Thin sections of the holotype, x 10.
20. Longitudinal section. 21. Tangential section, passing partly through a monticule.

Text-figs. 22-23.—Stenopora pustulosa, n. sp. Thin sections of the holotype, x 10. 22.
Tangential section, passing through a monticule near the centre of the diagram. 23
Longitudinal section.

STENOPORA JOHNSTONI Etheridge, 1891. Text-figs. 24, 25.

Stenopora johnstoni Etheridge, 1891, Pl. vii, fig. 7, and p. 59; Stenopora johnstoni
Etheridge, Hummel, 1915, 74, T. viii, figs. 4, A, B.

Neotype: The slide figured by Etheridge, and the specimen from which it was made
(here considered the type of the species), are lost. Since this specimen was collected
from Tasmania, from either Porter’s Bay or Maria Island, specimen 5433, Sydney
University Collection, from the marine beds which outcrop along the shore below the
fresh water Porter’s Hill Beds, is selected as neotype.

Horizon and locality: Tasmania: Marine beds just below fresh water Porter’s Hill
Beds, on shore below Porter’s Hill, Sandy Bay, near Hobart (holotype); just above
Eurydesma horizon, Darlington, Maria Island (recorded by Etheridge and Hummel) ;
New South Wales: Allandale Stage, Lower Marine Series, Por. 34, Par. Middlehope,
near Eelah Road crossing of North Coast Railway (5432, Sydney University Collection),
same Stage, above Eurydesma Conglomerate in railway cutting east of Allandale Station
(5448, Sydney University Collection); same Stage, “Maluna’, Pokolbin (F 12233,
Australian Museum Collection).

Frondescent Stenopora, zoaria rather thin, flattened, often of large extent; tube walls
with numerous crowded monilae in the mature zone, axial zone with arcuate rows of

BY JOAN CROCKFORD. 21

5; mesopores not
abundant; diaphragms absent; inconspicuous monticules developed on the surface.

The zoaria are frondescent, arising from small encrusting bases; typically they
appear about 4 to 5 mm. thick—the axial zone is readily crushed, and this has usually
reduced the thickness of the zoaria considerably, perfect zoaria being about 11 mm. thick;
the thickness of the colony has in some specimens been increased by overgrowths on
the surface, where part of the colony has been rejuvenated; the zoaria taper gradually
near the growing tip of the frond. The largest zoarium examined (from near Helah)
is an incomplete undulating frond about 12 cm. long and 5:5 em. wide. Very slightly
raised monticules, up to 3 mm. in diameter and with their centres spaced 6 to 10 mm.
apart, occur rather regularly on the surface. The zooecia are much thicker-walled, the
acanthopores more crowded, and the mesopores usually more abundant in these
monticules.

The zooecia are tubular, and the apertures are usually rather elongated parallel to
the direction of growth. The tubes are sub-circular to oval in section at the level of the
monilae, where their size usually ranges (in specimens from the type locality) from
about 0:24 to 0:33 x 0°36 to 0-45 mm., although groups of larger zooecia occur. The
thickness of the walls at the level of the monilae is usually about 0-15 mm., but may be
up to 0:25 mm.; between the monilae the walls are very thin, and when the zooecia are
cut at this level they are angular and are proportionately larger. The mesopores are
similarly either sub-circular or angular in cross-section when cut at different levels;
they vary, when cut at the level of the monilae, from a very small size up to about
0-25 x 0-17 mm. in diameter. Large acanthopores are abundant, usually in a single
row, in the zooecial walls, up to about 18 surrounding each aperture; a few of these
acanthopores are usually very much smaller than the majority. No diaphragms occur
in the neotype, but extremely infrequent complete diaphragms occur in some of the
sections of other specimens. The central thin-walled part of the zoarium is readily
crushed; in the few specimens with this central part better preserved, its width is up
to about 5°55 mm. Arcuate rows of thickening, each composed of a single row of
monilae, cross this central part. Monilae are closely crowded and often confluent in
the walls in the mature zone; in the neotype, this zone is 1:5 to 2-2 mm. thick and
10 to 14 rows of monilae occur in this distance; similarly crowded monilae occur where
the thickness of a zoarium has been increased by overgrowths. In specimens in which
the thickness of the zoarium is greater than in the neotype, the width of the mature
zone and the number of rows of monilae are proportionately increased. There are from
23 to 38 zooecia, and from 1 to 10 mesopores, in 7 sq. mm. in specimens from the type
locality; in specimens from Eelah, there are 25 to 42 zooecia, and 3 to 23, but usually
less than 8, mesopores in the same area. Mesopores do not appear as abundant at the
surface as in sections, probably due to infilling.

Stenopora frondescens Crockford, 1943, from the Westley Park Tuffs of the Upper
Marine Series at Gerringong, is similar in the form of its zoarium to this species, but
differs widely in the details of its external, as well as its internal, structure. Stenopora
spiculata, n. sp., which occurs with this species at Eelah, is a massive form, but does
show a general resemblance to this species in some details of its internal structure;
in it, however, the zooecia are rounded rather than oval, the acanthopores smaller and
less numerous, the mesopores more generally abundant, and the monilae are larger in
the axial zone and are more widely spaced in the mature zone.

monilae; acanthopores numerous, in a single row on the interspaces ;

Genus STENODISCUS, Nn. gen.
Stenopora Lonsdale, Lee, 1912, 147; Bassler, 1929, 54; [non] Stenopora Lonsdale,
1844, 178; [non] Stenopora Lonsdale, Bassler, 1941, 173.
Ramose Batostomellidae, internal structure as in Stenopora, except that thin complete
diaphragms occur fairly frequently in the zooecial tubes.
Range: Carboniferous to Permian.
Genotype: Stenodiscus moniliformis, n. sp.

bo
bho

STENOPORIDS FROM THE PERMIAN OF N.S.W. AND TASMANTA,

STENODISCUS MONILIFORMIS, nh. sp. PI. ili, figs. 1-3; Text-figs. 20, 21.

Holotype: F 37475, Australian Museum Collection.

Horizon and locality: Berriedale Limestone, Collinsvale Quarry, near Hobart,
Tasmania.

Ramose Stenodiscus, with broad mature zone, in which the tube walls are strongly
moniliform; surface with imconspicuous monticules; acanthopores small, fairly
numerous, in a single row in the interspaces; mesopores not abundant; thin, complete
diaphragms of frequent occurrence.

The zoarium is ramose, with cylindrical branches, circular to sub-circular in cross-
section, and from 13 to 17 mm. in diameter. Incomplete specimens are up to 10 cm.
long; bifurcation of the branches occurs at distant intervals. The surface is in general
smooth, but inconspicuous, irregularly placed monticules, 2 to 5 mm. in diameter, occur;
these are only very slightly raised, but in them the zooecia are larger and thicker-walled
than usual, and the mesopores are rather more abundant. The interspaces between the
zooecia at the surface are flat or slightly rounded, and the acanthopores are not
prominent.

Text-figs. 24-25.—Stenopora johnstoni Etheridge. 24. Tangential section of a topotype
(64038, Sydney University Collection), x 10. 25. Longitudinal section of the neotype, x 10.

The zooecia are tubular, sub-circular in section; the tube-walls are thin, except
where they are crossed by remote arcuate zones of small single monilae, in the axial
zone, which comprises one-half to one-third of the radius; the tubes bend gradually to
the mature zone, in which moniliform thickenings are strongly developed. In section
the monilae are pear-shaped, with their greatest thickness (usually up to 0:17 mm.,
but sometimes more) near their upper ends; they are very crowded, there being about
18 rows of monilae in the outer 3 mm. of the colony. At the level of the menilae the
zooecia are normally from about 0-19 to 0:29 x 0-14 to 0-22 mm. in diameter, but the
zooecia in the monticules are considerably larger, being up to 0-43 mm. in their longer
and 0-38 mm. in their shorter diameter; in the thin-walled zones between the monilae
the zooecia are proportionately larger. Small mesopores of varying size occur
occasionally. Normally there are from 70 to 75 zooecia, and about 4 to 8 mesopores, in
7 sq. mm., but where a monticule is included in the field there are from 48 to 58 zooecia,
and 8 to 19 mesopores, in the same area. Small acanthopores, rather granular in appear-
ance, occur generally in a single row in the interspaces between the zooecia; up to 12
occur around each aperture. Thin, slightly concave, complete diaphragms are
frequently developed in the tubes. The growth of the zoaria has in places been discon-
tinuous, two or three successive layers of zooecia frequently occurring around part of
the mature zone; these do not increase the size of the zoarium, but serve to bring the
surface, in parts of the colony in which growth has been interrupted, up to the general
level of the surface. In specimens of this species, the axial thin-walled zone has
frequently been crushed, or else broken down, the space so formed being filled either
with clear calcite or with fine sediment.

BY JOAN CROCKEFORD.

bo
oS)

Brown masses comparable to those described by Cummings and Galloway (1915, 351),
but which are not enclosed by cysts, although they are usually resting upon a diaphragm,
are rather commonly shown in the zooecial tubes of specimens of this species; in
addition to these larger brown masses, smaller rounded (occasionally elongate) brownish
globules, about 0:15 mm. in diameter, frequently occur, mostly in groups immediately
above a diaphragm; although these structures are not described in detail here, photo-
graphs of them are given on PI. iii.

Stenoporids in which complete diaphragms are developed have been described from
the Carboniferous of Great Britain and North America, and the Permian of India,
Russia and Timor, but none of the species so far described compares closely with this
species in the details of its internal structure.

SUMMARY.

In this paper ten species of Stenoporids are described and figured from the Permian
of New South Wales and Tasmania; of these, seven are described as new species of
Stenopora Lonsdale, and revision is made of two previously described species of
Stenopora, S. crinita Lonsdale, 1844, and S. johnstoni Etheridge, 1891; one species from
the Permian of Tasmania is described as the type of Stenodiscus, n. gen.; a discussion is
also given of the frequently recorded Stenopora ovata Lonsdale, 1844. In addition, a
short general discussion of the occurrence and distribution of Bryozoa in the Eastern
Australian Permian is included in the introduction.

BIBLIOGRAPHY.

Bassuser, R. S., 1929.—The Permian Bryozoa of Timor. Paldontologie von Timor, xvi Lief.,

XXViii.
, 1941.—Generic Descriptions of Upper Paleozoic Bryozoa. J. Wash. Acad. Sci.. 31

(CH) 2. TR

CROCKFORD, JOAN, 1943.—Permian Brvyozoa of astern Australia, Part iii: Batostomellidae and
Fenestrellinidae from Queensland, New South Wales, and Tasmania. J. Roy. Soc. N.S.W.,
“6 (for) 1942)): 258.

Dana, J. D., 1849.—Fossils of’ New South Wales. In Wilke’s U.S. Exploring Expedition, x,
Geology, Appendix and Atlas.

DUNCAN, HELEN, 1939.—Trepostomatous Bryozoa from the Traverse Group of Michigan. Mich.
Univ. Mus. Paleont., Contr. 5 (10): 171.

ETHERIDGE, R., Jnr., 1891.—A Monograph of the Carboniferous and Permo-Carboniferous
Invertebrata of New South Wales. Mem. Geol. Surv. N.S.W., Palaeont., 5 (2).

HuMMEL, K., 1915.—Uber einige Fossilien aus der unteren Dyas von Tasmanien. WN. Jb. Jin.
Geol. Palaont., 1 (1915): 68.

Ler, G. W., 1912.—The British Carboniferous Trepostomata. Mem. Geol. Surv. G.B., Palaeoint.,
iL (83) 3: assays

LONSDALE, W., 1844.—(In Darwin, C.) Geological Observations on Volcanic Islands.

, 1845.—(In Strzelecki, P. E, de) Physical Description of New South Wales and Van
Diemen’s Land.

NIcHOLSON, H. A., and ETHERIDGE, R., Jnr., 1886.—On the Tasmanian and Australian Species
of the Genus Stenopora Lonsdale. Ann. Mag. Nat. Hist., (5) 17: 178.

Nikirorova, A. I., 1933.—Stratigraphic Range of the Carboniferous Bryozoa of the U.S.S.R.
Trans. Un. Geol. Prosp. Serv. U.S.S.R., Fasc. 268.

Watkom. A. B., 1913.—Stratigraphical Geology of the Permo-Carboniferous System in the
Maitland-Branxton District: with some Notes on the Permo-Carboniferous Palaeogeography
of New South Wales. Proc. LINN. Soc. N.S.W., 38 (1): 114.

WHITEHOUSE, F. W., 1930.—Report on a Collection of Fossils made by Mr. J. H. Reid, in the
Springsure District. Qd. Govt. Min. J., 31: 156.

EXPLANATION OF PLATES II-III.
Plate ii.

Fig. 1.—Stenopora rugosa, n. sp. Part of the surface of the holotype, x 1.

Figs. 2-3.—Stenopora pustulosa, n. sp. 2. Topotype (5437, Sydney University Collection),
showing the prominent monticules characteristic of this species, x 1. 3. Holotype, x 1; the
monticules are less prominent in this specimen, since the surface is rather weathered.

Fig. 4.—Stenopora etheridgei, n. sp. Holotype, x 1; the furrow along the centre of each
branch is caused by the crushing and flattening of the zoarium.

Fig. 5.—Stenopora johnstoni Etheridge. ‘Surface of a small part of a topotype (F 2702,
Australian Museum Collection), showing the large but only slightly raised monticules.

24 STENOPORIDS FROM THE PERMIAN OF N.S.W. AND TASMANIA.

Fig. 6.—Stenopora hirsuta, n. sp. Holotype, x 1. (A small fragment of a very fine ramose
zoarium, indicated by an arrow, belongs to a species of Streblotrypa; this genus has not
previously been recorded from Eastern Australia, although it is very abundant in the Permian
of Western Australia and Timor.)

Plate iii.

Figs. 1-3.—Stenodiscus moniliformis, n. sp. 1. Part of the holotype, x 1. 2. Longitudinal
section of the holotype, x 20. 3. Tangential section of the holotype, x 20. The large brown
masses and smaller brown globules which are commonly shown in thin sections of specimens
of this species are shown in these last two photographs.

Fig. 4.—Stenopora spiculata, n. sp. Tangential section of the holotype, x 20.

a a ial ea ti a att a

25

THE CRANIAL NERVES OF NEOCERATODUS.
By H. LetgHTon KESTEVEN, D.Sc., M.D.
(Four Text-figures. )

[Read 28th March, 1945, ]

INTRODUCTION.

Early in 1939 Mr. Heber Longnan, Director of the Queensland Museum, sent me a
very perfectly preserved specimen of Neoceratodus (Epiceratodus). I wished to check
again the innervation of certain of the muscles, and for his very prompt reply to my
request I thank him very much. It seemed a pity to destroy so fine a specimen for so
small an investigation, so I decided to dissect out the roots of the cranial nerves and
describe them together with their distribution and their relations to other structures.

Before commencing the description of the nerves I wish to remark on that which
I regard as the most interesting result of the work. I was very forcibly struck by the
fact that although the brain was very much longer than that of the fishes the anterior
end of the medulla was in precisely the same position relative to the otocrane as that
of the fishes. It is probable that the presence of the lateral cranial fenestra emphasized
this identity of relationship. I then rediscovered an obvious fact, one which, of course,
has been known to all anatomists for very many years. The rediscovery was interesting
because, although so obvious, it has never, so far as I can ascertain, been categorically
stated nor has it been given recognition. The failure to take cognizance of the relation
of the medulla to the otocrane has resulted in the misinterpretation of archaic fish
skulls by palaeontologists, notably Watson (1925), Save-Séderbergh (1936), and Westoll
(1948).

Although the brain of Neoceratodus anteriorly to the medulla is very much longer
than in any of the fishes, the hind-brain is, as already stated, in precisely the same
position relative to the otocrane as that of the fishes. Room has been made for this
elongation in the sphenoidal region of the skull. In the fishes this part of the cranial
cavity is occupied by the olfactory peduncles. In the amphibians the sphenoidal part
of the cranial cavity is occupied by the elongated fore-brain and olfactory lobes as in
Neoceratodus. In the Sauria there is a return to the fish condition, only the olfactory
peduncles are lodged in the sphenoidal cavity. In the Theria room is made for the
expanding brain, and the cavity is terminated abruptly by the lamina cribrosa and there
is no sphenoidal cavity. Throughout the Vertebrata the anterior end of the medulla is
to be found in very close proximity to the transverse plane of the anterior limit of the
otocrane, and of course the pituitary body is always situated just a little farther forward.

All this is almost platitudinous, the facts are so obvious and have been well known
for decades. But although that is so, it has not been taken cognizance of and on that
account is well worthy of restatement.

A reference to the palaeontological papers referred to will reveal that these
investigators have located the fore end of the hind-brain in the anterior part of the
sphenoidal cavity and the pituitary body actually in the ethmoidal region. The incon-
gruity of their conclusions has not been realized by them for they have agreed to
designate the bone beneath the hypothetical pituitary fossa basisphenoid. We are
presented with a basisphenoidal bone divorced by nearly half the total length of the
skull from the otocrane, or in the alternative a basisphenoid bone which underlies the
whole length of the orbital region.

DESCRIPTION OF THE Roots AND DISTRIBUTION OF THE NERVES.

The olfactory peduncles are very short and thick, and, moreover, they expand
almost at once so as to embrace the postero-medial one-third of the olfactory capsule
(Fig. 1).

r

fishes.

THE CRANIAL NERVES OF NEOCERATODUS,

This shortness of the peduncles is in marked contrast with fish brains. The fore-
brain has grown much further forward in the skull than is the case in any of the
Comparing the condition here with that of the fishes, accommodation for the
increased size of the brain had been obtained by utilizing and enlarging the sphenoidal

weet tt tse

-
,

eee We al-

|

Semi-schematic dorsal view of the distribution of the cranial nerves.

Ch.t., chorda tympani. My., anterior myloid nerve. My.p., posterior myloid nerve, the
motor nerve to muscle CSV1. II1., optic nerve. IIi., oculomotor nerve. IV., the pathetic nerve.
V., ophthalmic division of the fifth nerve. V.2, mandibular division of the fifth nerve. V.®*,
maxillary division of the fifth nerve. ‘The motor twigs to the muscles of mastication not
shown. V-VII l.a., anterior division of the trigemino-facialis lateralis nerve. V-VII l.p.,
posterior division of the same nerve. VIII hy., hyomandibular division of the facialis nerve.
VII m., motor twigs of the facialis nerve to the CS2, interhyoideus and levator hyoidei muscles.
VII pal., palatine division of the same. VIII 1, labyrinthine division of the eighth nerve.
VIII v., vestibular division of the same. IX br., branchial division of the ninth nerve. IX pal.,

palatine division of the same nerve. X br., branchial branches of the vagus nerve. X g. i.,
gastro-intestinal division of the same nerve.

Fig. 1.—Neoceratodus.

X-V-VII lat., posterior lateralis nerve.

i

BY H. LEIGHTON KESTEVEN. 27

extension of the cranial cavity which, throughout the fishes, contains only the long,
narrow olfactory peduncles. In the fishes, in the great majority of instances, the
anterior limit of the brain is located no further forward than the posterior limit of
the orbit; in Neoceratodus the fore-brain lies medially to the orbit, extending forward
almost as far as its anterior boundary. This is not due to a forward shifting of the
whole brain, because the pituitary body and the optic chiasma are located in the same
position relative to the orbit and to the rest of the skull as it is in the fishes.

Another factor determining the brevity of the olfactory peduncles is that the
olfactory capsules have expanded in a caudal direction by the utilization of that
cavitation of the ethmoidal mass, which is so constant a feature of the fish skull.

The nasal capsule is remarkably large. Its sensitive mucosa is apparently confined
to the roof and a small segment of the posterior wall. It does not appear to be generally
appreciated that Neoceratodus has no external nostril. The anterior narial apertures
are situated on either side of the mid-line within the margin of the upper lip. The
lip itself is broadly grooved immediately within its border and this groove leads to the
anterior naris on the roof of the mouth. The posterior nares are also located in the
roof of the mouth, further back and a little more laterally. There is no partition or
other obstruction on the floor of the olfactory capsule between these two apertures; a
probe introduced into either may be pushed straight out through the other if it is but
kept against the floor with a little pressure to pull that floor downward. Just in front
of the angle of the mouth both upper and lower lips are grooved to provide an entrance
to a peculiar, laterally compressed cavity (bursa innominata) lined by the oral mucosa.
This appears to have been regarded by some, at least, of the recent students of the
fossil Crossopterygians as the external naris of the Dipnoi. The cavity in question
is not connected with the olfactory capsule at all. It extends dorsally in the side of
the mouth and backward until it lies medially to the depth of the anterior boundary
of the orbit, and also forward until it lies laterally to the lateral wall of the olfactory
capsule. The two cavities together call to mind the single cavity in the ethmoidal
region of Latimeria (Smith, 1940).

The optic nerves of Neoceratodus are rather thinner than might have been expected,
and there is nothing of particular interest to record in connection with their course or
relation to contiguous structures.

The location of the orbit relative to the skull is, however, of some interest. The
eye itself is small (a feature wherein the Dipnoan resembles the rest of the Amphibia
rather than the fishes) and has been located far forward in the space between the two
main massives of the cranium. The static location of the hinder half of the brain is
again emphasized by consideration of the varying location of the eye to the skull. It is
placed further back in the generality of the fishes than it is in Dipnoans, and further
forward in the rest of the Amphibia than it is in these. The optic chiasma, however,
remains in the same relative location and the length of the optic nerve is varied to meet
the different distances to the eye.

The Oculomotor Nerve.

The third nerve arises from the base of the brain a short distance caudal to the
pituitary body, and, as is usual, by a single root. The point of origin is medial to that
longitudinal eminence on the ventrum of the brain which is due to the descending
pyramidal fibres. The nerve runs forward and laterally for a short distance across the
ventrum of the mid-brain, then laterally through the loose connective spongework* in
which the brain is packed within the cranial cavity, to its own canal in the lateral wall.

with the fishes rather than the rest of the Amphibia. In the Dipnoan the space between the
brain and the wall of the cranial cavity is, as in the fishes, quite extensive. It is occupied by
fine strands of connective tissue none of which has any tensile strength and all of which
are well separated from each other, so that the brain may be said to be lightly suspended
in a bath of serous fluid by a multiplicity of fine, weak strands of tissue. Of course it rests
upon the floor of the cavity. In the rest of the Amphibia the brain case is more nearly a
reasonable ‘fit’? for the brain, the empty space is much less and the packing tissue more
plentiful and stronger. ¥ :

28 THE CRANIAL NERVES OF NEOCERATODUS,

This canal is situated behind and ventral to the optic canal; it runs through the wall
diagonally forward and laterally. The nerve emerges on to the outer surface of the
skull behind and below the optic nerve but above the ophthalmicus profundus. It crosses
this last nerve before it reaches the “wall of the orbit’, but has divided into two
branches before doing so.

Its distribution to the ocular muscles is without comparative interest.

The Pathetic Nerve.

The fourth nerve arises, by a single root, from the dorsum of the mid-brain half-way
between the posterior boundary of the pineal body and the anterior boundary of the
cerebellum. It is a larger nerve than might have been anticipated and quite readily
found. Its course is parallel to that of the third nerve but further forward and dorsally.
It also emerges from the cranium through a foramen of its own. This runs nearly
parallel to the optic canal and almost directly above it but just a little behind it. The
nerve emerges dorsally to the optic and has but a short course before penetrating the
“wall of the orbit’”,* close to its inner, deep end.

The Trigeminal Nerve.

The fifth nerve arises from the anterior end of the medulla, laterally to both the
pyramidal and lateral columns. It was not possible to separate the roots of this nerve;
they are so closely bound together that the nerve appears to arise by a single root
composed of a number of bundles of strands of fibres. The number of the bundles was
quite indeterminate and varied in four cases, apparently with the skill and patience
devoted to their separation.

The root passes across the intervening space and enters the prootic foramen before
the gasserian ganglion is reached. From the ganglion, the R. ophthalmicus profundus
is first given off. This runs directly forward in the thickness of the cranial wall. The
early embryonic stages, however, indicate quite clearly that the portion of the wall
lateral to the nerve is the processus ascendens quadrati and that the posterior boundary
of that structure is the anterior wall of the canal through which the rest of the nerve
runs to reach the external aperture of the prootic foramen.

The anterior aperture of the ophthalmicus profundus canal is situated low down on
the side wall of the cranium a short distance posterior to the posterior wall of the
orbit. As soon as the nerve emerges it splits into smaller superior and larger inferior
divisions. The former runs forward and dorsad and divides into two fine terminal’
branches which pass forward and round the orbit, medially, in the subcutaneous tissues.
The inferior division passes forward close to the inferior edge of the skull, it rises
slightly so as to pass above the optic and oculomotor nerves and the attachment of the
dorsal periphery of the orbital wall to the cranium. The nerve lies against the ascending
process of the palatine as it passes dorsad to reach the dorsal surface of the skull just
at the lateral end of the fronto-prefrontal suture. In this situation it lies behind the
dorsal wall of the olfactory capsule and divides here into four terminal branches. Two
of these run forward in a canal lodged in the cartilaginous tectum nasi, but dorso-
laterally to the capsule, the other two branches are similarly situated but pass dorso-
medially, lying ventrally to the terminal branches of the ophthalmicus superficialis VII.
This branch of the fifth nerve appears to carry no lateralis fibres, for none of its twigs
was traced to a special cutaneous sense organ or canal.

After giving off the profundus trunk the gasserian ganglion fuses with that of
the lateralis trunk of the facialis nerve. The actual exchange of fibres effected by this
fusion cannot be determined by dissection, but it appears that fibres from the trigeminalis
root pass to the lateralis nerve and that fibres from the lateralis pass into the ramus
maxillaris trigemini.

* This term “wall of the orbit’ has been used above and it might be well to explain that
all the oculomotor muscles and the terminal segments of their nerves, the optic nerve,
ophthalmic artery and the eye itself are wrapped about by a loose, but fairly dense and
relatively thick sheath of connective tissue which is attached to the deep layers of the skin
all round the eye. It is this sheath which is being designated as above.

BY H. LEIGHTON KESTEVEN. 29

The whole of the ganglionic mass is lodged within the prootic canal.

From the external aperture of this canal the following nerves emerge, and in the
following order from before backwards. The short motor twigs of V3 to the muscles of
mastication, the mylohyoid nerve, the ramus maxillaris trigemini, the ramus ophthal-
micus superficialis VII and three or four twigs from lateralis VII. The main trunk of
the lateralis VII turns caudally at the outer end of the canal and enters the lateralis
canal instead of emerging with the other nerves (Fig. 2).

The motor twigs to the muscles of mastication are four in number; they enter the
contiguous surfaces of the two great muscle masses very soon after emerging from the
canal, two to each muscle.

The rest of the mandibular division of the fifth nerve continues forward in the
layer of loose connective tissue which intervenes between the contiguous surfaces of
the two muscles. Just posterior to the fusion of the tendons of the muscles, this
“myloid” nerve reaches the surface. From here it runs around the combined tendons
close to their insertion, first forward, then laterally, and finally caudally until the
posterior opening of the myloid canal is reached.

RMartr ROSVIT a! Cran vad Surface

GEA eon
Canal

Rook V Rock Wi

Fig. 2.—Neoceratodus. The roots, ganglia and branches of the fifth and seventh nerves.

R. max. V., ramus mawxillaris of the fifth nerve. R.o.p. V., ramus ophthalmicus profundus
of the fifth nerve. R.o.s., ramus ophthalmicus superficialis of the seventh nerve. Other lettering
as in Fig. 1.

For a very short distance after entering this canal the direction of the nerve is
transversely out from the mid-line and ventrad. It then turns abruptly forward and
runs along the full length of the mandible to the posterior limit of the very broad
symphasis. At the point where this nerve turns forward it gives off the posterior
myloid nerve, which is apparently a purely motor nerve to the M. intermandibularis. As
it reaches its termination the nerve gives off a number of small branches.

A very short distance in front of the point where the posterior myloid nerve is
given off the main nerve is joined by a communicating branch of the hyomandibular
division of the nervus facialis (the interior mandibular nerve) which entered the
myloid canal with it but which does not fuse until after the smaller branch has been
given off (Fig. 3).

The maxillary division of the trigeminal nerve runs forward between the muscles,
above the mandibular division. It emerges from between the muscles and continues

30 THE CRANIAL NERVES OF NEOCERATODUS,

forward until the posterior wall of the orbit is reached. It then runs round the outer
side of this and below it and then divides into two branches. The smaller, dorsal
braneh breaks up into its terminal twigs in the tissues of the lip, and innervates the
posterior portion of the innominate pouch. The larger, ventral branch runs right
forward and its terminal twigs are distributed to the mucosa of the antero-lateral part
of the palate, the mucosa of the lip and that of the anterior portion of the innominate
pouch. Though it cannot be stated definitely, it is confidently believed that terminal
twigs of both branches of this nerve supply both lateral line and pit organs along the
edge of the upper lip nearly to the extreme forward end.

Nv.ch.t.

1

Nv.my

Fig. 3.—Neoceratodus. Semi-schematic presentation of some of the branches of the Vth and
VilIth nerves. C.hy., Ceratohyoid; Nv. ch-t, Communicating branch of the VIIth to the myloid
branch of the Vth; Nv. my., Myloid nerve; Nv. my.’, Posterior myloid nerve; Temp., Tendon of
the tempo-masseteric muscle

The sixth nerve arises from the ventro-lateral surface of the medulla behind the
roots of the seventh and eighth nerves, by three small roots. The nerve runs forward
below the brain and was unfortunately torn loose so that its course could not be
determined.

The Facialis Nerve.

The seventh nerve arises by three, relatively, widely separated roots, the lateralis, -
combined sensori-motor and otic. The lateralis root arises so high on the side of the
anterior end of the medulla that it may be said to arise from the lateral lip of the
anterior end of the fourth ventricle. The combined sensori-motor root arises directly
ventral to it just above the closely clustered roots of the eighth nerve. These two roots
are of very nearly equal size, the otic root is less than half the thickness of these and
arises in front of the inferior roots of the eighth nerve. The origin of this root is very
close to the labyrinthine roots of that nerve and it passes between the two vestibular
rami on its way to join the general sensori-motor root (Fig. 4). Both the larger roots
split into larger and smaller divisions, and these smaller divisions each pass to the
other root. The otic root joins the general sensori-motor root before that is joined by
the lateralis contribution and after it has given off its own contribution to the lateralis.
In the result the nervus facialis roots have now been collected into two main trunks,
the nervus lateralis dorsally and the facialis proper ventrally.

The fusion of the lateralis ganglion with the gasserian has already been described,
and it was noted that three branches of the lateralis nerve branched from the lateralis
portion of the ganglion.

The ramus ophthalmicus superficialis VII is probably a mixed general and special,
lateralis, cutaneous sensory nerve, and in all probability contains the whole of the fibres
received from the sensori-motor root by the lateralis nerve. This nerve divides into
anterior and posterior divisions almost before it leaves the ganglion. The posterior
division, after a very short course between the muscles of mastication, breaks up at their
surface into four to seven separate strands whieh are distributed to the: subcutaneous

BY H. LEIGHTON KESTEVEN. 51

tissues between the orbit and the anterior boundary of the fleshy wall of the branchial
chamber, but reaching towards the mid-dorsal line and stopping short of the post-orbital
lateral line canal, which runs caudally just above the level of the angle of the mouth.
The main part of the nerve runs forward superficially to the ramus maxillaris trigemini,
gives off several small twigs to the tissues of the skin and lateral line organs immediately
behind and below the eye, and passing, in the subcutaneous tissues, medially to the orbit
terminates in branches which are distributed to the pit and lateral line organs of the
snout, medially to and in front of the location of the olfactory capsules.

Fig. 4.—Neoceratodus. Semi-schematic presentation of the roots of nerves VII and VIII.

The lateralis nerve turns caudally close to the external aperture of the prootic canal
and enters the lateralis canal. This is lodged in that extraordinary expansion of the
otic root of the quadrate which constitutes the cartilaginous roof of the branchial
chamber and which, as Edgeworth (1925) demonstrated, has incorporated into it and
completely obliterated, the diminutive hyomandibular cartilage. The course of the »
canal is curved with the coneavity of the curve medially. It runs caudaily, dorsally and
medially. Whilst within thé canal five twigs are given off. Each of these runs in a
canal through the cartilage to its lateral edge. In this situation the nerve emerges
immediately deep to the lateral line canal and each is distributed to a segment of it.
Just before the lateralis VII reaches the posterior aperture of the canal it is crossed
superficially by a branch of the lateralis X which runs parallel to the branches just
mentioned. Immediately beyond this the two lateralis nerves meet and fuse, the
lateralis X reaching the other through a canal which it enters after leaving the main
vagus trunk. Immediately beyond the point of fusion another laterally trending twig
is given off from the combined nerve.

The subsequent course of the combined lateralis VII-X is caudad, parallel and deep
to the lateral line canal; at Jeast it is so assumed; the nerves were not traced beyond
the commencement of the fleshy wall of the branchial chamber.

The sensori-motor division of the nervus facialis has no communication with the
fifth nerve other than through the inferior mandibular nerve to be described later. Its
root enters a canal in front of the otic capsule and there lies the geniculate ganglion,
completely separated by cartilage from both the gasserian and lateralis ganglia. The
ramus palatinus leaves the ganglion in a ventral direction; lying still within its own
canal the nerve bends forward around the posterior wall of the canal for the internal
cerebral artery and then continues forward enclosed in the cartilage until the posterior
margin of the pterygopalatine bone is reached. The nerve now turns more markedly
laterally and emerges from beneath the bone at the transverse level of the teeth. The
vidian canal is provided with a floor by the pterygopalatine bone, but is lodged in
cartilage above the parasphenoid bone.

32 THE CRANIAL NERVES OF NEOCERATODUS,

The hyomandibular division of the nerve runs laterally and caudally through the
hyomandibular canal between the very much enlarged otic and the basal process of the
quadrate and emerges above the short vertical posterior margin of the quadrate deep
to the antero-dorsal corner of the operculum. Here it divides into several branches.
The first of these penetrates a foramen, in the posterior margin of the otic process, which
lies immediately superficial to the external aperture of the hyomandibular canal. This
is a sensory branch which apparently communicates with the first of the laterally
trending branches of the lateralis nerve wnose canal terminates just anteriorly to this
foramen, but the tissues here are exceedingly tough and permeated by bony spicules,
related to the lateral line canal, which make dissection very difficult. The second
branch is the inferior mandibular nerve already mentioned. This runs ventrad along
the posterior margin of the quadrate for a short distance and then turns forward
through a canal in the quadrate cartilage close to the surface and not far above the
articular head. This canal opens anteriorly about half-way across the quadrate body,
and from here the nerve runs forward on the surface of the cartilage and under cover
of the lower edge of those fibres of the M. quadratomandibularis which here take origin
directly from the cartilage. Turning ventrad just a little further forward, in the tissues
of the cheek just behind the angle of the mouth, the nerve enters the myloid canal as
‘already described.

The third branch of the nerve runs parallel to the inferior mandibular but continues
on in the same direction after that turns forward. ‘This nerve thus comes to reach the
tissues overlying the posterior end of the lower jaw. Here it turns forward and breaks
up into twigs distributed to the subcutaneous tissues behind and superficial to the
posterior part of the Csv..1. muscle. No twigs could be traced to that muscle.

The remaining branches of the hyomandibular division of the facial nerve are the
motor twigs to the Mm. levator operculi, interhyoideus and Cs. 2. They are distributed
over the surface of these muscles and many of their twigs, very definitely, terminate in
the muscles; others of the twigs, however, do not so terminate, and these are probably
cutaneous sensory fibres.

The Auditory Nerve.

The eighth nerve arises by two vestibular roots and four labryinthine. All arise
close together ventral to and just behind the sensori-motor root of the facial nerve, from
the side wall of the medulla close to the inferior margin. The vestibular roots are —
between the labyrinthine and the facial root.

The two vestibular roots pass directly to the vestibule, side by side, and break up
into numerous fine terminal twigs on its medial wall. In similar manner the labyrinthine
roots proceed to the medial wall of the otolith chamber and end in a “fan” of strands
each of which divides again into fine twigs distributed over the wall of the chamber.

The Glossopharyngeal Nerve.

The ninth nerve arises by a single root from the side of the medulla. Its point of
origin is ventral to the vagus roots and a short distance in front of the most anterior
ot them. Bing and Burckhardt (1905) represent the roots of these two nerves and those
of the fifth, seventh and eighth nerves very much more crowded together than has been
the condition in any of the four specimens which have been available for the present
investigation.

No trace of any communication between this and the vagus nerve was seen.

The root is remarkably long, resembling that of Scaphyrhynchus as described by
Norris (1925). It runs caudally and laterally along the dorso-lateral margin of the
vagus roots until it has passed the otic capsule; it then turns laterally and enters a
canal which runs around the posterior and ventro-lateral wall of that cavity and opens
on the ventrum of the skull posteriorly to the basal root of the quadrate. The glosso-
pharyngeal ganglion lies in this canal towards its external aperture. Leaving the
ganglion, the nerve divides into palatine and branchial divisions. The palatine division
runs forward along the lateral margin of the parasphenoid bone. Where the two bones
make contact at this margin the nerve passes on to the ventral surface of the pterygo-

BY H. LEIGHTON KESTEVEN.

(eX)
wo

palatine bone. As it runs forward across this bone it gives off a number of small
twigs and terminates in the thick mucosa just behind the palatine teeth.
The branchial division was not dissected out.

The Vagus Nerve.

The tenth nerve arises by several roots from the side of the medulla dorsally to both
the pyramidal and lateral columns. There are the usual two main divisions of the
roots. The anterior lateralis is a single stout collection of fibres which arises directly
dorsally to the root of the ninth nerve. The posterior division is composed of four
groups of strands, the first of which arises a short distance behind the anterior division
at the same level, and the other three in the same line, each in contact with that in
front of it. Gathered together into one rounded bundle these enter the vagus foramen
and at once enter the vagus ganglion. This ganglion is imperfectly divided into two
portions. The depth of the vagus foramen communicates with two wide canals, that for
the lateralis trunk and that for the remainder of the nerve. The ganglion extends into
both canals and there is quite a marked increase in the girth of the lateralis portion
just within its canal. The lateralis canal turns laterally; it is relatively short and ends
in the posterior end of the lateralis VII canal. Beyond the branches already mentioned
the ramifications and course of the lateralis X were not investigated.

The ramus gastro-intestinalis was only followed as far as the posterior end of the
branchial chamber. The ganglion is very large and extrudes through the posterior
aperture of the canal, and one small and three large branchial nerves arise directly from
the extruded portion. These are spread out slightly so as to have the appearance of
arising from the ventral side of the thickened base of the main nerve.

The Occipital Nerve.

This eleventh nerve is a typical spinal nerve. It arises by single dorsal and ventral
roots which leave the cranial cavity by their own canals, join in a ganglion, which lies
in the cartilage and then divide into dorsal and ventral nerves. The origin of the roots
is in line with those of the rest of the spinal nerves.

REFERENCES TO LITERATURE.

Bine, R., and BuRCKHARDT, R., 1905.—Semon. R. Zool. Forsch... Aust. u. Malay Archipel., Vol. 1,
IP, By

EXDGEWORTH, FEF. H., 1925.—J. Anat., Vol. 59. :

HOLMGREEN. N., and VAN pDER Horst, C. J., 1925.—Acta. Zool., Vol. 6.

IXESTBEVEN, H. LEIGHTON, 1944.—Mem. Aust. Mus., Vol. 8, No. 3, Pt. ii.

NORRIS, Hi. W., 1925.—J. Comp. Neurol., Vol. 39, No. 3.

SAVE-SODERBERGH, G., 1935.—Arkiv fur Zool., Vol. 26.

SMITH, H. B., 1940.—Tvans. Roy. Soc. S. Afr:., Vol. 28.

WATSON, D. M. S., 1928.—Proc. Zool. Soc.. Lond.

WiHSTOLGE. hs s:, 1943 ——Biol. Rer:, Viol. 18, Nie. 2:

CORRELATION OF SOME CARBONIFEROUS SECTIONS IN NEW SOUTH WALBS.
WITH SPECIAL REFERENCE TO CHANGES IN FACIES.

By Aan H. Vorsry, M.Sc., Lecturer in Geology and Geography at New England
University College, Armidale.

(Plates iv—v.)

[Read 18th April, 1945.]

INTRODUCTION.

This paper may be regarded as a sequel to that by Carey and Browne (1938) which
deals with the stratigraphy, tectonics and palaeogeography of the Carboniferous rocks
of New South Wales and Queensland. It takes into account the writer’s investigations
in the North Coast district of New South Wales, particularly those which were made
since the publication of the work referred to above. A short discussion of the important
Carboniferous sections in New South Wales is given and the transition from sequences
which are partly terrestrial and partly marine to the wholly marine sequence of Rock-
hampton, Queensland, is noted (see Plate iv).

The changes in facies as shown by the character of the sediments are then dealt
with and, finally, possible palaeogeographical conditions are considered.

The writer is indebted to the Commonwealth Research Grant to the University of

Sydney for some of the expenses of the field-work.

PREVIOUS LITERATURE.

The Carboniferous Period in eastern Australia was discussed briefly and a compre-
hensive list of related literature was given by Sussmilch (1935). Carey and Browne
(1938) followed this with a review of Carboniferous stratigraphy, tectonics and palaeo-
geography. The writer has referred to Carboniferous rocks on the North Coast of New
South Wales in a series of papers (Voisey, 1934, pp. 336-338; 1936a, pp. 185-189; 1936b,
pp. 157-158; 1938, pp. 454-458; 1939a, pp. 246-247; 1939b, pp. 260-261; 1939c, pp. 387-388;
1939d, pp. 399-401; 1940, pp. 193-209).

In addition to the above, reference is made to the work of others, particularly
Sussmilech and David (1919), Browne (1926) and Osborne (1921, 1922, 1926, 1938) in
the Hunter Valley, and Carey (1937) in the Werrie Basin.

NOMENCLATURE.

The nomenclature adopted is that set out by Carey and Browne (19388, p. 602) except
that the name Kullatine Series is used instead of Upper Kuttung Series for equivalent
beds north of the Manning River Fault System (Voisey, 1939a).

As pointed out by Carey and Browne (1938), the names Kuttung and Burindi have
always referred to terrestrial and marine beds respectively. The marine equivalents
of the Upper Kuttung Series in the Drake area have been called the Emu Creek Series
(Voisey, 19360). The constituent rocks there are very different lithologically from those
in the Macleay, Manning and Hunter districts to the south.

Now, in the Manning district, there are glacial beds corresponding to those of the
terrestrial Upper Kuttung Series in the south but containing marine fossils. Therefore,
in order to avoid violating the principle that Kuttung refers to the terrestrial series
only, it is suggested that the name of Kullatine Series (Voisey, 1934) be retained for
the equivalents of the Upper Kuttung Series in the Manning—Macleay area.

eet eS ae ae ee

a

woe i

BY ALAN H. VOISEY. 30

DISCUSSION OF SECTIONS.
Hunter Valley.

The Hunter Valley section (Plate iv) is based on the work of Osborne (1922). It is
generally regarded as the type-section for the Carboniferous System in New South
Wales.

Osborne’s later work, together with Browne’s mapping of Gosforth (1926), demon-
strates the fact that between Scone and the Coast there is much variation within the
terrestrial Lower and Upper Kuttung sequences.

Werrie Basin.

Carey (1937, pp. 355-359) discusses the generalized section (Plate iv). He notes
the importance of the oolitic limestone as a marker horizon in the Lower Burindi Series
and records the presence of Protocanites lyoni near the base of that series. The Lower
Kuttung beds are compared with those of the Hunter Valley. The presence of the
marine Amygdalophyllum-Lithostrotion limestone horizon in the midst of a terrestrial
sequence is important.

A coarse conglomerate at the base of the Upper Kuttung Series was traced by Carey
for 50 miles. He compares it with the conglomerate at the base of the Glacial Stage
in the Hunter Valley. He divides the series into three stages, viz., the Lower Glacial
Beds, the Interglacial Beds and the Upper Glacial Beds. The Interglacial Stage consists
of Rhacopteris-bearing strata, varves and tillites being developed in the two glacial
stages.

Gloucester.

The Lower Burindi strata occurring on the limbs of the Gloucester Trough to the
north are similar to those near Dungog and Clarencetown.

No big development of conglomerate corresponding to that at Wallarobba has been
seen, so that a lava flow has been taken as the base of the Upper Burindi Series (Voisey,
1940). The lithology of this series is somewhat similar to that of the Lower Kuttung
Series in the south but the sequence appears to be wholly marine and a rich fossil fauna
has been found.

The Upper Kuttung Series consists of conglomerate and mudstones with Rhacopteris
overlain by keratophyres. and rhyolites associated with much pyroclastic material
including some spectacular agglomerates and breccias. The volcanic suite was grouped
by Sussmilch (1921) under the general name of Gloucester Rhyolites but there is more
fragmental material than lava in most sections examined.

The change from terrestrial beds in the southern portion of the Gloucester Trough
(see Osborne, 1938) to marine ones in the northern must take place between Weismantels
and Stratford. It seems from the continuity of the structure that outcrops could be
traced on the limbs of the Trough on each side of this axis though the country is rough
and some of the field-work would be difficult.

The apparent absence of Upper Kuttung glacial beds in the sequences of the
Gloucester Trough is most significant. It should be noted here that the laminated shales
mentioned by Carey and Browne (19388, p. 598) are now believed to be in the Lower
Burindi Series and are not likely to be glacial in origin.

Wingham—Mount George.

The Lower Burindi Series in this district does not seem to include any limestone.
It passes upwards into the Upper Burindi Series and into the Kullatine Series without
much change in lithology until the tillites are met. It has not been possible to divide
what appears to be a continuous marine sequence into portions corresponding with the
three series of the type-area. The marine origin of the tillites is proved by the presence
in them of Spirifer at Kimbriki and Crinoid stems at Killawarra (Voisey, 1939qa).
(Spirifer sp. was found on a geology excursion in June, 1944.)

Taree.
The Lower Burindi Series retains its usual lithological characteristics in the neigh-
bourhood of Taree. As at Barrington, thick beds of massive tuffs occur near the top of

36 CORRELATION OF SOME CARBONIFEROUS SECTIONS IN N.S.W.,

the series, the base of the Upper Burindi being taken immediately above these. The
useful Lithostrotion limestone horizon follows, associated with mudstones and tuffs.

The Kullatine Series is characterized by tuffs and tillites, the latter being found so
low in the sequence that it is apparent that the Rhacopteris-bearing beds of the Main
Clastic Zone are missing or have an insignificant development.

Kendall,

A comparatively small area of Carboniferous rocks is in a block, faulted upwards
into the central region of the Lorne Triassic Basin between Heron’s Creek and Ross Glen
(Voisey, 1939b). A series of tuffs, mudstones, cherts, conglomerates and coal seams is
developed. Rhacopteris ovata occurs in a chert band exposed in a cutting beside the old
road opposite Kew Public School. A conglomerate which may mark the base of the
Main Glacial Beds immediately overlies the horizon.

The occurrence of these terrestrial beds is most important as they do not appear
to be represented at Taree less than thirty miles to the south.

Marine fossils have been found in road cuttings between Kew and Kendall evidently
from beds stratigraphically below the Rhacopteris horizon.

Hastings Valley.

Poor outcrops have militated against the measurement of the sequence in the
Hastings Valley, but essentially the same units as those met in the Yessabah—Wittitrin
area (Voisey, 1936a@) are represented. Typical Burindi sediments are. to be seen in
cuttings along the Oxley Highway on the Wauchope side of the Comboyne Road junction.
Similar beds outcrop in the Cooperabung Mountains to the north. The lower portion of
the Kullatine Series—the equivalent of the Main Clastic Zone is well developed between
Telegraph Point and Rolland’s Plains. The Main Glacial Beds appear at intervals
between the Broken Bago Range and the Hastings River.

Macleay Valley.

The Lower Burindi Series is still characterized by grey and olive-green mudstones
interbedded with a number of different kinds of tuff. Subordinate conglomerates and
breccias occur. The most conspicuous rock is a crinoidal felspathic tuff. This must be
present also in the Upper Burindi Series since it has not been possible to divide the
Boonanghi Series (Voisey, 1934) (= Lower and Upper Burindi probably) into two parts.

The Kullatine Series of the Macleay and Hastings Valleys resembles the Upper
Kuttung Series of the Hunter Valley type-area more closely than do equivalent beds in
any of the intervening districts. It is characterized by a lower portion corresponding to
Osborne’s Main Clastic Zone and comprising tuffs and volcanic breccias with mudstones
and cherts containing fossil wood and Rhacopteris ovata near the top. This is followed
by the Main Glacial Beds with a conglomerate, probably fluvio-glacial at the base. The
remainder of the sequence is made up of tillites and tuffs with subordinate varve-shales.
The tillites are very similar lithologically to those in the Manning district in which
marine fossils have been found, so it seems that they, too, are marine.

It is possible that many of the beds previously grouped to form the Kempsey Series
(Voisey, 1934) belong to the Carboniferous suite but little further information than that
previously recorded has been obtained.

Drake. é

The Hmu Creek Series, which resembles the Neerkol Series of Queensland, is
probably the equivalent of the Upper Kuttung Series as forecast by Osborne (1921)
and Reid (1930) and supported by Whitehouse (Voisey, 1936, p. 163). It is wholly
marine and so far has not been found to include glacial beds. An important horizon is
that containing Productus pustulosus and other forms, some of which are similar to
those in the Upper Burindi Series of Gloucester. Neither the upper nor the lower limit
of the series has been determined.

Rockhampton, Queensland.
In the Rockhampton district of Queensland the entire Carboniferous sequence is
marine and is divided into the Rockhampton and Neerkol Series. The important
Lithostrotion limestone horizon is present near the top of the Rockhampton Series.

BY ALAN H. VOISEY. 37

TRANSITION FROM TERRESTRIAL TO MARINE SEQUENCES.
A study of the Carboniferous rocks of eastern Australia reveals the following facts:

1. The Lower Burindi Series is always marine.

2. The Lower Kuttung Series of the Hunter Valley and Werrie Basin is
terrestrial except for a horizon of marine beds in the Babbinboon area.

3. Between Weismantels and Stratford the terrestrial Lower Kuttung Series
changes to the marine Upper Burindi Series but the lithology is very
similar in both series. ;

4. Between Gloucester and Wingham there has been a change in lithology
from the Kuttung to the Burindi type of sediment in the Upper Burindi
Series.

5. No terrestrial Lower Kuttung rocks are known in the coastal area between
the Manning River and Rockhampton (Queensland).

6. The main Clastic Zone of the Upper Kuttung varies in detail in the
Hunter Valley and the Werrie Basin. It is present in the Gloucester
Trough where it consists of much volcanic material: lavas, tuffs, breccias
and agglomerates. '

7. The Main Clastic Zone does not seem to be represented in the Manning
Valley (excluding the area drained by its tributaries, the Gloucester and
the Barrington). It reappears, however, at Kendall and is well developed
in the Macleay Valley.

8. The Main Glacial Beds of the Upper Kuttung are well developed in the
Hunter Valley and Werrie Basin areas, but are absent from the Gloucester
Trough. In the Manning Valley some of the tillites, at least, are marine
and the same may be true of corresponding beds in the Macleay Valley.

9. Probably the marine Emu Creek Series is synchronous with the Upper
Kuttung Series.

10. In Queensland the change to a wholly marine sequence is complete, the
two series being the Rockhampton and Neerkol.

CHANGES IN FACIES.
Lower Burindi Series.

The rhythmical deposition of the beds of tuff and mudstone throughout the series
is indicative of oscillations in the relations of land and sea during Lower Burindi time.
No marker horizons have been established for the series throughout the region though
some have a local application; e.g., the oolitic limestone at Barrington (Voisey, 1940)
has been useful. Although the series has not been studied in great detail, a few general
observations have some point. Sussmilch (1935, p. 89) drew attention to the fact that
a coarse crinoidal limestone has been found in many places near the base of the series
while oolitic limestones characteristically occur near the top.

Except at Barrington, where conglomerates are present, the basal beds of the Lower
Burindi Series have not been studied in the coastal areas. At Babbinboon, Carey (1937)
discovered basal conglomerates conformably overlying the Barraba Series of the
Devonian. Except for the limestones and occasional conglomerates, the remainder of
the sequence consists of tuffs and grey and olive-green mudstones. The tuffs vary
considerably in their lithology throughout the deposition area. So far no limestones
have been found in the series in coastal areas of New South Wales north of the Manning
River.

The probable position of the Lower Burindi strand-line was indicated on a map
given by Carey and Browne (1938, p. 610) (Plate v, fig. 2A).

The Wallarobba Disturbance (Sussmilch and David, 1919) at the close of Lower
Burindi time raised some parts of the area of marine sedimentation above sea-level.

Upper Burindi (= Lower Kuttung) Series.
As indicated by Carey and Browne (1938, p. 610), it is possible to separate an area
of terrestrial deposition from one of marine deposition during Viséan time.

38 CORRELATION OF SOME CARBONIFEROUS SECTIONS IN N.S.W.,

Further changes in the relations of land and sea can now be indicated as shown on
the three maps (Plate v, figs. 2B, 2C and 2D). The presence of the Lithostrotion lime-
stone at Babbinboon among terrestrial sediments makes it necessary to show these
changes.

The Lower Burindi sequence passes upwards into the Lower Kuttung in the type-
area of the Hunter River—it being difficult to draw a sharp boundary between the two
(Osborne, 1922). The Wallarobba Conglomerate, taken as the basal unit, occurs as far
north as Weistmantels and west of Bullah Delah but is not represented at Barrington
where a lava flow has been taken as the lowest unit (Voisey, 1940). Tuffs and
conglomerates form the basal beds of the Lower Kuttung Series to the west in the
Currabubula district.

The terrestrial origin of these sediments forming the Basal Stage of the (Lower)
Kuttung (Osborne, 1922) is demonstrated by the presence, in tuffs overlying the
conglomerates in the Clarencetown and Currabubula areas, of well-preserved remains
of Lepidodendron, Ulodendron, Stigmaria and Pitys (Osborne, 1922; Carey, 1937).

Towards the sea, fresh-water sediments merge into marine ones, and it is not always
possible to divide the Carboniferous sequence into series corresponding exactly to those
of the type area.

In the Gloucester area a littoral marine facies takes the place of the terrestrial one,
though the sediments are somewhat similar in each case. A shelly fauna including the
large Productid Productus barringtonensis has been found. Beyond this the sediments
change to those of the Lower Burindi type as in the Mount George-Wingham area and
along the coastal strip between the Manning and Karuah Rivers.

At Taree the Lithostrotion limestone was formed while the Gloucester beds seem to
have still remained littoral in character. The seas extended far to the west to reach
Babbinboon and Rocky Creek. Oolitic grits and conglomerates in the first-named locality
indicate proximity to the shore-line there. Terrestrial conditions obtained in the Hunter
Valley all through the Lower Kuttung.

The change back from marine to terrestrial conditions at Babbinboon following the
marine transgression is indicative of an easterly retreat of the sea. Again, neither the
Gloucester nor the Hunter district seems to have been affected.

The areas of terrestrial deposition during the Lower Kuttung are characterized by
volcanic lavas of great variety (Osborne, 1922, etc.; Carey, 1937). Only in the Rocky
Creek and Gloucester areas have lavas been found associated with a marine sequence of
Viséan Age in New South Wales. It would seem, therefore, that the centres of volcanic
activity ran parallel to the strand-line and were more or less restricted to the coastal
strip.

Upper Kuttung.

Another major change in the relations of land and sea took place as a result of the
Drummond Movement at the close of Viséan time. The sea retreated to the north-east
and the strand-line moved to the north of the Macleay River. How far north cannot be
determined as the deposits have been removed by erosion.

Terrestrial sedimentation in early Upper Kuttung time took place apparently in a
number of isolated basins. Some areas, for example, the Mount George—Taree area may
have remained dry land, since no rocks of this age have been found there.

The Main Glacial Beds which follow the Main Clastic Zone do not appear to have
been deposited over much of the area between the Karuah and Manning Rivers, i.e., in
the neighbourhood of the Gloucester Trough. This may well have been high land
formed as a result of great outbursts of volcanic activity and the deposition of the Main
Clastic Zone. It is noteworthy that sediments of the Lower Marine Series (Kamilaroi)
are also absent from this area, unless the Gloucester Coal Measures are their terrestrial ,
equivalents.

The glacial beds of the Hunter Valley are well known. They appear to be terrestrial
in origin, while the particular interest of those north of the type-area mentioned above
is that some of them at least are marine. It is possible that marine conditions prevailed
there through the epoch, indicating a southern advance of the shore-line.

BY ALAN H. VOISEY. 39

In the north-east portion of the State it is probable that marine conditions were
maintained throughout Carboniferous time as indicated by the sediments of the Emu
Creek Series (Voisey, 1936b, pp. 157-158).

CONCLUSION.

Our knowledge of the Carboniferous system in New South Wales is still far from
complete, but sufficient is known to demonstrate the fact that there are represented
beds laid down under a great variety of conditions. It has been possible to indicate
roughly by means of palaeographical maps the principal facies during successive stages
in the formation. These changes have led to difficulties of nomenclature since terrestrial
and marine beds have been laid down synchronously, and the boundaries of the areas of
deposition have been changed from time to time.

Acknowledgements.

I desire to thank Dr. W. R. Browne for his help and advice and Mr. A. H. Robinson
for correcting the manuscript.

EXPLANATION OF PLATE V.

Fig. 2A.—Lower Burindi. The areas occupied by land and sea during Tournaisian times
have been taken from a map by Carey and Browne (1938).

Figure 2A is the key to place names which are indicated only by letters elsewhere. Other
information appears as required and carries on for subsequent maps.

Fig. 2B.—Upper Burindi-Lower Kuttung. The Viséan beds are considered in three parts:
1, those below the Lithostrotion Limestone; 2, the horizon of the Lithostrotion Limestone and its
equivalents; 3, those above the Lithostrotion Limestone.

Figures 2B, 2C and 2D show the distribution of land and sea and the area of terrestrial
deposition in each case.

Fig. 2H.—Upper Kuttung-Main Clastic Zone. Within the dotted area were fresh-water
lakes and thus a regression of the sea is indicated. The absence of sediments in the Taree-
Wingham area suggests that this was dry land.

Fig. 2F.—Upper Kuttung-Main Glacial Zone. The Gloucester-Stroud area was probably an
elevated block with some, at least, of the glacial beds to the north being laid down under marine
conditions.

Fig. 2G.—Carboniferous Lava Flows. Proven Carboniferous lavas are restricted to the
shaded area. They are mostly interbedded with terrestrial sediments.

Fig. 2H.—Kamilaroi-Lower Marine. This map has been included to demonstrate the
relationship between Carboniferous deposition areas and those of the subsequent period.

Note the extension of the land area around Gloucester which was instrumental in preventing
some of the characteristic Queensland fauna from migrating to the Hunter River Province.

The actual glacial beds are not continuous but the marine beds which follow are found
through much of the area indicated.

REFERENCES.

BROWNE, W. R., 1926.—The Geology of the Gosforth District, N.S.W. J. Roy. Soc. N.S.W.,
(HO) 2 Pils
— , and WALKom, A. B., 1911.—The Geology of the HEruptive and Associated Rocks of
Pokolbin, New South Wales. Ibid., 45: 379.
Carny, S. W., 1937.—The Carboniferous Sequence in the Werrie Basin. Proc. LINN. Soc. N.S.W.,
62: 341.
————.,. and BRowNeE, W. R., 1938.—Review of the Carboniferous Stratigraphy, Tectonics and
Palaeogeography of New South Wales and Queensland. J. Roy. Soc. N.S.W., 71: 591.
OSBORNE, G. D., 1921.—A Preliminary Examination of the Late Palaeozoic Folding in the
Hunter River District, New South Wales. Ibid., 55: 124.
———__, 1922.—-The Geology and Petrography of the Clarencetown-Paterson District. Proc.
InN. Soc. N.S.W., 47: 161, 519.
, 1926.—Stratigraphical and Structural Geology of the Carboniferous Rocks in the Mt.
Mirannie and Mt. Dyrring Districts, near Singleton, N.S.W. Ibid., 51: 387.
, 1938.—On Some Major Geological Faults north of Raymond Terrace and their Rela-
tion to the Structure of the Stroud-Gloucester Trough. J. Roy. Soc. N.S.W., 71: 385.
SussmincH, C. A., 1921.—The Geology of the Gloucester District of New South Wales. Ibid.,
By) 2 ABC
, 1935.—The Carboniferous Period in Eastern Australia. Pres. Add., Sect. 6,
A.N.Z.A:A.S., 22: 838.
, and Davi, T. W. E., 1919.—Sequence, Glaciation and Correlation of the Carboniferous
Rocks of the Hunter River District, New South Wales. J. Roy. Soc. N.S.W., 53: 246.

40) CORRELATION OF SOME CARBONIFEROUS SECTIONS IN N.S.W.

Voisgy, A. H., 1934.—A Preliminary Account of the Geology of the Middle North Coast District
of New South Wales. Proc. Linn. Soc. N.S.W., 59: 333.

——— —, 1936a.—The Upper Palaeozoic Rocks around Yessabah, near Kempsey, New South
Wales: Je Roy: Soc) NeS3W., 7102 183°

——., 1936b.—The Upper Palaeozoic Rocks in the Neighbourhood of Boorook and Drake,
Nis: Wi. Proc LINN. Soc NESiW., 61): 255.

, 1938.—The Upper Palaeozoic Rocks in the Neighbourhood of Taree, N.S.W. Ibid.,
63: 453.

————, 1939a.—The Upper Palaeozoic Rocks between Mount George and Wingham, N. S.
Wales. Ibid., 64: 242.

, 1939b.—The Lorne Triassic Basin and Associated Rocks. Ibid., 64: 255.

————, 1939c.—The Geology of the County of Buller, N.S.W. Ibid., 64: 385.

, 1939d.—The Geology of the Lower Manning District of New South Wales. Ibid.,
64: 394.

————, 1940.—The Upper Palaeozoic Rocks in the Country between the Manning and Karuah
Rivers, New South Wales. Ibid., 65: 192.

41

EVIDENCE OF AN EUSTATIC STRAND-LINE MOVEMENT OF 100 TO 150 FEET
ON THE COAST OF NEW SOUTH WALES.

By Winson H. Mazer, M.Sc., Lecturer in Geography, University of Sydney.
(Four Text-figures and one Map.)

[Read 28th March, 1945. ]
‘Evidence of eustatic strand-line movements have been established in many parts of
the world. In New South Wales, a level about 20 feet above the present sea-level which
may have an eustatic origin has been noted by many writers—David (1907), Hedley
(1924), Maze (1933), Voisey (1934), and Jutson (1939), etc. Only a few records—

BROKEN
BAY

\'sYDNEY C)
PARRAMATTA®
ele

MILES
ie} 10 20

Map 1.—The central coastal area of New South Wales.

The area for which the altimetric frequency curve in

Fig. 1 was constructed is enclosed by the dashed line.

The altimetric frequency curves in Figs. 2 and 3 are
for the areas marked A and B respectively.

42 AN EUSTATIC STRAND-LINE MOVEMENT ON THE COAST OF N.S.W.,

Browne (1926), Voisey (1934) and Sussmilch (1940)—have been made of silt terraces
and gravels along the coastal rivers at elevations of 100 to 150 feet higher than the
present river levels. Field observations and analysis of topographic maps reveal that
this higher level has a wide distribution.

EVIDENCE FROM ALTIMETRIC FREQUENCY CURVES.

The introduction and use of altimetric frequency curves for land-form analysis has
already been discussed (Maze, 1944). It consists of the statistical analysis of spot
heights or the heights deduced from the contours of the highest points in uniform
squares on topographic maps.

The one inch to a mile topographic maps for the central coastal area of New South
Wales, as defined in Map 1, have been examined and the values of the heights in each
of the 1,000 yards grid squares have been listed and arranged so as to give the frequency
of occurrence of each height. From these frequencies a frequency curve has been
constructed (Fig. 1) to show the altitudinal distribution of the “high points’ over the

1000

—{800
700

600
4 BE
ee

TE :

FREQUENCY OF OCCURRENCE .

a ;
100

Sun ovaS
Oo wo

v ) a 7
ALTITUDE IN FEET.

Fig. 1.—Altimetric frequency curve for the central
coastal area of New South Wales.

700

BY WILSON H. MAZE. 43

area. On an altimetric frequency curve a well-marked frequency maximum indicates
the presence of a fairly level bench or platform. In Fig. 1, a maximum occurs for the
elevations between 0 and 50 feet. This indicates the widespread areas, mainly consisting
of depositions which have been partly exposed by the recent small negative movement of
sea-level. [See David (1907) and Maze (1933), etc.] The frequency curve drops for
elevations between 50 and 100 feet and then rises abruptly for elevations between 100
and 150 feet. There is thus over the whole central coastal area of New South Wales
a greater frequency of elevations between 100 and 150 feet than any other elevation
except those between 0 and 50 feet.

The altimetric frequency evidence is even more striking when smaller areas are
considered in more detail. Fig. 2 is an altimetric frequency curve for area A as shown
in Map 1. This area extends along the southern side of the Hunter River from Newcastle

120

100

80

60

40

FREQUENCY OF OCCURRE NCE

20

(o)

fo) ° fo) fe) fo) fo) fo) ° ro) ra) rs) ° °

fo) ro) ,o) fo) fo) to) ts) fe) re) 3} ro) fo) ° fo)

z 32) N = ° o ro) 5S re) o zt ~* a =
(Ew wCHOEm WN) ists a

Fig. 2.—Altimetric frequency curve for the Newcastle
district, or area marked A on Map 1.

to Hast Maitland and south to Lake Macquarie. All the low-lying depositional land-
forms (less than 25 feet above sea-level) such as swamplands and recent sand deposits
have been omitted in the analysis of the heights, so that Fig. 2 is an altimetric frequency
curve of an erosional surface. In it the maximum frequency for elevations between
100 and 150 feet is even more apparent. This is amply borne out by field-survey, which
reveals that gently sloping spurs and benches between these elevations are a charac-
teristic feature of the area.

Fig. 3 is an altimetric frequency curve for area B, or the Sydney district, as shown
on Map 1. This curve also shows the frequencies rising to a maximum for the elevations
between 100 and 150 feet.

The altimetric frequency curves thus point to the widespread existence of a bench
along the coast at an elevation between 100 and 150 feet.

EVIDENCE FROM VALLEY-IN-VALLEY AND LONGITUDINAL RIVER PROFILES.
Evidence of valley-in-valley forms is common in the Sydney district, particularly in
the tributaries of Middle Harbour. Fig. 4, which was constructed from field-survey
observations, illustrates the longitudinal profile and two cross-sections of Flat Rock

44 AN EUSTATIC STRAND-LINE MOVEMENT ON THE COAST OF N.S.W.,

Creek, a tributary of Middie Harbour. The longitudinal profiles of the main stream
and its tributary both have well-marked knickpoints, and the cross-sections show well-
defined valley-in-valley forms. Many. pitfalls and assumptions attend any attempts at
extrapolation and reconstruction of cross-sections [see Johnson (1938) and Miller
(1939) ]. In the case of Flat Rock Creek the extrapolation and reconstruction required
are only for a relatively short distance and a valley-in-valley section is available for the

20

o
)

uJ

VU

Z

uJ

(od

a

5)

TS)

TS)

2 100

(he

e)

>

©)

Zz

lu

=)

3a 50

WW

a

(i
ie)
je) fo) fe) fo) fo) (2) fo)
fo} ie) ro) (e) fo) e)
o o (09) fat) =

T
NEWS TN PEE a

Fig. 3.—Altimetric frequency curve for the Sydney
district, or area marked B on Map 1.

lower reaches of the Creek (Section B). If the reconstructed valley-floor heights of 160
feet from Section A and 140 feet from Section B are used, then the dashed line in
Section C is an attempt to reconstruct the pre-rejuvenation profile of the stream. This
would suggest a former base-level or sea-level at an elevation of 100 to 150 feet higher
than at present.

The necessity for detailed field-survey to obtain the precise data for stream profile

reconstruction makes it impossible, for the time being, to attempt this work on a large |

scale. Aneroid observations and the use of the 1 inch to 1 mile topographic map reveal
that Moore’s Creek, another tributary of Middle Harbour, has a well-defined knickpoint
and valley-in-valley forms of the same order as Flat Rock Creek. Similarly on the
eastern side of Lane Cove River, Sydney, in the vicinity of Bradfield, there is an extensive
bench in the valley side about 150 feet above sea-level.

Observations to date do not reveal similar knickpoints in many of the coastal streams
of New South Wales, but in the Hunter Valley, valley-in-valley forms are confirmed by
the wide distribution of terrace and river gravels from Newcastle up to Scone. At Hast
Maitland there is a marked bench (Sussmilch, 1940) covered with river sands and gravels
about 125 feet above the present river level. Browne (1926) has also described relics
of former flood-plains, which are found up to a maximum elevation of 120 feet above
the river level at Gosforth. Similar river gravels occur on terraces above the Hunter
River near Muswellbrook and west of Kingdon Ponds near the town of Scone.

On the Macleay River, Voisey (1934) has also described gravels, 100 feet and more
above the present river level.

CONCLUSION.

Evidence from field observations and from a large-scale statistical analysis of the
topographic maps indicates a wide distribution of a level of deposition and erosion along
the central coastal area of New South Wales at a level between 100 and 150 feet above
sea-level. The existence of such a surface can best be explained by postulating a period
of prolonged stability of sea-level corresponding to a maximum of transgression at about
150 feet. -It would also appear fromthe evidence of the valley-in-valley forms that the

.

BY WILSON H. MAZE. 45

time which has elapsed since the withdrawal of the sea and subsequent rejuvenation of
the valleys has been small compared with that taken for the earlier broad valleys to be
graded to the higher base level.

4 MILES

O SECTION B
PROFILE

2@ MILES

-

O SECTION A
8

ee hee
A
Fig. 4.—Cross-sections (A) and (B) and longitudinal stream profile (C) for Flat Rock Creek, Middle Harbour, Sydney.

9 secTION Cc

46 AN EUSTATIC STRAND-LINE MOVEMENT ON THE COAST OF N.S.W.

Further detailed work is required to see if a range of terraces exists such as are
described for the Mediterranean and Atlantic coasts. The 100- to 150-feet level for New
South Wales may be correlated with the Tyrrhenian (30-35 metres) terrace of the
Mediterranean, the 100-feet beach of Britain or the 30-metre terrace of eastern North
America. The latter terraces have been attributed to Pleistocene strand-lines formed by
oscillations of the sea-level during the inter-glacial stages.

REFERENCES.

BROWNE, W. R., 1926.—The Geology of the Gosforth District, N.S.W. Part i. J. Roy. Soe.
N.S.W., 60: 248 and 272.

Davin, T. W. E., 1907.—Geology of the Hunter River Coal Measures. Mem. Geol. Surv. N.S.W.,
Geology No. 4.

HEDLEY, C., 1924.—Differential Elevation near Sydney. J. Roy. Soc. N.S.W., 58: 61-66.

JOHNSON. D., 1938.—Stream Profiles as Evidence of Eustatic Changes of Sea Level. J. Geo-
morphology, 1: 178-181.

JuTSON, J. T., 1939.—Shore Platforms near Sydney, New South Wales. Ibid., 2: 237-250.

Maze, W. H., 1933.—Pot holes at Green Point, Broken Bay, N.S.W., and their Significance in
Coastal Uplift. Awst. Geographer, 2: 52-55.

, 1944.—The Geomorphology of the Central Eastern Area of New South Wales. Part i.

J. Roy. Soc. N.S.W., 78: 30 and 31.

MIuugrR, A. A., 1939.—Attainable Standards of Accuracy in Determination of Preglacial Sea
Levels by Physiographic Methods. J. Geomorphology, 2: 95.

SussmMILcH, C. A., 1940.—The Geomorphology of the Hunter River District, New South Wales.
Proc. LINN. Soc. N.S.W., 65: 317.

Voisny, A. H., 1984.—The Physiography of the Middle North Coast District of New South Wales,
J. Roy. Soc. N.S.W., 68: 95-96.

47

ON AUSTRALIAN DERMESTIDAER. PART IV.
NOTES AND THE DESCRIPTION OF A NEW GENUS AND FOUR NEW SPECIES.
By J. W. T. ARMSTRONG.
(Three Text-figures. )

[Read 28th March, 1945.]

The following tabulation of those genera of the family Dermestidae represented in
Australia is largely based on those given by Leconte and Horne (Coleoptera of N.
America, p. 142) and Blackburn (Trans. Roy. Soc. S. Aust., xxvii, 1908, p. 159).
Megatoma is not included, as the only Australian species still placed in it is
M. tenuifasciata Reitt., which probably does not belong there (see my foot-note, these
‘PROCEEDINGS, Ixvii, 1942, p. 328).

Tabulation of Genera of Australian Dermestidae.
eM LOntalOcelluSvabsSent os fa sc cyef ccciols ce pele erecelee 4 vlelel ore. ete sleiglels Piet nice eters RO RATE Dermestes
AA. Frontal ocellus present.
B, Mesosternum narrow, middle coxae not widely separated, antennal fossae wanting ......
ROMAN cer oh yhs tate cle Vers ce masti cite ieee a alee asa En UR UAT sire ice Seen os Getto naSNS aa ocetiedal ub ole ee dal atlewscavleta wt Attagenus

Subgen. Telopes
BB. Mesosternum broad, divided or emarginate, receiving the tip of the prosternum, middle

coxae widely separated, antennal fossae present, but very wide and indefinite in

Psacus.
C. Femora and tibiae strongly compressed.
Deetleadinormailssantennale fossae) INCONSPICUOUS <i as ce: ees © cles eles oi piers Adelaidea

DD. Head abnormal, produced forward and upward above eyes, antennal fossae distinct

PR ee Ne Hee Cereals pelt trans soul es saearciaitmtsny ara israel nvetehe: Seavey de af aus Ghenacs Myrmeanthrenus, n. gen,
CC. Tibiae at least normal.

D. Vestiture consisting of hairs.
EK. Antennal fossae lateral, sulciform.
PATE TIVa Cratl ab Cll teins sacesacdercucusssed eibeawlatews core codeine re Gr elastic aucuad oleitelinn cies) alauenes usiieanie Psacus

EE. Antennal fossae anterior, foveiform.
IN. ANaKSroa NL Glo) pe Wav TOMI Locococnecndobuoopos bu ourboobneodas< Anthrenocerus
FF. Antennal club of two very unequal joints, the last circular .......... Orphinus
HMhMaAntennal club) of one joint. very largeriny Gio. .5. ssn eee on. Thaumaglessa

DD. Vestiture consisting of scales.

EH. Form rotund, head scarcely visible from above ............. Bocce Anthrenus
EE. Form narrower, pronotum anteriorly raised, head plainly visible from above
RR STOR RCM TS hee TSR Lee ene TIC cen ee UCR eee Loiah, oc HE Ree GL Gee Neoanthrenus

Genus DERMeESTES L.*

Those species of Dermestes that have been introduced to Australia, may be distin-
guished among themselves by the following tabulation.

PAM © OL OUMAUTNT ROTI a Ne meso e ete iioue ces ei oine al eilsliailen ctisttel osiselstre IMUM NaS ca tie eke tier ecec eho eres) Gaetoe! alia tae 6 cadaverinus F.

= felinus F.
AA. With a pattern of lighter coloured pubescence.
eee CLO CHIN SeaO fre ly EA MUTT O RIG .s!).0 si cnone sorskal oer ec oe) el cies eke phcusaeileps caueusire yerttep were sue one seer sa anebs vulpinus F.
= australis Macl.
BB. Clothing of elytra bicolorous.
C, Elytra with a broad sub-basal band of ashen pubescence enclosing six dark spots,

clouhinewotLeaventraliesurtacemcdankwenessiei eden me cuacihorielsuclaclssclickucnedsts lardarius L.
CC. Elytra with numerous small spots or speckles distributed evenly thereon, ventral surface
thickivaclothedmwithvwinitishs DUbeSCeNnCe) misc clesacelce soko siee sisi murinus F.

* Since I submitted this paper for publication, Mr. H. S. Barber of the United States National
Museum has sent me his note, “Some synonymy in Dermestes (Coleoptera)’’, Bull. Brooklyn
Ent. Soc., xxxvii (5), 1942, pp. 174-176, in which he points out that Dermestes ater De Geer
1774 has priority over D. cadaverinus F. 1775 and D. maculatus De Geer 1774 over D. vulpinus
F.1775 and that there is no good reason known why De Geer’s names should not stand. Froggatt
in his book ‘‘Australian Insects’’ has the names D. cadaverinus F. and D. vulpinus F. transposed,
which explains some of the confusion in Australian collections.

48 ON AUSTRALIAN DERMESTIDAE. Iv,

D. murinus F. is listed in Masters’ Catalogue from Queensland, but I have not seen
an Australian specimen nor any other reference to it.

Genus ATTAGENUS Latr.

There are four species of Attagenus represented in Australian collections, of which
the only one that may be endemic is A. (Velopes) undulatus Motsch. Brachysphyrus
iwroratus Blackb. is a synonym of this species (Arrow, Ann. Mag. Nat. Hist., (8) xv,
1915, p. 426). I have seen numerous specimens of an insect that agrees with Castelnau’s
brief description of his A. annulifer (Hist. Nat. Col., ii, 1840, p. 36), which is placed
as a synonym of A. cinnamomeus Roth. in the Junk Catalogue. A. gloriosae F.* is
placed under the same name.

‘The four. species-may be distinguished as follows:

A. Head, ” pronotum and antemedial elytral fascia clothed with golden yellow pubescence ......

PE A Puts pha Staats: cy ei gs Sp avast Oa as BA ee he ee oc EO SN Be SOTO S CREE
AA. Not thus.

B. With 5 small conspicuous white spots, 3 on pronotum situated near each posterior angle
and on ‘the medial lobe, 2 on elytra placed medially on either side of suture .......

Saba SE a go. RUorahs cays tier aihe, ISRTNE Mi ecPapeetctmeiet aeiclons (gt ners nitions RU Gr eR Oa pe gE TR CR pellio L.
BB. Not thus.

Cc. More ovate, numerous small White pubescent spots on dorsal surface

a djatabaten Soke SGeholiguemencehetese choliegchat sh eueli ch al stent pellet neem aulcuermactnc’ aha neuen RLS CUaEa ISL MTL IUCL OU L(LDILS Sev LOUSG Ine

CCe Moretelongatesclothin a sumitonmnser scien cei iclerd eacieicien eel ee eemerene sens (ite piceus Ol.

Gene

Genus Psacus Pasc..

ne ‘genus, originally placed in the Rhipidoceridae, can only be separated from
Ty ogoder ma by the great development of the male antennae. P. mastersi Macl. is
evidently not a Dermestid as Mr. K. C. McKeown, of the Australian Museum, in a letter,
states: “I have made a careful examination of the type of P. mastersi Macl. There is
no sign of.an ocellus as far as I can see and the antennae are strongly flabellate over
almost their whole length; superficially the appearance is very like that of a Trogoderma,
put it is much narrower and the base of the prothorax is almost straight, not roundly
excavated on either side, as in the Dermestids.” This leaves P. attagenoides Pasc. as the
only described species. A second is now added.

PEN Te PSACUS CALLUBRIENSIS, N. Sp.
mlonenee! ovate, fusco-piceous, nitid, rather strongly and closely punctate, clothed
with moderately long semi-erect fusco-piceous setae, legs and antennae fusco-rufus.

: Pronotum widest at base, this moderately bisinuate, sides evenly rounded to apex,
not noticeably expanded at middle, posterior angles acute. Elytra slightly wider than
prothorax, two-thirds as wide as long, as wide as pronotum at base, thence expanding a
little to shoulders, apex evenly rounded. Antennae noticeably hairy, ¢ segments 1 and 2
moniliform, 3 very small, 4 strongly pectinate, 5 to 10 strongly flabellate, 11 elongate-
clavate; 9 segments 1 and 2 moniliform, 3 to 6 small, 7 to 10 larger serrate, 11 subovate,
rather pointed at apex. No defined prosternal fossae, but antennae received under lateral
margin of prothorax much as in Attagenus.

~ Size:'2-8 mm. x 1:3 mm.; 2 mm. x 1 mm.

~ Hab—N.S.W.:. Bogan R., Nandewar Range (J. W. T. Armstrong).

_ _ Cotypes in the British, South Australian and Australian Museums and the author’s
collection.

It is with hesitation that I place this species with P. attagenoides Pasec. The
prothorax is not greatly expanded to take the antennae of the male, but otherwise it
agrees fairly well. There are fifteen examples before me, and they are at once
distinguished from Pascoe’s species by being much smaller and far less hairy. They are
to be found on the flowers of the wilga’(Geijera parviflora), mostly in company with

* Since submitting this paper for publication I forwarded a specimen of the species referred
to above as Attagenus ? gloriosae F. to Mr. G. J. Arrow of the British Museum of Natural
History for verification. In a letter just received he writes: ‘“‘Rather to my surprise, I have
found that the unknown Attagenus is not A. gloriosae but a species unknown to me.’’ In this
case A. gloriosae has not been recorded from Australia, and the species here referred to remains
to be determined.

BY J. W. T. ARMSTRONG. 49

Trogoderma boganense Armst., with which they are easily confused, owing to their small
size and dark colour, until examined with a glass.

Genus TrogopERMA Berth.
TROGODERMA PICINUM, Nl. Sp.

Elongate-ovate, piceous, nitid, clothed with short semi-depressed piceous setae,
antennae and legs fuscous.

Pronotum widest at base, this moderately bisinuate, posterior angles acute, sides at
first gradually then evenly rounded towards apex, rather finely and closely punctate.
Elytra two-thirds as wide as long, as wide as prothorax at base, slightly expanding to
shoulders then subparallel for half length, then evenly rounded to apex, more coarsely
and closely punctate than pronotum. Antennal club ovate, 3-segmented. Prosternal
fossae wide, triangular, closed.

Size: 1:9 mm. x 1:1 mm.; 1:75 mm. x 0-9 mm.

Hab.—N.S.W.: Mullaley (J. W. T. Armstrong).

Cotypes in British and South Australian museums and the author’s collection.

This species would be associated with 7. parvum Armst., excul Blackb. and lindense
Blackb. in my tabulation (these Procrerepines, Ixvii, 1942, p. 321). It is narrower than
any of these and the 3-segmented antennal club at once separates it from 7. parvum and
lindense. T. excul is lighter in colour and has very different clothing. There are five
specimens before me.

TROGODERMA NIGRONITIDUM, Nl. Sp.

Ovate, convex, black, nitid, clothed with short black semi-erect setae, tarsi fusco-
piceous.

Pronotum widest at base, medial lobe wide, deep and narrowly rounded at apex,
posterior angles acute, sides evenly rounded to apex, disc deeply and fairly closely
punctate, more closely so at sides. Elytra three-fourths as wide as long, base as wide
as prothorax, expanding in same line as sides of pronotum to shoulders, these evenly
rounded, thence narrowing in a gentle curve to near apex which is evenly rounded,

_ coarsely and closely punctate. Antennal club wide, compact, elongate-ovate,

5-segmented, 2 3-segmented. Prosternal sulci wide, moderately deep, triangular, closed.

Size: 3°3 mm. x 2 mm.; 3 mm. x 1:8 mm.

Hab.—N. Qd.: Townsville (G. F. Hill), on grass flowers, Cooktown (W.J.T.).

Cotypes in the National Museum and F. EH. Wilson’s and the author’s collections.

Four specimens from northern Queensland represent a species that falls beside
T. morio Er., in my tabulation referred to above, from which it is distinguished by its
greater convexity, narrowing elytra, and its antennal club being much wider and more
compact.

, Genus MYRMEANTHRENUS, nN. gen.

Body compact, finely setese. Femora and tibiae strongly compressed, tarsi slender.
Head abnormal, produced widely and anteriorly elevated, with a large cavity between
the eyes, at the tip of which the ocellus is situated; eyes visible from above. Prosternum
produced anteriorly concealing mouth parts, with large foveate antennal fossae situated
along the anterior margins. Mesosternum narrow, entirely bisected. Antennae short,
11-segmented, segments 1 and 2 moniliform, 3 to 8 short, strongly compressed, 9 to 11
forming a stout ovate club.

Genotype, Myrmeanthrenus frontalis, n. sp.

The abnormal head at once separates this genus from all others known to me. It is
further distinguished from Anthrenocerus, to which it seems closest, by its compressed
legs, from Trogoderma by the position of the antennal fossae and from Neoanthrenus
and Anthrenus by the setose clothing. The single species described below was found
associated with ants which would account for the compressed appendages and frontal
fovea.

MYRMEANTHRENUS FRONTALIS, D1. Sp. Figs. 1-3.

Ovate, brunneus, nitid, sparsely clothed with long fine brunneus setae, antennae and

legs castaneous.

I

50 ON AUSTRALIAN DERMESTIDAE. Ivy,

Head produced anteriorly above eyes and strongly elevated at apex, underside of
this projection with an extensive deep fovea continuing down between eyes, having the
margins thickly rounded. Pronotum transverse, widest at base, convex, sides evenly
rounded to apex, base moderately bisinuate, posterior angles acute, base and sides
slightly marginate, sparsely and lightly punctate. Elytra three-fourths as wide as long,
base slightly narrower than prothorax, widening to shoulders thence almost parallel till
apical one-third, this evenly rounded, coarsely and fairly closely punctate with two
perceptible striae on either side of suture.

(e

Figs. 1-3.—Myrmeanthrenus frontalis, n. sp. 1. Lateral view. 2. Dorsal view of head
and prothorax. 3. Head from below and in front showing frontal cavity.

Size: 2°75 mm. x 1:5 mm.

Hab.—Vict.: Tallangatta (Hjnar Fischer), in ants’ nests under stones.

Holotype and allotype in the authoyr’s collection.

There are two specimens of this remarkable inquiline before me, given me by
Mr. F. E. Wilson, whom I have to thank for the opportunity of describing them. They
seem to be sexes as one has the antennal club less regularly ovate and with the
segments more uniform in length. They represent the most abnormal species of this
family so far described from Australia. Figure 3 is to some extent a sketch, from in
front and below the head, and must not be considered accurate in detail.

Genus ANTHRENUS Geoffroy.
ANTHRENUS VORAX Waterh.
In November, 1943, Mr. F. H. Taylor sent me for identification five specimens of
A. vorax Waterh. taken in an Australian port on an Australian warship, which had
presumably been in Indian waters. They were infesting horse-hair fillings of chairs,
etc., and there would seem to be considerable risk that the species might be introduced
to this country. It is more robust and more brightly coloured than A. verbasci L., and
is more circular in outline. The clothing of the ventral surface is snowy white.

ANTHRENUS SCROPHULARIAE L.

There is a reference to this species in the Report of the Entomologist and Vegetable
Pathologist (A.R. Dept. Agric. and Stock, Qd., 1918-19, p. 40), as follows: “Beetle larva
(Anthrenus scrophulariae) attacking woollen clothes; two instances, Brisbane.” This,
in the absence of any further evidence, would seem to be a mis-determination and I do
not propose to admit the presence of this species in Australia. This report has been
cited by Griswold and Greenwald (Corneil Univ. Agric. Exp. Stat., Mem. 240, pp. 59 and
74), who also give a sufficiently alarming account of the ravages of both this species
and A. vorax. A. scrophulariae can be easily distinguished from the species already
introduced, by the orange-red vitta along the elytral suture.

ConcLupDING NOTES.
As this paper completes a series based on the material available to me up to the
present, the following check-list is given as a summary. Ninety-one species are listed,

|
|

BY J. W. T. ARMSTRONG. 51

of which two (Dermestes murinus F. and Thaumaglossa concavifrons Reitt.) are doubt-
fully recorded as Australian. This is an increase of just over 50 per cent. on the number
(60) given by Tillyard (Insects of Australia and New Zealand, 1926). Those species
marked with an asterisk occur on the Bogan River above Nyngan, in central New South
Wales, and the majority of these, with the exception of the two species of Dermestes, are
to be found on the flowers of the wilga (Geijera parviflora) during the spring and early
summer. On several occasions I have taken specimens of Trogoderma tolarnense Blackb.
infesting my insect collection and also in tins containing a little powdered milk.
Attagenus ? gloriosae F.+ has only made its appearance in the Bogan River district in
recent years and is rapidly increasing.

The numbers in brackets refer to the list of references and indicate the paper in
which the original description is to be found.

List of Australian Dermestidae including Introduced Species.

Dermestes cadaverinus F.* (Introduced. ) picinum, nN. sp.
= felinws BH. (19) reitteri Blackb. (6)
lardarius lL. (Introduced. ) riguum Hr. (10)
murinus EF. (Introduced, if, in fact, present rufipenne Armst.* (2)
in Australia. ) ‘ setulosum Armst. (2)
vulpinus F.* (Introduced.) singulare Blackb. (5)
= australis Macl. (14) socium Lea (12)
Attagenus ? gloriosae F.*7 tasmanica Armst. (2)
pellio L. ie ; tolarnense Blackb.* (8)
piceus Ol.* varipes Blackb. (6)
undulata Motsch. : whitei Armst. (2)
= Brachysphyrus irroratus Blackb. (8) yorkense Blackb. (5)
'Megatoma tenuifasciata Reitt. (18) Psacus attagenoides Pasc. (15)
(Unknown to me.) callubriensis, n. sp.*
Thaumaglossa concavifrons Reitt. (18) Adelaidea rigua Blackb. (5) (Unknown to

(Doubtfully recorded from Tasmania. me. )

Unknown to me.) Myrmeanthrenus frontalis, n. gen. and sp.
nigricans Macl. (14) Anthrenocerus australis Hope (11)
= Orphinus rufopygus Pic = erichsoni Reitt. (18)

Trogoderma adelaidae Blackb. (5) bicolor Arrow* (4)
alpicola Blackb. (5) ‘ blackburni Armst. (3)
antipodum Blackb. (5) chalceows Armst. (3)
apicalis Macl.* (14) concolorous Armst. (3)
= pectinifer Arrow (4) condensus Armst. (3)
apicipenne Reitt. (17) confertum Reitt.* (18)
= baldiense Blackb. (5) = flindersi Blackb. (5)
blackburni Lea (13) convexus Armst. (3)
boganense Armst.* (2) maculosus Armst. (3)
carteri Armst.* (2) niger Armst.* (3)
consors Arrow (4) pulchellus Arrow (4)
debilius Blackb. (8) quadrifasciatum Blackb. (8)
difficile Blackb. (5) signatus Armst. (3)
ellipticum Armst. (2) terzonatum Blackb.* (8)
excul Blackb.* (8) trimaculatus Armst.* (3)
explanaticolle Armst. (2) variabile Reitt.* (18)
eyrense Blackb.* (5) Anthrenus verbasci L.* (Introduced. )
frater Arrow (4) = varius EF.
froggatti Blackb. (6) pimpinellae F. (Introduced)
hobartense Armst. (2) Neoanthrenus frater Arrow (4) =
inconspicuum Armst. (2) niveosparsa Armst. (1)
laevipenne Armst. (2) ocellifer Blackb.* (5)
leai Armst. (2) : parallelus Armst. (1)
lindense Blackb. (5) Orphinus atrowus Armst. (3)
longius Blackb. (8) australicum Blackb. (5)
macleayi Blackb. (5) caswuarinae Blackb. (8)
marginicolle Armst. (2) | ceciliense Blackb. (8)
maurulum Blackb. (8) interioris Blackb. (5)
meyricki Blackb. (5) minimus Arrow (4)
morio Er. (10) ; nealense Blackb.* (8)
nigrobrunneum Armst. (2) occidentalis Armst. (3)
nigronitidum, n. sp. quornense Blackb. (7)
occidentale Blackb.* (5) woodvillense Blackb.* (5)
parvum Armst. (2) = eucalypti Blackb. (8)

+ See footnote, p. 48.

52 ON AUSTRALIAN DERMESTIDAE. IV,

REFERENCES. *

(1). ArRmstRONG, J. W. T., 1941.—PrRoc. LINN. Soc. N.S.W., 66: 388-390.
(2): , 1942.—TIbid., 67: 321-330.

(3). ————, 1943.—Ibid., 68: 57-63.

(4). ArRRow, G. J., 1915.—Ann. Mag. Nat. Hist., (8) 15: 425-451.

(5). BLACKBURN, T., 1891.—Trans. Roy. Soc. S. Aust., 14: 123-133.

(6). , 1892.—Ibid., 15: 34 and 207-208.

(7))c , 1894.—Proc. LINN. Soc. N.S.W., (2) 9: 93.
(8): , 1903.—Trans. Roy. Soc. 8S. Aust., 27: 159-173.
(9). ————, 1907.—Ibid., 31: 232.

(10). BrRicHson, W. F., 1842.—Arch. f. Naturg., 8 (1): 152.
(11). Hops, F. W., 1843.—Ann. Mag. Nat. Hist., xi, 1843, “Proc. Ent. Soc. Lond. for June, 1842”,

10s asdb)>
(12). Lea, A. M., 1895.—Proc. Linn. Soc. N.S.W., (2) 10: 228.
(13). ———, 1908.—Proc. Roy. Soc. Vict., 20 (2) (New Series) : 155.
(14): Macipay, W. J., 1871-—Trans. Ent. Soc. N.S.W., 2 (3))3 170-174.

(15). Pascon, FE. P., 1866.—J. Hnt., 2: 446.

(16). Pic, M., 1933.—HAnt. Nachr. Bl., 7: 71.

(17). Rerrrer, H., 1881.—Deuts. ent. Z., 25: 232.

(18). , 1881.—Verh. naturf. Ver. Briinn., 19 (1880): 36, 42-438 and 55-56.
(19). -Fapricius, J. C., 1787.—Mant. Ins., 1: 34.

* References to introduced species are not included.

CORRIGENDA. .
Armstrong, 1942 (Part ii of this series), page 322, line 2 from bottom: for less read more.

, 1943 (Part ili of this series), page 57, line 17 from bottom: after ‘clothing bicolorous”
insert ‘‘or of more than two colours”.

Proc. Linn. Soc. N.S.W., 1945.

PLATE

Morphology of the Pouch Young of Thylacinus cyinocephalus.

Ie

Proc. Linn. Soc. N.S.W., 1945. PrArE air

Stenoporids from the Permian of N.S.W. and Tasmania.

Proc. Linn. Soc. N.S.W., 1945. PLATE III.

Stenoporids from the Permian of N.S.W. and Tasmania.

Lunn. Soc. N.S.W., 1945.

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RELATION OF THE ORCHID FLORA OF AUSTRALIA TO THAT OF NEW ZEALAND.
WITH THE DESCRIPTION OF A NEW MONOTYPIC GENUS FOR NEW ZEALAND.
By H. M. R Rupp, B.A., Sydney, N.S.W., and EH. D. Hatrcu, Auckland, N.Z.

[Read 27th June, 1945.]

This paper may be conveniently divided into the following sections:
I. A general survey of the orchid genera of both countries, with certain data

in regard to their distribution.

II. The probable origin of these genera.

III. Orchid species common to Australia and New Zealand.

IV. Possible explanations of the close relation existing between the two orchid
floras.

Vv. The description of a new monotypic orchid genus for New Zealand.

I. GENERAL SURVEY.

To the student of orchidology, the relation between the orchid flora of Australia and
that of New Zealand is so striking, and in some respects so remarkable, that a survey
of the subject seems long overdue. The present attempt to provide this can scarcely be
regarded as more than preliminary, but the authors are hopeful that it may at least
clear up some obscurities, and pave the way for other workers who may be able to reach
satisfactory conclusions as new light is thrown upon the subject in the course of time.

The figures given below, in connection with the numbers of orchid genera and
species, must be taken as approximate only. They are as nearly correct as it is possible
for us to make them at the time of writing. But no comprehensive census nor catalogue
is available, including every genus and species effectively published for Australia and
New Zealand up to the present time; and so far as Australian orchids are concerned,
descriptions of new species have been published in so many different journals (not all
of them Australian), that it is possible we have missed a few. We have been as accurate
as our sources of information permitted.

(a). The Orchid Fiora of Australia.—This is distributed among 71 genera, containing
about 470 known species. The number will probably be substantially increased before a
comprehensive census becomes possible; new species are being added every year, and
comparatively little is known as yet of the orchids of the tropics between the north-
west of Western Australia and the Cape York Peninsula. In the table which follows, the
number of known species each of the six Australian States is given, with an extra column
for the Northern Territory.

Number of Known Species.

Genus. Qd. N.S. W. Vict. Tasm. S. Aust. W. Aust. INCAS
Habenaria, lL. 8 9
Thelymitra, Forst. 2 1} 22 10 16 20
Epiblema, R.Br. iL
Diuris, Sm. 9 29 ial U 8 ut
Orthoceras, R.Br. 1 1 1 al iL
Microtis, R.Br. 2 4 5 4 4 9
Goadbyella, Rogers 1
Corunastylis, Fitzg. 1
Prasophyllum, R.Br. 9 44 27 14 13 18
Caleana, R.Br. 2 3 3 2 2 al
Spiculaea, Lindl. “I 2 2 al

4

Drakaea, Lindl.
J

54 i RELATION OF THE ORCHID FLORA OF AUSTRALIA TO THAT OF NEW ZEALAND,

Number of Known Species.
Genus. Qd. N.S.W. Vict. Tasm. S. Aust. W. Aust. INGE

Chiloglottis, R.Br. il 5 5
Acianthus, R.Br. 5 4 3
Townsonia, Cheesmn. by
Eriochilus, R.Br. 1 1 il il
Leptoceras, Windl. 1 1 1
Calochilus, R.Br. % 3

Rimacola, Rupp

Lyperanthus, R.Br. alt
Burnettia, Lindl.
Caladenia, R.Br.
Adenochilus, Hook. f.
Glossodia, R.Br.
Corybas, Salisb.
Nervilia, Comm. ex Gaud.
Didymopleaxis, Griff. iL
Cryptostylis, R.Br.
Pterostylis, R.Br.
Galeola, Lour.
EHpipogum, Gmel.
Gastrodia, R.Br.
Rhizanthella, Rogers a
Cryptanthemis, Rupp 1

Spiranthes, Rich.
Zeuxine, Lindl.
Anaectochilus, Blume
Goodyera, R.Br.
Corymborchis, Thou.
Hetaeria, Blume
Cheirostylis, Blume
Microstylis, Nutt.
Liparis, Rich.
Oberonia, Lindl.
Phaius, Lour.
Calanthe, R.Br.
Spathoglottis, Blume
Pholidota, Lindl.
Geodorum, Jacks.
Eulophia, R.Br.
Cadetia, Gaud.
Dendrobium, Sw.
Fria, Lindl.

Phreatia, Lindl. :
Pachystoma, Blume - 1
Bulbophyllum, Thou.
Dipodium, R.Br.
Cymbidium, Sw.
Luisia, Gaud.
Phalaenopsis, Blume
Sarcanthus, Lindl.
Camarotis, Lindl.
Schoenorchis, Schltr.
Drymoanthus, Nich.
Saccolabium, Blume
Vanda, Jones.
Ornithochilus, Wall.
Taeniophyllum, Blume
Sarcochilus, R.Br.
Chiloschista, Lindl.
Thrixspermum, Lour.

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Of these seventy-one genera, only thirteen appear to be strictly endemic, viz.:

Epiblema. Drakaea. Rimacola. Rhizanthella.
Goadbyella. Eriochilus. Burnettia. Cryptanthemis.
Corunastylis. Leptoceras. Glossodia. Drymoanthus.

Spiculaea.

BY H. M. R. RUPP AND E. D. HATCH. 55

Nine of the endemic genera are monotypic. The exceptions are Spiculaea (3 species),
Drakaea (4 species), Hriochilus (5 species), and Glossodia (5 species).

(0) The Orchid Flora of New Zealand.—In Cheeseman’s Manual of the N.Z. Flora
(1925 ed.), p. 331, the orchids are distributed among twenty-two genera. One of these,
Cyrtostylis R.Br., is now absorbed into Acianthus R.Br.; but the number will remain the
same, as the genus Aporostylis, n. gen., described in section V of this paper, must be
added. In the table of distribution which follows, four geographic areas are recognized,
and are indicated by abbreviations thus: N.I., North Island; S.I., South Island; Stewart,
Stewart Island; and Sub-ant., the sub-antarctic Auckland, Campbell, Chatham, and other
groups. i

Number of Known Species.

Genus. N.I. S.1 Stewart. Sub-ant.
Thelymitra 14 3 2 2
Orthoceras iL il
Microtis 1 1
Prasophylluimn 5 2 1 al
Caleana © i
Chiloglottis .2 i! 1 1
Aporostylis, n. gen. 1 1 1 1
Acianthus 2 2
Townsonia iL
Calochilus 3 i
Lyperanthus 1 il 1 1
Caladenia ul 2 1 1
Petalochilus Rogers. 2
Adenochilus 1 1
Corybas 7 5 4 2
Pterostylis 12 9 3 2
Gastrodia 2 o 1 1
Spiranthes i iL
Karina Lindl. 3 2 2 1
Dendrobium 1 1 1
Bulbophyllum 2 2
Sarcochilus i al 1 iff

It will be observed that of these genera, only three—Aporostylis, Petalochilus and
Harina—are not in the preceding table of Australian genera. The first two are endemic
in New Zealand, with one and two species respectively. Harina will be mentioned again
in section II. It is surely obvious from a comparison of the tabulated orchid genera of
the two countries that a close relationship exists; and the evidence for it becomes still
clearer as we discover how large a proportion of the New Zealand orchid species are
actually identical with Australian species. This will form the subject-matter of
section III.

Il. THE PROBABLE ORIGIN OF AUSTRALIAN AND NEW ZEALAND ORCHID GENERA.

We think it necessary to preface our remarks on this subject by stating that, broadly
speaking, we accept the conclusions of Cockayne and Marshall in regard to the geological
and geographical history of the distribution of land-masses in the south-west Pacific.
This means that we reject the theory of any direct land-connection between Australia
and New Zealand later than early or middle Mesozoic time. To put it in Marshall’s words,
we believe that ‘“New Zealand has been separate from AusStralia at least for the period
that has elapsed since that continent received its reptilian, amphibian, insect and
mammalian fauna and the characteristic flora”. (Rep. Aust. and N.Z. Ass. Adv. Sci.,
Sydney, 1932, p. 411.) Therefore, though from an orchidological point of view there
would seem to be quite a substantial amount of circumstantial evidence favouring the
theory of a land connection across the Tasman Sea during the above period, we think
that the facts relevant to our subject are better explained on the hypothesis of an
extensive antarctic continent in early Cretaceous time (see Cockayne, Veget. of N.Z.,
1928, p. 422 et seq.).

Excluding the thirteen endemic Australian genera specified in section I, we have
fifty-eight non-endemic Australian genera. In 1923 Rogers (Trans. Roy. Soc. 8S. Aust.,

56 RELATION OF THE ORCHID FLORA OF AUSTRALIA TO THAT OF NEW ZEALAND,

xlvii, p. 331) gave the following as true generic types originating in Australia, but not
endemic:

Calochilus. Pterostylis. Cyrtostylis.
Thelymitra. Caleana Caladenia.
Orthoceras. Acianthus. Adenochilus.
Prasophyllum. Lyperanthus. Chiloglottis.
Microtis.

If we substitute Townsonia for the now obsolete genus Cyrtostylis, the number
remains the same in this list. But a careful study of the distribution of the species
comprising these genera reveals difficulties in the way of accepting an Australian origin
for all. It appears to us far more likely that some of the genera, or their ancestral
forms, originated in the antarctic (Palaeozelandic) continent already alluded to. This
continent is believed to have extended northward to include the land-masses now repre-
sented by New Zealand and its small island dependencies, and also Lord Howe Island,
and, less certainly, Norfolk Island; from Lord Howe Island there was an extension to
New Caledonia, Melanesia and New Guinea, with a probable land-connection there to
Australia. Further west the Palaeozelandic continent threw out another northern
extension to what is now Tasmania, which was then in direct land-connection with the
south of Australia. The development and distribution of the following orchid genera
lead us to regard them as having had their origins in this Palaeozelandic continent,
whence they spread northward to New Zealand and in some instances beyond it, and
also through the Tasmanian extension to the mainland of Australia:

Thelymitra. | Lyperanthus. Caladenia.
Chiloglottis. Townsonia. Pterostylis.

We agree, with certain reservations, that the remaining genera—Diuris, Orthoceras,
Microtis, Prasophyllum, Caleana, Calochilus, Acianthus and Adenochilus—may be
considered as probably of Australian origin, though not endemic there. The reservations
concern Microtis, which is represented by one or two species over a large area of eastern
Asia, and may conceivably have had an Asiatic origin; Acianthus, the remarkable
development of which in New Caledonia (thirteen species) seems to require explanation
if the genus was originally Australian; and Adenochilus, which may possibly have been
Palaeozelandic. With regard to the monotypic genus Townsonia (see also section III),
it would appear that its progress beyond Tasmania was arrested by the formation of Bass
Strait.

Eliminating, then, the Australian endemic genera (13), the genera here admitted
as of Australian origin though not now endemic (8), and those we regard as of Palaeo-

zelandic origin (6), there still remain forty-four genera of Australian orchids to be.

accounted for. According to Rogers (Presidential Address, Botany Section, Rep. Aust.
and N.Z. Ass. Adv. Sci., Sydney, 1932, p. 339), in the great majority of instances these
can be traced back through New Guinea or the Malay Archipelago, into the continent of
Asia proper. They may be considered as of Asiatic or other origin, or at least it may be
confidently affirmed that they did not originate in the south-west Pacific. They are as
follows:

Habenaria. Goodyera. Geodorum. Phalaenopsis.
Corybas. Corymborchis. Eulophia. Sarcanthus.
Nervilia. Hetaeria. Cadetia. Camarotis.
Didymoplewis. Cheirostylis. Dendrobium. Schoenorchis.
Cryptostylis. Microstylis. Eria. Saccolabium.
Galeola. Liparis. Phreatia. Vanda.
Epipogum. Oberonia. Pachystoma. Ornithochilus.
Gastrodia. Phaius. Bulbophyllum. Taeniophyllum.
Spiranthes. Calanthe. Dipodium. Sarcochilus.
Zeuxine. Spathoglottis. Cymbidium. Chiloschista.
Anaectochilus. Pholidota. Lwuisia. Thrixspermum.

Turning now to the New Zealand genera, we find that all but three—arina,
Aporostylis and Petalochilus—are already accounted for in the foregoing lists. The
Asiatic element, which probably came over the Palaeozelandic continent via New
Caledonia, is small:

Corybas. Spiranthes. Bulbophyllum.
Gastrodia. Dendrobium. Sarcochilus.

BY H. M. R. RUPP AND E. D. HATCH. 57

Genera of Australian origin:

Orthoceras. Caleana. Calochilus.
Microtis? Acianthus. Adenochilus?
Prasophyllum.

Possibly Adenochilus should be transferred to the next list. There is one species in
New Zealand and one in Australia, and they may have developed from a common Palaeo-
zelandic ancestral form.

Genera of Palaeozelandic origin:

Thelymitra. Lyperanthus. Caladenia.
Aporostylis. Townsonia. Pterostylis.
Chiloglottis. 2

The genus Harina presents some difficulties. There are three species in New Zealand,
and others occur in New Caledonia, Fiji, and other south Pacific Islands; but the genus
is not represented either in Australia or Indo-Malaya. It may have developed in the
Palaeozelandic continent after the severance of Tasmania from the latter; but it is not
impossible that it is of Polynesian origin. With regard to the two endemic genera
Aporostylis and Petaiochilus, the former is sufficiently dealt with in section V, while
Petalochilus is a somewhat anomalous genus very closely allied to Caladenia, but
entirely lacking the labellar glands so characteristic of the latter. It appears to be
strictly a ‘local development” for the only two species discovered are confined to a very
small area in the extreme north of the North Island.

This is a suitable place for a note on the orchids of Lord Howe and Norfolk Islands,
both of which lie between Australia and New Zealand. On Lord Howe Island there is
one Australian species of Dendrobium (D. gracilicaule F. Muell. var. Howeanum Maiden),
one endemic species of the same genus (D. Moorei F. Muell.), one New Zealand species of
Bulbophyllum (B. tuberculatum Col.), one endemic species of Sarcanthus (S. erectus
(R. D. Fitzg.) Rupp), and one species of Microtis (M. unifolia (Forst.) Reichb. f.). Thus
there are connecting links here with both Australia and New Zealand. On Norfolk
Island, however, with the exception of the ubiquitous Microtis unifolia, the orchid flora
is entirely endemic, consisting of two species of Dendrobium (D. brachypus Reichb. f.
and D. macropus Benth. & Hook. f.), one species of Bulbophyllum (B. argyropus Reichb.
f.), and one of Phreatia (P. obtusa Schltr.). If then Norfolk Island was originally part
of the Palaeozelandic continent, the development of its orchid flora suggests that it
became isolated at a very early period. In both islands the presence of Microtis unifolia
might be explained by seeds carried by wind or birds from Australia, where this orchid
is very common.

III. Orcuip SPECIES COMMON TO AUSTRALIA AND NEW ZEALAND.

In 1932, Rogers (l.c., p. 341) put the number of these at twenty, with two others
doubtful. Our own investigations have increased the number to twenty-eight, with
four others sufficiently doubtful to require further study when fresh material is avail-
able on both sides of the Tasman Sea.*

These conspecific orchids may be conveniently dealt with under two divisions, viz.:
Those which we regard as entirely identical except for such slight and unimportant
variations as occur in all plant species (Table A) and those which, although specifically
identical, show sufficient constant variation from the type to be ranked as named varieties
(Table B).

In tabulating the species in Table A, we give in the first column the valid name of
each species, and in the second column the synonyms in those cases where it has been

* Since the completion of this paper, further investigation has conviced us that three more
species must be added to the list of those which are common to both countries. The New
Zealand Corysanthes Matthewsii Cheesmn. is undoubtedly conspecific with the Australian
Corybas unguiculatus (R. Br.) Reichb. f., and should be known in future by the latter name.
On the other hand, the Australian Chiloglottis Muelleri R. D. Fitzg. and Caladenia alpina Rogers
are conspecific respectively with the New Zealand Chiloglottis cornuta Hook. f. and Caladenia
Lyallii Hook. f. Our opinion in regard to these has been fully endorsed by Messrs. J. H. Willis
of the Victorian National Herbarium, and W. H. Nicholls, Hon. Curator of the Melbourne
University Herbarium. The name Corysanthes Matthewsii therefore becomes a synonym of
Corybas ungwiculatus; the name Chiloglottis Muelleri a synonym of OC. cornuta; and the name
Caladenia alpina a synonym of C. Lyallii.

58 RELATION OF THE ORCHID FLORA OF AUSTRALIA TO THAT OF NEW ZEALAND,

necessary to restore the valid name. Further information on such cases will be found
in the notes which follow Table B.

TABLE A.
Valid Name. Synonymy.
Thelymitra ixioides Sw.
longifolia Forst.
aristata Lindl.
paucifiora R.Br.
venosa R.Br.
Matthewsii Cheesmn. .. .. .. YZ. D’Altonii Rogers.
Orthoceras strictum R.Br. :
Microtis unifolia (Forst.) Reichb. f.
Prasophyllum patens R.Br.
Rogersii Rupp
Caleana minor R.Br.
Chiloglottis formicifera Fitzg.
Calochilus Robertsonii Benth. .. .. .. .. OC. campestris sensu Cheesmn., non R.Br. (see
note 4 below).
paludosus R.Br. :
Townsonia viridis (Hook. f.) Schltr. .. .. Acianthus viridis Hook. f.: Townsonia deflexa

Cheesmn.
Corybas aconitiflorus Salish. we ee ee ee Corysanthes Cheesemanii Hook. f. ex T. Kirk.
Pterostylis nutans R.Br. ay ee eee ath nenusiimaCheesmn.
nana R.Br. TAU CER aba Torey eC puberula Hook. f.
PUMPOCn Koh JALAN ah S Si- Lalo og 8.0 micromega Hook. f.
foliata Hook. f. Slvr uaslisn Pouiks , Nas gracilis Nich.

mutica R.Br.
barbata Lindl.
Gastrodia sesamoides R.Br.

Spiranthes sinensis (Pers.) Ames .. .. .. S. australis Lindl.
TABLE B.
Named varieties.
Valid Name. Synonymy.
Caladenia carnea R.Br: var. pygmaea Rogers! .. .. .. .. Caladenia minor Hook. f.
carnea R.Br. var. exigua (Cheesmn. )
WON) | oo oo Se Ge aT On hence Caladenia exigua Cheesmn.

Thelymitra carnea R.Br. var. imberbis (Hook.

f.) Rupp and Hatch we Be Nae Tyson ee ens a tata (as Thelymitra imberbis Hook. f.
Acianthus reniformis (R. Br.) Schltr. var.

oblongus (Hook. f.) Rupp and Hatch .. .. . Cyrtostylis oblonga Hook. f.

It would not be profitable to discuss here the Hout doubtful species alluded to above.
‘But some notes on the changes of nomenclature involved in the foregoing tabulation are
desirable, and in a few instances we briefly comment where no change of name has been
necessary.

1. Thelymitra Matthewsti Cheesmn.—In 1930 Rogers (Trans. Roy. Soc. 8S. Aust.,
liv, p. 42) described a species of Thelymitra from the Victorian Grampians under the
name 7. D’Altonii, which was subsequently found in eastern Victoria also. At it became
better known, its resemblance to the New Zealand JT. Matthewsti was generally recog-
nized; and Nicholls (Vict. Nat., lvii, 1940, p. 83) records T. D’Altonii as a synonym only.

2. Prasophyllum Rogersii Rupp.—This was described as a new species from the
plateau of Barrington Tops, N.S.W. (these PROCEEDINGS, liii, 1928, p. 340). Shortly after-
wards the author received specimens from the late H. B. Matthews of a Prasophyllum
collected at Kaitaia, N.Z., which he determined as identical with his Barrington Tops
species. It has subsequently been found near Hobart (A. M. Olsen), and, according to
a personal communication from W. H. Nicholls, in the far east of Victoria.

3. Chiloglottis formicifera Fitzg.—To anyone familiar with the habitat of this
comparatively rare species in Australia (where it is restricted to a limited area in New
South Wales), its occurrence in New Zealand is very puzzling. This will be discussed
further in section IV.

4. Calochilus Robertsonii Benth.—Cheeseman (Manual of N.Z. Flora, p. 357),
recording for New Zealand what he took to be C. campestris R.Br., stated that specimens
exactly matched R. D. Fitzgerald’s plate in Aust. Orch., i (4), 1878. But Fitzgerald was
mistaken in his interpretation of the species; the plant he depicts as C. campestris is

BY H. M. R.-RUPP AND E. D. HATCH. 59

not that species at all, but the pale-flowered form of C. Robertsonii (see Rupp, these
PROCEEDINGS, 1xix, 1944, p. 277). In Rupp’s herbarium there are typical specimens of
C. Robertsonii collected by H. B. Matthews at Rotorua. An admirable plate of
C. campestris may be seen in Bot. Mag., 1832, t. 3187. In our opinion this species has not
been found in New Zealand.

5. Townsonia viridis (Hook. f.) Schltr—tIn Vol. ii of his Flora Tasmaniae, 1850,
under “Additions and Corrections”, Hooker described a small orchid of mountain gullies
as Acianthus viridis. In the 1906 edition of Cheeseman’s Manual of N.Z. Flora, the
author created a new genus, Townsonia, the single species being named 7. deflexa
(p. 692). Schlechter (Fedde, Repert., ix, 1911, p. 249) transferred Hooker’s Acianthus
viridis to Townsonia. Rupp (Vict. Nat., 1, 1933, p. 18) discussed and illustrated both
plants, expressing the opinion that they were conspecific. We are now completely
satisfied that this view is correct. It seems a pity that the name of the author of this
monotypic genus should be excluded from the valid nomenclature of the species; but
according to international rules it must stand as 7. viridis (Hook. f.) Schltr.

6. Corybas aconitiflorus Salisb—There is no doubt that, as Cheeseman himself
hinted (Manual of N:Z. Flora, p. 364), Corysanthes Cheesemanii Hook. f. ex Kirk is
identical with C. bicalcarata R.Br. Since the International Council for Nomenclature
decided against the conservation of Brown’s nomenclature, that of Salisbury must be
adopted (see Rupp, Vict. Nat., lix, 1942, p. 60).

7, 8. Pterostylis nutans R.Br.; Pterostylis nana R.Br.—There can be no doubt
that these are the valid names respectively for P. Matthewsti Cheesmn. and P. puberula
Hook. f.

9. Pterostylis furcata Lindl.—We have very carefully compared specimens of this
with P. micromega Hook. f. The latter appears to have slightly more acute sepals and
petals; otherwise they agree perfectly, and should be regarded as conspecific.

10. Pterostylis foliata Hook. f—Nicholls (Vict. Nat., xliii (11), 1927, p. 324)
described a Pterostylis found in Victoria and Tasmania as P. gracilis. It has since been
recognized as identical with P. foliata Hook. f., though we have been unable to trace any
previous publication of the identity.

11, 12. Caladenia carnea R.Br. var. pygmaea Rogers, and var. exigua Cheesmn.—
Rupp (these Proceeprnas, Ixix, 1944, p. 74) reduced the New Zealand C. minor Hook and
C. exigua Cheesmn. to the above varieties of C. carnea. Since then, additional material
of C. carnea var. pygmaea has raised some doubt as to whether it would not have been
better to retain Hooker’s name as that of a distinct variety. But even the varieties of
C. carnea are themselves so liable to vary, that for the present at all events we think it
best to let the matter rest. We feel no doubt that both the New Zealand forms really
belong to C..carnea, and both can be matched freely in Australia.

13. Thelymitra carnea R.Br. var. imberbis (Hook. f.) Rupp and Hatch.—We have
reduced Hooker’s 7. imberbis to varietal rank with some hesitation. Specimens in
Rupp’s herbarium received from H. B. Matthews are more robust than any form of
T. carnea he has seen, and the column is stouter. But the morphology of the flowers is
almost identical, and there does not seem to be any distinction warranting specific
separation. :

14. Acianthus reniformis (R.Br.) Schltr. var. oblongus (Hook. f.) Rupp and
Hatch.—The New Zealand plant seems consistently more diminutive than the typical
form of A. reniformis, but apart from this and the oblong leaf we can find nothing to
distinguish them. Hatch is convinced that Cheeseman was right in: sinking C0. rotundi-
folia Hook f. (Manual of N.Z. Flora, p. 356). In the Australian plant the leaf is by no
means always reniform, but is often orbicular or even cordiform.

IV. PossiBLE EXPLANATIONS OF THE CLOSE RELATION EXISTING BETWEEN THE ORCHID FLORAS
or AUSTRALIA AND NEW ZEALAND.

The fact of this relation will have been made abundantly clear in the foregoing
sections of this paper. Various explanations have been offered for it. The hypothesis of
a primeval antarctic continent, to which we expressed our adherence in section II,
explains much, but it certainly does not explain everything. It provides a rational

60 RELATION OF THE ORCHID FLORA OF AUSTRALIA TO THAT OF NEW ZEALAND,

explanation for the development of such genera as Thelymitra and Pterostylis along
similar lines in both countries; and if certain species were in process of being evolved
into their present forms when the Palaeozelandic continent was broken up, no one can
assert that there was anything in the new conditions to prevent the continuance of their
development into identical forms. But if ancestral forms of Caladenia were also included
in the Palaeozelandic orchid flora, why did they develop with such remarkable richness
of colouring and great variety of form on the Australian side, and so poorly on the New
Zealand side? Take the case of another genus—Chiloglottis. A common ancestral form
in the ancient continent might well develop into C. cornuta in New Zealand, and into
C. Gunnii, C. Muelieri, and perhaps C. Pescottiana in Australia; their close affinity is
obvious. But we cannot believe that in the far north of New Zealand they would evolve
so different a form as (©. formicifera, completely identical with a relatively rare New
South Wales species. We have much to learn yet. This particular species is of special
interest. In Australia it occurs, nowhere in great abundance, but in considerable
“colonies”, from the Hunter River on the north to the Shoalhaven River on the south.
It inhabits well-shaded forest gullies. How did it reach New Zealand? Was the seed
carried across the 1,200 miles of the Tasman Sea by dust-storms? We know that in
times of drought very considerable amounts of Australian dust are occasionally deposited
on New Zealand; and orchid seeds are extremely minute. But is it likely that seeds of

a dwarf terrestrial orchid from forest gullies of the coastal belt would be caught up and

transported by a dust-storm from the dry interior? Another suggestion is that the
seeds might have become attached to the feathers of migratory birds. They might;
and that is all we can say at present. We do not deny that both birds and dust-storms
may have been responsible for the appearance in New Zealand of certain orchid species.
But is it not remarkable that, although New South Wales has nearly thirty species of
Diuris on record, not a single species of this genus has been seen in New Zealand?
Many species produce seeds quite freely, and some grow in almost any type of country.
Or take the genus Cymbidium. The North Island of New Zealand should afford conditions
suitable at least for C. candliculatum and C. suave. Both produce immense quantities of
fine, dust-like seeds, easily carried by wind from their arboreal homes. Yet neither
occurs in New Zealand. Moreover, if birds and winds have transported orchid seeds
successfully across the Tasman Sea, are they doing it still? We do not pretend to answer
these questions, but we think they should be faced, and patient research and study will
no doubt in time be rewarded by glimpses of the truth. As yet there is no complete
explanation of the relation between the two orchid floras. They are explained in part
if we accept the theory of the Palaeozelandic continent, for that allows us to believe in
an inflow of allied ancestral forms both from north and south; in part perhaps by the
agencies of birds and winds; more than this we cannot say. The distribution of orchids
may not seem a subject of great importance in itself. But it must be remembered that
the Orchidaceae now rank as the largest family of flowering plants; and, in the words
of the late Dr. R. S. Rogers, facile princeps among Australasian orchidologists, “It is
obviously desirable that such a matter as their distribution should be established as
accurately and as early as possible. On it may depend, to some extent, the solution of
much greater questions concerning the former disposition of land-masses, the origin
of our flora, and the true relation of our continent” (and, we may add, of New Zealand)
“to other portions of the globe”. (Trans. Roy. Soc. 8. Aust., xlvii, 1923, p. 322.)

V. A New OrcHID GENUS FOR NEW ZEALAND.
APOROSTYLIS, nN. gen.

Genus monotypicum. Planta terrestris 7-23 em. alta, plerumque pubescens,
luberibus parvis. Bractea basalis lata, acuminata, bracteae caulinae absentes. Folia
duo, inaequalia, fere basalia vel folium minus altius quam folium majus; patentia, breviter
petiolata, 3-7 cm. longa; majus magnopere latius quam minus. Flos solitarius, albus vel
puniceus, cum sub ovario bractea laxe vaginante. Sepalum dorsale lanceolatum, erectum,
circiter 15 mm. longum: sepala lateralia, tam longa quam dorsale: petala similia, paulum
breviora. Labellum sessile, prope basem erectum, deinde paulum recurvum, obovatum
vel fere orbiculare, apice rotundo et marginibus laevis, circiter 12 mm. longum: discus

OO EE

BY H. M. R. RUPP AND E. D. HATCH. 61

cum glandium flavidorum ordinibus duobus. Columna illae Chiloglottis instar, sed alis
non pone antheram extendentibus.

A monotypic genus created to absorb the anomalous species Caladenia bifolia Hook.
t. (Fl. Nov. Zel., i,.18538, p. 247). The description of the genus is therefore that of the
solitary species, Aporostylis bifolia (Hook. f.) Rupp and Hatch.

A terrestrial herb 7-23 cm. high, usually pubescent or even hirsute but occasionally
glabrous, with small tubers. General habit that of Chiloglottis. Sheathing bract at the
base of the stem broad, acuminate; cauline bracts absent. Leaves two, unequal, almost
basal or the smaller one above the larger; spreading, shortly petiolate, 3-7 cm. long;
the larger leaf usually very much broader than the smaller one but varying from linear-
lanceolate to ovate-oblong or almost orbicular, mucronate; the smaller one elliptical to
broadly linear, acute. Flower solitary, white or pink, with a loosely-sheathing bract
subtending the ovary. Dorsal sepal erect, lanceolate, about 15 mm. long; lateral sepals
broad-linear, about as long as the dorsal; petals similar but a little shorter. Labellum
sessile, the basal portion erect, then gently recurved, obovate or almost orbicular, with
rounded apex and entire margins, about 12 mm. long: disc with two rows of yellow calli
extending from the base to about the middle. Column resembling that of Chiloglottis,
but with wings neither lobed nor produced behind the anther.—Caladenia bifolia Hook.
f., l.c.; Cheeseman, Man. N.Z. Fl., 1925, p. 360, and Jllustr. N.Z. Fl., ii, 1914, t. 197 B;
C. nacrophylla Colenso, Trans. N.Z. Inst., xxvii, 1895, p. 396; Chiloglottis Traversti
F. Muell., Veg. Chath. Is., 1864, p. 51; Ch. bifolia (Hook. f.) Schltr., Hngl. Bot. Jahrod.,
shy, UG, io), eter

Distribution.—New Zealand: North and South Islands, Stewart Island, Chatham and
Auckland Islands. Usually alpine or sub-alpine, but descending to sea-level in Stewart
and the outlying islands.

“A curious plant, the genus of which is doubtful’ (Cheeseman, Man. N.Z. FI1., l.c.).
Hooker placed it in Caladenia, Mueller removed it to Chiloglottis, and Schlechter endorsed
this, restoring Hooker’s specific name. In either case it exceeds the limits of the generic
character, and we believe that the most satisfactory way out of the difficulty is to make
it the type of a new genus. It probably originated as an inter-generic cross between
ancestral forms of Chiloglottis cornuta Hook. f. and Caladenia Lyallii Hook. f. Its
distribution is very similar, and it is reasonable to suppose that these species, or their
ancestral forms, were spread over the ancient Zelandic continent which arose in the
Cretaceous period. The affinities of Aporostylis with Chiloglottis and Caladenia are
obvious; but the anomalous character of the column distinguishes it from either. The
general habit, the occasionally glabrous surface of stem and leaves, and the structure of
the column apart from its wings, are all reminiscent of Chiloglottis; but the sub-erect,
gently recurved labellum with two rows of yellow calli, the broad column wings neither
lobed nor produced behind the anther, and the common pubescence of stem and leaves,
are more suggestive of Caladenia. The great variability of the leaves, and their alterna-
tion between the glabrous and pubescent forms, seem to indicate a hybrid origin.

62

GEOGRAPHIC VARIATION IN THE LIZARD HEMIERGIS DECRESIENSIS
(FITZINGER).

By STEPHEN J. COPLAND, B.Sc.
(Plate vi; five Text-figures.)

[Read 27th June, 1945.]

Contents.

Page
I. Introduction ae rh CC ae) cared sh cesuamTe Memes BeeU IAT ceHe atin Woeree cae Reese rep male Sees Benen (31>
Il. Hemiergis decresiensis Gaitzineer) ee ae aie Pe Riis tas Ce tale teu Batre cnet 3
Ill. Hemiergis decresiensis decresiensis GBtinteee) Pecan srry in Pan WEN ELC Son rope ithe tea eu ml~ i ayel
IDV TEAC CMS COCTOH GNSS) COMHUCO DUES Ms SNS 65 65 sa $6 oe bo bo oo on 5a
V. Hemiergis decresiensis talbingoensis, n. subsp. SES NSPE Ue cate noua, Urol waar Aiea trae oman a ae ae Ti
Wile JsICMHGIOIS CEGTONCIMS COIN, WW. GUND co 08 case oo oc of 60d 66. oo so 7
VII. Relationship of the four subspecies .. Seg Ma eR ETE es Rea teeth OE OSIM R oe cae ms ean
VIII. Conclusions ots Ale etna aa SEAS RUT RENEE BE aa a ats, Ses o's Dake arenes oo hae Bee eT
IX. Western Mistral ecorde 30 6.4 cken Syne eee, os BG ads ae Sie hae tes; testo
xX. Acknowledgements ep ge bin ea a syteooe "is hl ct chk heneg Oelg rcrper kc Bag era p
x 1 Bibliography Be ee eng tests Fe SEN 2 ers Snr ate ee Cane, eae ate Pea aoe in Sauleo an Cul

I. INTRODUCTION.

An attempt has been made to deal with one species of Australian lizards as
comprehensively as possible from the taxonomic aspect. Every specimen of the skink
Hemiergis decresiensis (Fitzinger) from the collections of the Australian Museum,
Sydney, and the South Australian Museum has been examined as well as a few from the
National Museum, Melbourne, and large New South Wales series in the author’s collection.
As a result the species has been divided into four subspecies. A standard description
is given of a topotype of the typical race H.d. decresiensis from Kangaroo Island, to -
which it is restricted as far as is known at present. The mainland race from South
Australia, and probably western Victoria, which approximates most closely to the
nominate form, is shown to differ in size and other characters and has been named
H.d. continentis. The two remaining subspecies—H.d. talbingoensis from the Southern
Tableland of New South Wales, and probably the north-eastern highlands of Victoria,
and H.d. davisi from the Northern and Central Tablelands of New South Wales—are
shown to differ from one another and H.d. continentis in scale and other characters.
Records of H. decresiensis from Western Australia are queried.

The great majority of species of Australian lizards has probably been described, but
much work done on this section of the reptiles remains rather isolated in literature and
uncorrelated. There have been few papers on single species in which all material in the
museums of the different states has been brought together and correlated against every
reference to it in literature. Complete locality records and other references to many
species can normally only be found by painstaking search through numerous publications
in several languages. Final generalizations on Australian lizards cannot be made until
at least a large percentage of species has been given separate comprehensive treatment.
Present collections in museums will not permit these studies to be carried out in most
cases, either because of the small number of specimens or the fact that they come from
restricted localities. Studies approaching completeness will therefore in general entail
planned collecting at key points within the area of distribution of each species. The
present paper, although reasonably complete for New South Wales and South Australia,
suffers through lack of Victorian material.

BY STEPHEN J.. COPLAND. 63

It may be mentioned here that there is ample justification, in the author’s opinion,
tor retaining the genus Hemiergis in spite of the view of Malcolm A. Smith (1937, p. 213)
that “such groups as Siaphos and Hemiergis, which are merely assemblages, mainly of
‘degenerate species, and are not capable of being defined, have been abandoned”. In
Smith’s paper (p. 223 et seq.) all members of this purely Australian genus are placed in
Leiolopisma. It is clear that Hemiergis has been derived from an ancestral stock closely
allied to the modern Leiolopisma, but the links have disappeared. The fact that there
is marked differentiation and that it has occurred solely in Australia is sufficient to
validate the genus even if parallel series were evolved elsewhere.

In the present paper, all relevant literature known to the author has been noted,
except bare references the inclusion of which would have added nothing of significance.

There are so many small differences between the 250 odd specimens of the four
subspecies examined that to compare each holotype with the topotype of H.d. decresiensis
and then note variations within each race at another stage removed would result in
hopeless confusion. The unavoidable course of giving detailed descriptions of the
nominate race and H.d. talbingoensis and H.d. davisi has been adopted. The small
number of specimens (26) and the fact that they are closely allied allow the Kangaroo
Island and South Australian forms to be treated together.

II. HEMIERGIS DECRESIENSIS (Fitzinger).

Zygnis decresiensis Fitzinger, 1826, p. 53. Tridactylus decresiensis Cuvier, 1829,
p. 64: Gray, 1831, p. 72 and 1839, p. 333. Hemiergis decresiensis Duméril and Bibron,
1839, p. 766; Gray, 1845, pp. 87 and 272; Steindachner, 1867, p. 50, part; Gtinther, 1875,
Plate 6, fig. 5; Lucas and Frost, 1894, p. 24; Lucas and Le Souef, 1909, p. 255: Waite,
1927, p. 326, and 1929, p. 161. Hemiergis polylepis Giinther, 1867, p. 48. Lygosoma
decresiense Boulenger, 1887, p. 327; Lonnberg and Andersson, 1913, p. 9; Zietz, 1920,
p. 216. Lygosoma (Hemiergis) decresiense Werner, 1910, p. 481, in error; Proctor,
1923, p. 81. Hemiergis decresiense Loveridge, 1934, p. 368.

Duméril and Bibron (1839, p. 766) give the following synonymy, points of which
are dealt with in the preceding list or later in this paper, while, because of lack of data,
others could not be checked:

“Tridactylus Decresiensis Péron, Mus. Par. Zygnis Decresiensis Fitz., Neue classif.
Rept. Verzeichn., p. 53, No. 4. Seps aequalis Gray, Ann. Philosoph., tom. 10 (1825), p. 202.
Seps (Tridactylus Decresiensis Péron), Leuckart, Breves animal. quorund. Descript.,
p. 10. Seps (Tridactylus Decresiensis Péron), Cuv. Régn. anim., 2° edit., tom. 2, p. 64.
Seps (Tridactylus Decresiensis Péron), Griff. anim. Kingd. Cuv., tom. 9, p. 159..
Hemiergis Decresiensis Wagl., Syst. amph., p. 160. Péron’s seps. Gray, Synops. Rept.
in Griffith’s Anim. Kingd., tom. 9, p. 72. ? Siaphos aequalis, id., loc. cit., p. 72. Peromeles
aequalis Wiegm., Herpet. Mexic., pars 1, p. 11. Tridactylus Decresiensis Gray, Catal.
slender-tong. Saur., Ann. of natur. hist., by Jardine, tom. 1, p. 333.”

One of the references given by Boulenger (1887, p. 327) may be noticed: Hemiergis
decresiensis Gray, Zool. Ereb. and Terr., Rept., Pl. vi, fig. 5, but this plate was omitted in
1845 and, according to Giinther’s notes in the introduction, redrawn for the publication
of 1875 (see Giinther, 1875).

There is no doubt as to the validity of the specific name of Hemiergis decresiensis
(Fitzinger) and little doubt as to the genus, but considerable uncertainty as to the
author. The original lizard or lizards in the Paris Museum were apparently labelled
Tridactylus decresiensis by Péron, but there was no accompanying publication of the
name and description. Accordingly the name was given no standing. This specific name
has been since used by all authors (omitting modifications in word endings) with the
exception of Gtinther’s polylepis. First publication was by Fitzinger (1826, p. 53), and
I have to thank Mr. W. A. Rainbow, Librarian at the Australian Museum, for securing”
me a photostat copy of this rare work. Fitzinger says: “7. Zygnis decresiensis. m.
Decresische Z. (Tridactylus decresiensis. Mus. Paris). Ex Australia, Insula Decres.’’
This is the entire reference and decresiensis is marked as a nomen nudum in Sherborn’s
Index Animaltum. This would invalidate Fitzinger’s claim to authorship of the species.
Nevertheless there is not a bare mention of the name bare of context, and it indicates.

64 VARIATION IN THE LIZARD HEMIERGIS DECRESIENSIS,

the species better than many type descriptions which have been allowed in this and
other groups. Justification may reside in the nomination of the actual specimen or
specimens on which the name is based and the place where they were to be found, i.e.,
“Tridactylus decresiensis. Mus. Paris’, and the type locality “Insula Decres”. Placing
the species in the genus Zygnis automatically attributes it with the following characters
given in the same work (p. 23) in the key to the Scincoidea: “Pedes quatuor: Pori
femorales nulli: Digiti non dilatati: Plantae non pentadactylae: tridactylae”’, but as he
includes two other species in the same genus the characters cannot be considered a
species description. The next author, Cuvier (1829, p. 64), with: “Une a quatre doigts,
dont les postérieurs inégaux (le Tetradactylus decresiensis, Per.) et une a trois, d’ailleurs
trés semblable a la précédente (Tridactylus decresiensis, Per.). Toutes deux viennent
de Vile de Crés, et sont vivipares’”, adds little or nothing to Fitzinger, but the form of
the statement may entitle him to the authorship of the species, which would then be
Hemiergis decresiensis (Cuvier). If for any cause both authors were ineligible the
species must be attributed to Gray (1831, p. 72) with: ‘“Peron’s Seps. Tridactylus
Decresiensis, Peron. Toes 3, 3; hinder unequal, one short and two long, subequal; pale
brown, with long dark lines; beneath netted.” As far as generic names are concerned,
Zyenis Oken 1816, Lehrb. Nat., 3 (2), 284.—Rept. and Tridactylus Olivier 1789, Ency.
Méth., 4 (Ins.), 26——Orth., and Lacepéde 1799, Tabl. Oiseaux, 11.—Aves, are unavailable
and Hemiergis Wagler 1830, Syst. Amph., 160, must be used. These last four references
are in the form given in Neave’s Nomenclator Zoologicus, London, 1939.

III. HEMIERGIS DECRESIENSIS DECRESIENSIS (Fitzinger). Pl. vi, fig. 1.

Topotype. No. R.2191 in the South Australian Museum; Kangaroo Island, c. 35.50 S.,
137.20 E., 1885.

Description of Topotype.—Rostral* moderately high, area visible from above equal
to nearly one-half that of the frontonasal, long concave sutures with the nasals and
slightly concave, approximately vertical ones with the ist supralabials; the nearly
straight junction with the frontonasal is about one-third the width of the frontal. Nasals
large, not in contact, roughly quadrilateral, long convex sutures with the rostral, fronto-
nasal, and anterior loreal, nearly straight with 1st supralabial; round nostril slightly
behind centre, no sign of groove running from it to separate scale. No supranasals.
Frontonasal large, subequal in area to the frontal, with which it forms a suture about
one-eighth the width of the latter scale, also in contact with prefrontals, nasals, rostral

and anterior two-thirds of the upper margin of the anterior loreal. Prefrontals large,

well developed, four-sided, sutures long and slightly convex with frontonasal, nearly
straight with frontal, concave with ist supraciliary, concave against the anterior and
straight against the posterior loreal, point of contact with 1st supraocular. Frontal
kite-shaped, indented in front against the frontonasal and rounded behind between the
frontoparietals, pointed laterally where frontal, prefrontal, 1st supraciliary and 1st supra-
ocular touch, long, straight, postero-lateral sides against Ist and 2nd supraoculars, shorter
antero-lateral sutures with prefrontals. Frontoparietals paired, large, subequal in size
with the interparietal, left scale a rough crescent, twice as long as wide, inner convex
border against parietal, interparietal and its fellow, outer border nearly straight against
2nd, 3rd and 4th supraoculars, indented against frontal; right frontoparietal in contact
with the same scales, but more squat, roughly pentagonal, and pointed mediad between
interparietal and left frontoparietal, suture with frontal only about one-third the length
of the contact of the left frontoparietal with the same scale. The interparietal kite-
shaped, smaller than frontal, rounded behind, pointed in front and at sides, sutures long
and straight with parietals, shorter with frontoparietals, concave with left, sinuous with
right; a rounded milky area in the midline one-third the length of the scale from the
posterior end covers the pineal foramen. Parietals are the largest head shields, irregularly
shaped and at least twice as long as wide, meeting in an oblique suture behind the
interparietal, other sutures straight and long with the interparietal, shorter and slightly
coneave with frontoparietals, very short with 4th supraocular, 8th supraciliary, and 2nd
postocular, straight with upper secondary temporal, left scale in contact with two dorsal

* For designation of scales see illustration of H.d. davisi (Figs. 2 and 3).

BY STEPHEN J. COPLAND. 65

seales, the right with four, one at hardly more than a point. There are no nuchals, if a
large irregular scale on the left be excluded. Seven supralabials, the anterior three
roughly quadrilateral, their upper margins forming a nearly straight, horizontal line
with the nasal, loreals and lower preocular, postero-dorsal angles of the 1st and 2nd
project backwards, that of 3rd does not, the smaller 4th is under the 1st subocular, and
the 5th, which is much smaller again, under the 2nd and 3rd suboculars; 6th and 7th
are large, the posterior three scales are pentagonal, haystack-shaped, lower margins
horizontal, anterior and posterior sutures vertical, and the other two sides meeting in a
point dorsally, size in decreasing order, 6, 7, 2, 3, 1, 4, 5, 5th under centre of eye.
Primary temporal roughly oblong, two posterior borders against upper and lower
secondary temporals and 7th supralabial, anterior two against the 2nd and 3rd post-
oculars, 4th subocular, and 6th supralabial. There are two secondary temporals. Body
scales begin behind the parietals, secondary temporals and 7th supralabial. The two
loreals are squarish quadrilaterals, slightly higher than long, the anterior between nasal,
frontonasal, prefrontal, posterior loreal and 1st and 2nd supralabials, the posterior
between the anterior loreal, prefrontal, 1st supraciliary, upper and lower preoculars and
2nd supralabial. The single well-defined chain which forms the upper palpebral series
and the irregular scales forming the lower palpebral series abut against the upper
preocular, which is also in contact with the 1st supraciliary, the upper accessory palpebral
(a small scale intercalated between the lower margins of the anterior two supraciliaries),
lower preocular and posterior loreal. The lower preocular is twice the size of the upper
and lies between it, the posterior loreal, 2nd (narrowly) and 3rd supralabials, 1st
subocular and the lower accessory palpebral (a small scale below the anterior end of
the palpebral series). The first three of the four suboculars are pentagonal with points
running down between the 38rd, 4th, 5th and 6th supralabials; the 4th is oblong and
between the 3rd subocular, 6th supralabial, primary temporal and 3rd postocular. All
are in contact with small scales forming that portion of the eyelid before, behind and
below the transparent disc. The postoculars are three small scales, the 1st about half
the size of the 3rd, which is about half that of the 2nd, lying between the 7th and 8th
supraciliaries, parietal, upper secondary temporal, primary temporal, 4th subocular and
a group of postpalpebrals; the Ist, which is anterior, lies against the junction of the
2nd and 3rd, the latter scale being antero-ventrally placed. Of the eight supraciliaries
the Ist and Sth are by far the largest with the 7th much larger than the remaining five,
the 1st is a triangular scale meeting the frontal at a point and lying between prefrontal,
Ist supraocular, 2nd supraciliary, upper preocular, and posterior loreal; the 7th is
roughly triangular, between the 4th supraocular, 6th and 8th supraciliaries, the last of
the upper palpebral chain and 1st postocular; the 8th lies in the angle between 4th
supraocular and parietal and is in contact with the 7th supraciliary and 2nd postocular.
There are four large supraoculars, the 2nd being the largest, the frontal is in contact
with the 1st and 2nd, the frontoparietal with the 2nd, 3rd and 4th, and the parietal with
the 4th. The transparent dise in the lower eyelid is undivided, convex, lens-like, about
half the length of the eye. The lower palpebral series forms a rim above, underneath
which there are about 12 small, nearly equidimensional scales. A few small scales
margin the dise before, and postero-ventrally there are three larger scales, one against
the 4th subocular and two against the 3rd. The large mental and postmental are
followed by three pairs of chin-shields, the 1st pair in contact, the 2nd separated by a
large, azygous scale, and the 3rd by three small, rhombic scales. There are seven infra-
labials, the first three small; all except the 1st are elongated, the 4th being the longest.

The ear, the centre of which is about. seven scales behind the mouth, slants obliquely
downwards and forwards as a shallow, ill-defined, scale-covered depression. It is seven
or eight scales long.

Seales are 24 at midbody, subequal, but slightly larger dorsally. Caudal scales
larger, 14 rows around tail at length of hindlimb behind vent. Two much enlarged
preanal scales with two smaller ones at each side. Scales from above vent to parietals, 66.

Body much elongated, the distance between the end of the snout and the forelimb
is contained twice in the distance between axilla and groin. Limbs small and weak,
especially the forelimbs, separated by about one and a half times the length of the hind-

66 VARIATION IN THE LIZARD HEMIERGIS DECRESIENSIS,

limb when adpressed. Lamellar formula for fingers, 5, 6, 6, about 10 tubercles on palm.
Lamellar formula for toes, 6, 8, 7, about 10 tubercles on sole, larger ones around margin
surrounding the smaller, inner ones.

Measurements of R.2191 are given with those of the other topotypes.

Ground colour of the specimen, preserved about 60 years ago, was apparently golden-
brown. Heavy dark brown dorsolateral lines begin on the 1st supraciliary, run through
the upper secondary temporal and along the body occupying one-half to two-thirds of the
4th row of scales. The upper third is left clear along most of the body. Behind the
vent the line changes to the 3rd row. The tail is regenerated, but the line probably
continues normally to the tip, becoming increasingly irregular. Four longitudinal lines
running between the dorsolaterals are dotted in outline, most dots being small. They
are missing on many scales. The inner pair continues on to the tail, where the lines
become more pronounced. The outer, more prominent, pair dies out just posterior to
the hindlimbs. Limbs dorsally, and tail ventrally heavily spotted. Head shields with
large, irregular, dark brown markings. Throat and neck scales with brown-dotted
margins giving a reticulated pattern. Belly practically immaculate.

Specimens of Hemiergis decresiensis decresiensis examined.
6 (R.2191-6, S. Aust. Mus.), Kangaroo Island, S. Aust., 1885. 1.

In this and the following three lists the number of specimens is given first, followed
by the eatalogue number, locality, collector’s name if available, date, and finally a
number giving the locality on the accompanying map (Fig. 1). Every specimen on the
tour lists has been examined by me.

KANGAROO
| 1 SLAND

KEY

H Ss D s
DECRESIENSIS
CONTINENTIS
TALBINGOENS!S
DAV ISI

Fig. 1.—Locality map. Only type localities, States and State capitals are shown by names.
Other localities are represented by numbers which tally with the last figure of each entry in the .
four lists of specimens examined.

Variation in Topotypes (excluding Provisional Neotype).—In the other five specimens
of the subspecies available (R.2192-6), the rostral agrees exactly with that of R.2191.
There is an apparent discrepancy in R.2196 where the rostral is quite typical, but the
frontonasal is abnormally large so that the area of the former scale visible from above
is equal to cnly one-quarter of that of the latter. The suture with the frontonasal
includes about four-fifths of the upper margin of the anterior loreal in four cases, only
one-third in R.2195. In R.2193 the frontonasal is in contact with the ist supraciliary

BY STEPHEN J. COPLAND. 67

-and is widely separated from the posterior loreal and 1st supraocular. In all specimens
the Ist supraocular and prefrontal are in contact. No specimen agrees with R.2191 in
having the scales meeting at a point. In four cases there is a short suture while in R.2195
it is more than half the length of the 1st supraocular. Two specimens have the pre-
frontals separated by 1/20 the width of the frontal, one by ~;, one by 4 and one by 2.
Average separation (including R.2191) is 0-108. The frontal shows only slight variation
in its proportions. It does not touch the Ist supraciliary. The frontoparietals, in spite
of their normally irregular shape individually, differ very little through the series. In
R.2196 the suture of the right frontoparietal with the frontal is slightly longer than that
ot the left with the same scale, which is normally three times as long. In all specimens
the oblique suture between the parietals slants backwards towards the left. In R.2196,
as in R.2191, there are no nuchals but a single large irregular scale on the left. R.2193
has an irregular pair. In R.2192 the post-parietal rows of scales are somewhat irregular
while in R.2194 and R.2195 they extend back in uniform rows. Four specimens have
typical supralabials, but in R.2194 there are eight on each side apparently through the
interpolation of a high, narrow scale before the normal 4th. Unlike R.2191, the posterior
loreal touches the 3rd supralabial in-all specimens. In R.2193, where the ist supraciliary
extends abnormally forward, the upper preocular is in contact with the 2nd supraciliary
only; in the other four specimens with the 1st only. The lower preocular meets the 3rd
supralabial only, while in R.2191 it also has a suture with the 2nd. Suboculars show
only slight irregularity. In R.2194 the 4th is grooved transversely. Postoculars are most
uniform. The ist supraciliary is separated from the frontal. In R.2193 the supraciliary
chain is broken after the 4th by the 2nd supraocular which, except for two nodules,
meets the upper palpebrals. Here the 1st supraciliary is moved forward until it is in
contact with the anterior loreal and frontonasal. All five specimens have two scales |
against the anterior margin of the 4th subocular, one of the two scales being in contact
with the 1st postocular. In R.2191 the upper of the two scales is small or missing. All
specimens have 24 scale rows at midbody. There are six lamellae under the 1st toes,
eight under the median, while seven, eight and nine lamellae under the 3rd are repre-
sented by two specimens each.

The five specimens agree fairly closely with R.2191 in the brown dorsal ground
colour and the heavy, black, single dorsolateral lines. The outer of the two pairs of
black, longitudinal lines between the dorsolaterals may be nearly continuous (R.2194-5)
or faintly outlined by dots, but it is always more pronounced than the inner pair except
on the tail. The lizards are somewhat bleached, but tails ana throats were apparently
heavily spotted ventrally and the underside of the bocy practically immaculate.

Generally all six topotypes (including R.2191) agree closely in scale and colour
characters. They all belong to Group B in the table of coleur patterns given with para-
types of H.d. talbingoensis.

Measurements of H.d. decresiensis in mm.

Number .. we aid ie R.2191 R.2192 R.2193 R.2194 R.2195 R.2196

Snout-vent e 5 at 45 40 43 33 26 40
Tail x ae Me ae 36+ 43+ 34+ 35 + 19+ 16+
Snout-ear Bie Se Ss 7 7 7 6 5 6
Snout-forelimb Me ae an 14 13 12 10 9 10
Axilla-groin a to Aa 28 25 27 20 15 25
Head, length oe -% Ws 7 i 6 a 6 5 6
Head, width Ay. ate yy: 6 5 6 4°5 4 4-5
Body, width “e ay 6 5 6 5 4-5 —
Forelimb, length .. ui Ae 6 6 6 4:5 4 5
Hindlimb, length . . 8 9 8 6 5p Me

Unfortunately Fitzinger’s type of Hemiergis decresiensis, which may still be in
existence in Hurope, has not been examined. On the other hand, there is no doubt as
to the identification of the specimens dealt with here. The type locality, Kangaroo Island,
is a comparativly small area where, in spite of considerable collecting, only a single

68 VARIATION IN THE LIZARD HEMIERGIS DECRESIENSIS,

species of lizard with three fingers and three toes is known to exist. This fact and the
close agreement of the South Australian Museum specimens R.2191-6 with original
descriptions leave no reasonable doubt that they are identical with the type. A British
Museum specimen mentioned by Gray (1845, pp. 87 and 272) and Boulenger (1887, p. 327)
from the Paris Museum may be a cotype. Gray (1845, p. 272) says: “Inhab. Kangaroo
Island. Mus. Paris, 1 specimen.” A description of a topotype of H.d. decresiensis is
given in full in the present paper. The holotype does not appear to have been mentioned
specifically since the original description nearly 120 years ago. It may have been lost
and in any case is now inaccessible.

For these reasons R.2191 in the South Australian Museum may for the present be
regarded as a provisional neotype.

It is realized that six is an unsatisfactorily small number of specimens on which
to establish standards, but there is no choice. Hemiergis d. decresiensis is doubtless
very rare on Kangaroo Island. The six specimens dealt with are the only ones in the
South Australian Museum and they were collected in 1885. The holotype and the
three specimens in the British Museum are the only others mentioned specifically in
literature, bringing the total known to 10. Collectors such as EH. le G. Troughton who
have visited Kangaroo Island failed to find specimens. It is probable that the position
cannot be permanently clarified until some herpetologist spends several weeks collecting
on the island with this one problem in view.

Boulenger (1887, p. 327) has given the only full description of H.d. decresiensis of
which I am aware. One of his two specimens was a topotype from Kangaroo Island,
the other Giinther’s type of Hemiergis polylepis. Both had 24 midbody scale rows.
Boulenger’s key (1887, p. 223), drawn up when only the Kangaroo Island and possibly
the South Australian races were known, includes H. tridactylum and the four subspecies
defined in this paper.

Proctor (1923, p. 81) expressly mentions the Kangaroo Island individual: “One
specimen of this rare skink from Flinders Island. The British Museum has only four
specimens, one of which is from Kangaroo Island.” Flinders Island is one of the
Investigator’s Group off the coast of South Australia. Parker (1926, p. 203) makes this
Flinders Island specimen (No. 1922.11.8.32 in the British Museum) the type of a new
species, Lygosoma (Rhodona) terdigitatum, chiefly on the grounds of the ‘‘much larger
frontal, which is broader than the supraocular region and longer than the interparietal
and frontoparietals together, a larger transparent disc and the absence of suboculars
separating the upper labials from the orbit”. It may be noticed also that the small
and widely separated prefrontals separate the individual generically from Hemiergis
while the 20 scales at midbody are at variance with the 24 or 26 scales of the Kangaroo
Island and South Australian mainland subspecies.

Waite (1927, p. 328) remarks on distribution: “This species occurs throughout
southern Australia and is found sparingly on Kangaroo Island and on Flinders Island
in Nuyts Archipelago.” The Flinders Island record is doubtless that of Proctor’s single
specimen of Lygosoma (Rhodona) terdigitatum Parker.

Gray (1831, p. 159) says of Tridactylus Decresiensis (now Hemiergis decresiensis)
and Tetradactylus Decresiensis (now H. peronii) “both were from the island of Décrés
and are viviparous”.

Gray’s short description (1831, p. 72) and the references of Fitzinger and Cuvier
have been given earlier.

Giinther (1867, p. 48) gives the following description of his Hemiergis polylepis:
“very similar to H. decresiensis, but with smaller scales, the body being surrounded by
26 series (in H. decresiensis by 18 or 20). Also the toes are more developed, the
anterior as well as the posterior being conspicuously longer than the eye. Posterior
frontals well developed. 72 scales in a series between the axils of the fore and hind
limbs. South Australia. 4 inches long’. He later (1875, p. 14) gave the habitat of
H. polylepis as “South Australia, Kangaroo Island”. It is apparent that polylepis must
be placed in the synonymy of H.d. decresiensis, which has the same number of scale
rows and was described from Kangaroo Island 41 years earlier.

BY STEPHEN J. COPLAND. 69

Duméril and Bibron (1839, p. 766), after describing the head scales in moderate
detail, give the following comparison of Hemiergis decresiensis with “le Tetradactyle de
Décrés”, i.e., Hemiergis peronti: “Sous le rapport des formes, si lV’on en excepte la
différence qui existe entre le nombre des doigts, cette espéce représente exactement le
Tetradactyle de Décrés dans tous ces détails. Son mode de coloration est aussi absolu-
ment le méme que celui de ce dernier Scincoidien. Longueur totale 10” 2”; Téte,
long. 9”; Cou, long. 9”; Tronc, long. 3”; Memb. antér., long. 8”’; Memb. postér., long. 1”;
Queue, long. 5” 4”. C’est également a la Nouvelle-Hollande et particuliérement dans
lile de Décrés que se trouve la présente espéce d’Hémiergis.”

The last paragraph does not enable one to determine whether the specimen described
came from Kangaroo Island or the mainland. Relative proportions are similar to those
of either H.d. decresiensis or H.d. continentis, except for greater lengths of head and
limbs. If from Kangaroo Island the specimen is 3 mm. longer in snout-vent measure-
ment than the largest of the topotypes in the South Australian Museum R.2191.

IV. HEMIERGIS DECRESIENSIS CONTINENTIS, Nn. subsp. PI. vi, fig. 2.

Diagnosis: Hemiergis decresiensis continentis is separated from the typical sub-
species H.d. decresiensis by larger size, greater body length and stouter habitus, also the
minor differences given in the tables and descriptions.

Holotype. No. R.2190, South Australian Museum; Myponga, S. Aust., 35.25 S,
138.20 E. H. M. Hale. No date.

Description of Holotype—HExcept for larger size and stouter habitus, R.2190 differs
little from the nominate subspecies. The rostral tends to be higher and more strongly
developed. Three-quarters of the upper margin of the anterior loreal is in contact with
the frontonasal. The frontal is narrowly in contact with the ist supraciliary, separating
the prefrontal and 1st supraocular. In all Kangaroo Island specimens the prefrontal
and ist supraocular meet. There is a pair of large, irregular nuchals. Supralabials
are deeper and more pronounced, especially the anterior three, than in the insular form.
The lower preocular touches the 2nd supralabial at a point as well as being widely in
contact with the 3rd. The postoculars are normal except that the 1st is not in contact
with the 2nd. One of two comparatively large scales against the anterior edge of
the 4th subocular is in contact with the Ist postocular. Formulae for lamellae under
fingers, 5, 6, 6; under toes, 6, 8, 8. Heavy black dorsolateral lines extend from the eye
along the tail. The head is flecked with black and there are three black patches on the
median line behind the neck. The whole of the throat and underside to the forelimbs
is reticulated with black, each scale having a dark margin thickest posteriorly. ‘The
underside is yellowish-white. The tail is longitudinally spotted with black. The pattern
is that of Grcup C in the table given under paratypes of H.d. talbingoensis.

Specimens examined and Locality Records of Hemiergis d. continentis.

(R.2190, S. Aust. Mus), Myponga, S: Aust. (H. M. Hale). No date. 2.
(R.2197-200, S. Aust. Mus.), between Gawler and Tanunda, S. Aust. No date. 3.
(R.2201-2, S. Aust. Mus.), South Australia. No date.
(R.8434-6, Aust. Mus.), Adelaide, S. Aust. (Pres. L. Harrison). Oct., 1924. 4.
(R.8531, Aust. Mus.), Victoria (Pres. Thomas Steele). Nov., 1924.

2 (D.1715-6, Nat. Mus.), Victoria. No date.

The number of specimens does not always tally with that of the tags, which are sometimes,
as in the Australian Museum series from Adelaide, attached to more than one individual.

Variation in Auxiliotypes—The single specimen from Myponga has been made the
holotype. The 17 individuals (two from the National Museum were returned before
detailed comparisons were made) now compared with the holotype cannot be considered
paratypes as they are from different localities. There is apparently no accepted term
for this class of specimens; paratypes sens. lat. being inadvisable because of possible
confusion with the strict paratypes, which in the case of lizards must at least be topo-
types. Because they are used to help define a new form the name auxiliotype (supporting
type) has been applied to them in this paper. Auxiliotypes are defined as types, not
holotypes or paratypes, used by an author to assist his original descriptions of new
species or subspecies. In the valuable paper of Davis and Lee (1944, p. 18), the metatype
“a specimen compared and declared conspecific with the true type by the original

Pon »

70 VARIATION IN THE LIZARD HEMIERGIS DECRESIENSIS,

author” could include this category, but “both homoeotype and metatype are taken as
implying that the comparison post-dated the publication of the original description” and
exactitude is again sacrificed.

The rostral in mainland specimens tends to be higher and more developed than in
the nominate subspecies from Kangaroo Island, although in eight of the 17 specimens
there is no great difference. In R.2202 and one of R.8435 the area visible from above
is equal to two-thirds that of the frontonasal. Width of the suture between rostral and
frontonasal to the width of the frontal varies from one-third to one-eighth. Between
one-third and four-fifths of the upper margin of the anterior loreal is in contact with
the frontonasal. In five specimens the prefrontal meets the 1st supraocular in a point,
in nine in a short suture, while in three (which agree with the holotype) the two scales
are separated by the junction of the frontal and 1st supraciliary. Frontoparietals are
regular in 15 specimens. In R.2199, instead of the suture of the right frontoparietal
with the frontal being only about a third the length of the contact of the left fronto-
parietal with the same scale as normally, the position is reversed. In one of the R.8434
specimens the scales are abnormal and the left is cut off widely from contact with the
frontal by the right frontoparietal with which the anterior third of the left has evidently
fused. In 15 cases the suture between the parietals has the normal slant backwards
towards the left, but in two the direction is reversed towards the right. In R.2197,
R.2200 and one of the R.8435 series the parietal meets the 4th supraocular at little more
than a point. No specimen has nuchals, but there is a large irregular scale on the right
in R.8531 and two irregular scales on the left in R.2202. Post-parietal scales are regular
in nine specimens, irregular in six. Supralabials are deep and sharply cut as in the
holotype in 14 specimens. In three they are lower, being somewhat similar to those of
H.d. decresiensis. In R.2199 the supralabials are normal on the left side, but reduced to
six on the right, apparently by the fusion of the third and fourth scales. The posterior
loreal is narrowly in contact with the 3rd supralabial in six cases, three of which are at
a point. The lower preocular touches only the third of the supralabials in seven
specimens. In R.2199 the Ist and 2nd suboculars have fused on each side and on the
left the 5th supralabial reaches nearly to the margin of the eye behind the fused scales.
In R.2201, where the 3rd postocular is much enlarged, the usually comparatively large
and oblong 4th subocular is small and rounded. There is a tendency in all mainland
specimens for the 1st postocular to be moved forward and reduced in size. The 1st
postocular is not in contact with the 2nd in six individuals. In R.2201 the first scale
is nearly equal in size to the second. In R.2199, which is irregular in other scale.

Measurements of H.d. continentis in mm.

Head. Length.
Number. Snout- Snout- Axilla- Body. Fore- Hind-
Vent. Tail. Forelimb. Groin. Width. Length. Width. limb. limb.
R.2190 ee 52 58 15 35 6 8 8 5 8
R.2197 me 46 5333 12 31 4-5 6 5 6 8
R.2198 ah 43 31+ 12 27 5 6-5 6 6 8
R.2199 ae 44 18+ 12 28 5 6 5 5-5 8
R.2200 He 42 49 12 28 4-5 6 5 6 8
R.2201 46 49+ 14 31 5-5 a 7 6°5 8
R.2202 50 41+ 15 31 5-5 7 6 5 7
R.8434 47 38l+ 13 Sil 5:5 6°5 6 6 8
46 22+ 1133 32 6 iG 7 6 9
46 50+ 12 30 6 6°5 6:5 6:5 9
R.8435 54 64 14 36 6 6-5 i 6:5 9
47 52+ 14 33 6 6°5 6-5 7 8
53 54+ 13 35 6:5 7 a 6°5 8
R.8436 nS 49 28+ 12 31 5 7 6-5 5:5 8:5
48 38 + 13 30 5-5 6-5 5-5 Oo) GOD
52 60 14 32 Bobs 7 6-5 6 8
52 29+ 14 35 5°5 6-5 6 6-5 8
R.8531 “te 46 344+ 13 82 5 7 6 5 8
D.1715 2 44 — 1838) 25 —- — — — 8
— — — — 7

D.1716 56 33 = 11 20

|
|

BY STEPHEN J. COPLAND. 71

characters, the 2nd and 3rd postoculars have fused on the right side and partly on the
left. In R.2199 the anterior two supraciliaries have partly fused. Six specimens agree
with R.2190 in having one of the two large scales against the 4th subocular in contact
with the 1st postocular. Hight have two scales against the subocular but no contact
with the Ist postocular. In three cases there is a single scale. Colour and markings
of 14 of the 19 specimens agree essentially with those of H.d. decresiensis (Group B),
but three specimens from between Gawler and Tanunda agree with the holotype in
having both pairs of dorsal lines vestigial (Group C), while in two from Adelaide both
pairs are practically continuous and about equally prominent (Group A).

Main reasons for the separation of the mainland race from the closely allied typical
form are: greater body length (average 47 mm. against 37:83 mm.; no specimen of
H.d. decresiensis exceeds 45 mm., while 40 per cent. of H.d. continentis are in excess of
this figure); stouter habitus, A.d. continentis averaging practically a millimetre more
in width of body (6:25 mm. against 5-30 mm.); more robust rostral and supralabials;
' tendencies for the lst postocular to be moved forward and reduced in size, and for
frontal to meet ist supraciliary: besides small differences which give an individual
aspect to the two subspecies. In the author’s opinion separation of the island and
continental races is valid and agrees with the views of Mayr (1942), the mammalogist
Glover M. Allen (1938 and 1940), and the herpetologists Mertens (1931) and Pope (1935).
Mayr’s treatment of the whistler Myiolestes megarhynchus (p. 43) and the Asiatic bulbul
Microscelis leucocephalus (p. 83) and in Allen’s work of most subspecies, especially
those of the genera Ochotona (1938, p. 525), Callosciurus (1940, p. 626), and Rattus (1940,
p. 983) may be cited. It should be noted here that the differences between H.d.
decresiensis and H.d. continentis are much slighter than those between either of these
races and H.d. talbingoensis or H.d. davisi, or again the differences by which the two
latter forms are distinguished from one another.

Lonnberg and Andersson (1913, p. 9) record two specimens from Adelaide collected
on October 15, 1911.

Loveridge (1934, p. 368) gives brief notes on the four specimens of H. decresiensis
in the Museum of Comparative Zoology, Cambridge, Massachusetts. The lizards (M.C.Z.
33155-8) were collected by W. M. Wheeler at Mt. Lofty, S. Aust., in 1931. Midbody scale
rows number 24 or 26, and the largest skink measures 103 (49 + 54) mm. Both
Loveridge’s and Lonnberg and Andersson’s specimens may be taken as belonging to the
subspecies continentis.

V. HEMIERGIS DECRESIENSIS TALBINGOENSIS, nh. Subsp. Pl. vi, fig. 3.

Diagnosis: Hemiergis decresiensis talbingoensis is separated from the typical sub-
species H.d. decresiensis by the lower number of midbody scale rows (22 in the holotype
against 24: average for 94 specimens examined 21.74), lower number of lamellae beneath
mid-toe (seven in the holotype against eight: average for 96 specimens examined 7.18).
There are also differences in scalation, colour and size, these three points being dealt with
in the tables and descriptions.

Holotype. Author’s Collection, No. 2081; Talbingo, N.S.W., 35.34 S, 148.20 E. Alt. c.
1300 feet. 3.xii.1943.

Description of Holotype.—Rostral moderately high, area visible from above equal
to about half that of the frontonasal; long, concave sutures with the nasals and approxi-
mately vertical ones with the ist supralabials; the nearly straight junction with the
frontonasal is about a quarter the width of the frontal. Nasals large, not in contact,
roughly quadrangular, long convex sutures with the rostral and frontonasal and shorter
straight ones with 1st supralabial and anterior loreal. Oval nostril approximately central,
no sign of a groove from it to divide the nasal scale. No supranasals. Frontonasal large,
equal in area to the frontal, in contact with frontal, prefrontals, nasals, rostral, and
anterior half of upper margin of anterior loreal. Prefrontals large, well developed,
separated by a tenth the width of the frontal, sutures long and fairly straight with
frontonasal, sinuous with frontal, concave with 1st supraciliary, nearly straight and
horizontal with the two loreals, very short with 1st supraocular. Frontal kite-shaped,
rounded in front against the frontonasal and behind between the frontoparietals, pointed

72 VARIATION IN THE LIZARD HEMIERGIS DECRESIENSIS,

laterally where frontal, prefrontal, 1st supraciliary and 1st supraocular meet in a point.
Long, straight posterolateral sides against the 1st and 2nd supraoculars, concave antero-
lateral sutures with prefrontals. Frontoparietals paired, large, subequal in size with
interparietal, left scale a rough crescent twice as long as wide, inner border long and
convex against parietal, interparietal, and its fellow, outer border slightly concave
against the 2nd, 3rd and 4th supraoculars, indented anteriorly against posterior end of
frontal. Right frontoparietal in contact with the same scales, but more squat and
pointed mediad between interparietal and left frontoparietal, suture with frontal only
about one-third the length of the contact of left frontoparietal with the same scale. Inter-
parietal kite-shaped, subequal in size with the frontal, rounded behind, pointed in front
and at sides, sutures long and straight with parietals, shorter with frontoparietals,
concave. with left, sinuous with right; a rounded milky area in the midline one-third
of the length of the scale from the posterior end covers the pineal foramen. Parietals
are the largest head shields, forming a V-shaped pair enclosing the interparietal, irregu-
larly pentagonal, meeting behind the interparietal in an oblique suture slanting back-
wards towards the left; other sutures are straight and long with interparietal, short
and slightly concave with frontoparietals, very short with 4th supraocular, 8th supra-
ciliary and 2nd postocular, long and straight (about parallel to the one with the inter-
parietal) with upper secondary temporal, left scale in contact with two rows of dorsal
seales, the right with three, there being no nuchals. Seven supralabials, the anterior
three roughly quadrilateral, their upper margins forming a straight line with the nasal,
loreals and lower preocular, posterior four pentagonal with lower margins horizontal,
anterior and posterior sutures vertical, the other two sides meeting in a point dorsally.
Size in decreasing order, 7=6, 3, 2, 1, 4, 5; 5th under centre of eye. Primary temporal
roughly oblong, two posterior borders against upper and lower secondary temporals and
7th supralabial, anterior borders with 6th supralabial, 4th subocular, and 2nd and 3rd
postoculars. There are two secondary temporals (the upper being much the larger) and
a tertiary temporal. Body scales begin behind the parietals, upper secondary temporal,
tertiary temporal, and 7th supralabial. The two loreals are oblong, slightly higher than
wide, the anterior. between the nasal, frontonasal, prefrontal, posterior loreal and ist
and 2nd supralabials, the posterior between the anterior loreal, prefrontal, Ist supra-
ciliary, upper and lower preoculars and 2nd supralabial. The upper and lower palpebral
series abut against the upper preocular, which is in contact with the list supraciliary,
the upper accessory palpebral (a small scale intercalated between the lower margins of
the ist and 2nd supraciliaries), the posterior loreal, lower preocular and lower accessory -
palpebral. The lower preocular is twice the size of the upper and lies between the upper
preocular, posterior loreal, 3rd supralabial (meeting the 2nd in a mere point), Ist
subocular and lower accessory palpebral. The first three of the four suboculars are
pentagonal with downwardly-directed points lying between the supralabials, the posterior
seale is roughly oblong and lies between the primary temporal, 3rd postocular, 6th
supralabial and 3rd subocular. The postoculars are three small seales, the 1st one-third
to half the size of the 3rd, which is smaller than the 2nd, lying between the 4th subocular,
primary temporal, upper secondary temporal, parietal, 7th and 8th supraciliaries and
the small scales behind the eye. The 2nd and 3rd lying immediately against the anterior
border ot the primary temporal separate that scale from the ist postocular. Of the
eight supraciliaries, the 1st, 7th and 8th are by far the largest, the Ist a large triangular
scale between the prefrontal, lst supraocular, posterior loreal, upper preocular, upper
accessory palpebral and 2nd supraciliary; the lozenge-shaped 7th and 8th lie between
the 4th supraocular, 6th supraciliary, the last of the upper and lower palpebral chains, 1st
and 2nd postoculars, and parietal. There are four large supraoculars, the 3rd being
the largest; the frontal is in contact with the 1st and 2nd; the frontoparietal with the
2nd, 3rd and 4th; and parietal with the 4th. The transparent dise in the lower eyelid
is undivided, convex, and lens-like, slightly longer than half the length of the eye
aperture and larger than the pupil. The large mental and postmental are followed by
three pairs of chin-shields, the 1st pair in contact, the 2nd separated by a large azygous
shield, and the 3rd separated by three small scales. Infralabials are six if a small
scale behind the 6th is not counted; all except the Ist are elongated, the 5th being

BY STEPHEN J. COPLAND. 73

twice as long as wide. The ear about six scales behind the mouth runs obliquely down-
wards and forwards as a shallow, scale-covered, slit-like depression. There are about
four scales on its anterior border.

Seales 22 at midbody, subequal, but slightly larger dorsally. Tail scales larger,
15 rows around tail at length of hindlimb behind vent. Four preanal scales, inner
pair much larger than outer. Scales from above vent to parietals, 76. Body much
elongated. The distance between the end of the snout and the forelimb is contained
3-1 times in the distance between axilla and groin. Limbs small and weak, especially
the forelimbs, separated by nearly three times the length of the hindlimb when
adpressed. Lamellar formula for fingers, 5, 6, 5. Ten enlarged rounded tubercles on
palm. Lamellar formula for toes, 5, 7, 8. Five flattened tubercles on sole surround five
larger, more prominent ones.

Measurements of the holotype are given with those of the paratypes.

Colour. Ground colour of head, body and tail is light chocolate. A black dorso-
Jateral line begins just behind the eye in the black postoculars, runs through the upper
secondary temporal, which is half black, and along the third row of dorsal scales. The
line, half a scale wide, runs through the centre of the scales, leaving dorsal and ventral
quarters untouched. Just behind the vent, the line switches to the second row from
the midline and continues to the tip of the tail, being discontinuous along the posterior
half. Two black, longitudinal lines begin behind the parietals and follow the first row
of dorsal scales on each side. They are about a quarter scale wide and not quite
continuous, the black pigment normally occupying only the anterior three-quarters of
each scale. The lines become more discontinuous on the tail where they are very
irregular for the distal half. Traces of two other black lines on the second rows of
scales extend along seven or eight scales behind the parietals as dots and then as very
occasional flecks to the end of the body. Dark brown to black patches occur on all
head shields. Under high power they are seen to consist of hundreds of deep brown
dots. Sides of the body are of the ground colour becoming gradually lighter ventrally.
Small black dots tend to form ill-defined lines along each lateral scale row. Dark
markings crowd together on the sides of the tail. The underside is immaculate for
six scales from the forelimbs to near the vent where the clear area narrows to four
scales’ width. The throat and neck nearly to the forelimbs appear reticulated with
dark brown caused by the posterior third of each scale being beset with scores of tiny
brown dots. Mental, postmental, and chin-shields are especially dark. Scales behind °
vent are white with perhaps a quarter black, but each scale becomes progressively
darker posteriorly until the distal half of the tail is crowded with dark brown to black
spots. The pattern is that of Group H.

Variation in Paratypes.—The rostral is remarkably uniform in all 52 paratypes. The
only variation is in the length of the suture with the frontonasal compared with the
width of the frontal. Twelve specimens agree with the type, about 4 the width, but in
40 it is much less (29, 4; 10, 7,; while the nasals are nearly in contact in one). Nasals,
which are regular, are separated from the 2nd supralabial by a quarter to a third the
width of the 1st, but approach contact in the holotype and only one paratype. Relation
of the frontonasal to surrounding scales varies very little, most specimens agreeing with
the type in being in contact with about half the upper margin of the anterior loreal.
In 19 cases the length of the contact is about two-thirds, in seven about four-fifths, while
in A.C. 2008 and A.C. 2011 it is in contact with the entire upper margin. In A.C. 2059
the lower portion of the scale is cut off on the right to form a small triangular scale.
In 22 cases the prefrontals meet the 1st supraocular in a short suture. The two scales
are only separated in four specimens. In A.C. 2082 they are separated on one side but
meet on the other. The remainder have the scales meeting at a point or suture so
short that it may be classed as such. In A.C. 1687 the lower two-fifths of the right
scale is divided off by a transverse suture. Separation of the prefrontals to the width
Ciechemirontaly Ie see ae ib ese et te Te. Ad ei ao) (as! holotype); 6, 1/20) 5) an!
contact. The frontal is normally half as long again as wide, an occasional individual
being nearly twice as long as wide, while a few are nearly as wide as long. Most
frontal scales are smooth or have a few minute pits. A few, such as A.C. 2047, are

74 VARIATION IN THE LIZARD HEMIERGIS DECRESIENSIS,

heavily pitted. These remarks on pitting apply to most of the head shields. Lateral
angles of the frontoparietals sometimes approach closely but never touch the 8th supra-
ciliary. In A.C. 2083 the right is partly fused to the 2nd supraocular. A.C. 2013 has a
process running back between the interparietal and parietal. The interparietal varies
in its degree of elongation, sometimes being very squat. In A.C. 2014 it is abnormal
and only about half the size of the frontal, while in A.C. 2044 it is very well developed
and larger than the frontal. In A.C. 2087 the scale is unusually rounded with a
process between the right frontoparietal and parietal. The number of dorsal scales
touching the parietals varies from two to four on each side, but the number is normally
larger on the left, where in one case (A.C. 2085) it meets a fifth at a point. The number
of scales in contact depends generally on the presence of irregular nuchals or large
unpaired post-parietal scales. A.C. 2087 has the right parietal divided into three irregular
seales, two large with a smaller one between them. In only two specimens does the
suture between the parietals slope backwards towards the right. Thirty-four specimens
agree with the holotype in having no nuchals, the body scales running back sometimes
quite regularly, but often the anterior one or two rows are irregular. In seven cases
there is an irregular pair of nuchals, in four there is a large unpaired scale on the left,
in five on the right, while one specimen has two large unpaired scales on the right and
another two on the left. The 1st supralabial about as high as long in the type, is most
often longer than high. The 7th and 6th scales are always the largest but the others
may be practically equal in size and frequently the Ist ranks after the 7th and 6th.
Generally the order in size agrees with that of the holotype. A.C. 2015 has eight supra-
labials on each side due to an additional large scale being intercalated between the normal
3rd and 4th; the Ist scale remains normal but the next three are narrow and high. Out-
- lines of the remainder are as usual. The 6th (normal 5th) is under the centre of the
eye. Of interest in showing how abnormalities occur symmetrically, a small scale occurs
on the right and left sides of A.C. 2039 between the 2nd and 3rd supralabials, posterior
loreals and lower preoculars, yet in A.C. 2041 and A.C. 2058 a similar scale occurs on
the right side only. A.C. 2031 has the 5th and 6th scales irregular and fused on the
right side. In A.C. 2056 the right primary temporal is fused with the 2nd postocular
and thus brought into contact with the 1st postocular and Sth supraciliary. In A.C.
2059 where the upper secondary temporal is divided and abnormally developed on the
right side, the primary temporal is cut off by it from the lower secondary temporal. The
upper secondary temporal is divided in A.C. 2012. The posterior loreal meets the 3rd
as well as the 2nd supralabial in 10 cases. A.C. 2033 has-the upper and lower preoculars .
partly fused. Twenty individuals have the lower preocular meeting the 2nd supralabial
in a short suture as against a mere point in the type. Suboculars are generally most
regular. There are five on each side in A.C. 2015 and A.C. 2045. Three scales replace
the normal anterior two in A.C. 2020 on the right side. In one of R.12084 the first is
small and the second divided on each side. A.C. 2031 has the 3rd scale on the right
side broken down into an aggregate of small scales. A.C. 2054 has the 2nd and 3rd
separated by a process of the 5th supralabial. The postoculars vary greatly in size and
shape, being oblong or lens-shaped. The 2nd and 3rd are often comparatively large with
a long suture between them. The 1st is always very small. In A.C. 2012 the right 2nd
postocular is separated from the parietal and in A.C. 2041 the left. Supraciliaries are
most regular in number, order and size for such attenuated chains. A.C. 2016 has the
6th scale on the right divided to give nine supraciliaries on that side. Supraoculars
vary little in their relationships with other scales. The only noteworthy variation from
the holotype is that the 2nd scale is larger than the 3rd in about 80 per cent. of
specimens. A.C. 2014 has the 2nd much larger than the 3rd on each side with an
abnormal lenticular scale, truncated against the supraciliaries, on the right side.
A.C. 2020 has a large semi-circular seale cut off from the anterior margin of the post-
mental. The normally small scale behind the 6th infralabial is often large and assumes
the status of a 7th infralabial. Again the 6th is often large and unmistakably the
terminal scale. In the occasional case where the posterior border of the 7th infralabial
is in contact with the 7th supralabial, there is a small 8th scale behind it.

BY STEPHEN J. COPLAND.

-
or

It was found convenient to arrange colour patterns in groups to facilitate analysis
and recording. All 258 specimens of the four subspecies were lumped together and then
sorted into separate piles mainly on the basis of the relative prominence of the dorsal
and dorsolateral lines. All lizards fell into 18 groups with only a small percentage of
borderline cases. Sixteen and possibly the other two of the 18 groups indicate
geographical grouping of colour patterns; the most conspicuous being Group J, which

contains 65 individuals.
from the Northern Tableland.

All but one are H.d. davisi, including 27 of the 29 specimens
Full importance probably cannot be attached to Groups

Q and R, which are almost uniform in colouration, because lack of pigment may occur

independently from various causes.

Group. G, with faintly dotted dorsolateral and

dorsal longitudinal lines, also appears to be composite.

The table for all four

races iS Summarized here.

Abbreviations are: d.l., dorso-

lateral lines, 1 for inner and 2 for outer dorsal lines; H.d. decr. for H.d. décresiensis;
H.d.c. for H.d. continentis; H.d.t. for H.d. talbingoensis; and H.d.d. for H.d. davisi.

Table of Colour Groups of the Four Races of H. decresiensis.

Group A. d.l. wide, heavy and black. H.d.ec., 2 spms.: Adelaide.
- 1 and 2 practically continuous, about
equal. :
Group B. 4.1. wide, heavy and black. H.d. decr., 6 spms.: Kangaroo I.
1 traces. 2 practically continuous or H.d.c., 14 spms.: 8 Adelaide, 3 Victoria, 2
more prominent than 1. South Australia, 1 between Gawler and
Tanunda.
Group C. d.l. wide, heavy and black as in A H.d.c., 4 spms.: 38 between Gawler and
and B. Tanunda, 1 Myponga.
1 and 2 vestigial. edit. so. Spmsic is) Cullerin’ 125 Collectorsel
Talbingo.
H.d.d., 3 spms.: 1 nr. Abercrombie River, 1
nr. Porter’s Retreat, 1 Hampton.
Group D. d.l. distinct, but not very heavy and H.d.t., 19 spms.: 11 Talbingo, 6 Mount Kos-
inclined to be double. ciusko, 1 Marulan, 1 Goulburn.
1 and 2 lightly dotted.
Groupee Much, as DD) but ad: and 2) more H.d.t., 18 spms.: 11 Talbingo, 5 Mount Kos-
prominent. sciusko, 1 Cullerin, 1 Collector.
Group F. ad.Jl. much as in D and E. H.d.t., 11 spms.: 6 Talbingo, 2 Mount Kos-
1 about equal to d.l., 2 faint. ciusko, 2 Victoria, 1 Cullerin.
Group G. d.l. 1 and 2 dotted, fairly faint. H.d.d., 8 spms.: 1 Curraweela, 1 Duckmaloi
River, 1 Oberon, 1 Hampton, 1 Black
Springs, 1 Tarana, 1 Hartley Vale, 1
Capertee.
H.d.t., 7 spms.: 6 Talbingo, 1 Adaminaby.
Group H. d.l. very heavy. edste wus aSspMmssenaiecalbim Ons. Cullerinvas2
1 continuous or nearly so. 2 faintly Collector, 1 Goulburn.
outlined in dots or missing.
Group I. 4d.l. light. H.d.d., 4 spms.: 3 Hartley, 1 Lett River,-nr.
1 very distinct and continuous. 2 Hartley.
traces.
Group J. d.J. continuous or nearly so, about H.d.d., 64 spms.: 12 Salisbury, 8 Oberon, 6

Group K.

quarter scale wide.
las dl. 2 faintly dotted or missing.
Four-lined pattern.

As J, but 2 approaching that of L.

“Southern Australia’, 6 Hartley, 6 Armi-

dale, 4 Llangothlin, 4 Bendemeer, 4

Black Springs, 4 Tarana, 2 Rydal, 2

Little Hartley, 1 Lett River, nr. Hartley,

1 Duckmaloi River, 1 Hampton, 1 Forest

Reefs, 1 Liverpool Plains, 1 Lithgow.
H.d.t., 1 spm.: Mount Kosciusko.

Eld:ds; 2 spms::
tralia’.

1 Rydal, 1 “Southern Aus-

u

76 VARIATION IN THE LIZARD HEMIERGIS DECRESIENSIS,

Group L. dl. 1 and 2 continuous or practically
continuous black lines.
Six-lined pattern.

Group M. As L, but d.l. heavier.

Group N. dl. 1 and 2 composed of uniform
dots, each occupying about a
sixth of a scale.

Group O. 4.1. 1 and 2 dotted lines, which may
be double and scarcely distinct
from the lateral lines.

Group P. As O, but lines lighter.

Group Q. Close to R, but flecks and suggestions
of markings.

Group R. Uniform dorsal ground colour, with-
out markings.

—

H.d.d., 6 spms.: 1 Bendemeer, 1 Laggan,
Curraweela, 1 Hampton, 1 Armidale, 1
Bundanoon.

H.d.t., 1 spm.: Talbingo.

H.d.d., 5 spms.: 2 Curraweela, 2 Hampton,
1 Black Springs. é

H.d.d., 8 spms.: 2 9 m. N. of Abercrombie
River, 2 Curraweela, 1 5 m. N. of Aber-
crombie River, 1 Porter’s Retreat, 1
Duckmaloi River, 1 Hampton.

H.d.d., 17 spms.: 5 5 m. N. of Abercrombie
River, 4 11 m. N. of Abercrombie River,
2 Curraweela, 2 Porter’s Retreat, 1
Oberon, 1 Hampton, 1 Black Springs, 1
Tarana.

H.d.d., 11 spms.: 4 Black Springs, 2 Duck-
maloi River, 1 Hampton, 1 Hartley, 1
10 m. from Jenolan Caves, 1 9 m. N. of
Abercrombie River, 1 5 m. N. of Aber-
erombie River.

H.d.t., 16 spms.: 8 Talbingo, 5 nr. Adaminaby,
2 Mount Kosciusko, 1 Cullerin.

H.d.d., 7 spms.; 2 Hampton, 2 5 m. N. of
Abercrombie River, 1 Hartley, 1 10 m.
from Jenolan Caves, 1 Tarana.

EGeee spms.: 2 Talbingo, 1 nr. Adaminaby,
1 Mount Kosciusko.
H.d.d., 1 spm.: Hampton.

Summary of Table.

Group. ALOT BS. (CS DE Wha wine Gy MES iTS sed heey oli VS ON age soe) aha
Hd. decresiensis aS -. — 6— — — _—
H.d. continentis 214 4—- — — — — — — —-
H.d. talbingoensis ee 2 ae 16 OST Be abl ale 1 1 iG) we!
H.d. davisi a 3k se SS SS 8 PS 8 a 2 6 ea Seely, dee eee

All Talbingo specimens in the table are paratypes. Twenty-two of the 52 specimens
fall in Groups D and E (11 each) with distinct but not very heavy dorsolaterals, and
Six specimens occur in each of the three
following groups, one each in C and L and 10 in the nearly uniform Groups Q and R.

light or slightly more prominent dorsals.

Measurements of the Holotype and Ten Paratypes of H.d. talbingoensis in mm. (Specimens with Complete Tails Chosen.)

Number. Snout- Tail. Snout-
Vent. Forelimb.

A.C.2081 ae Lost GO. 74 14
A.C.1687 ee bt 43 60 13
A.C.2008 as is 55 69 14
A.C.2020 Be ae 37 46 11:5
A.C.2030 ae oe 53 65 13
A.C.2033 Bs ie 53 67 14
A.C.2043 Be 3 35 43 10
A.C.2046 te ns 50 66 15
A.C.2054 se she 59 71 15
A.C.2055 Aus Pre 54 67 13
A.C.2083 Ae ats 50 70 14°5

Length.
Axilla- Head. Fore- Hind-
Groin. Length. Width. limb. limb.
43 7:5 6:5 6 9
29 6-5 5 5e5) 8
37 7 6 6-5 8
23 6 4-5 4-5 7
35 7 6 6 8
35 7 5:5 6 8
22 6 4:5 Deo 6-5
32 (5 6 6 9
42 8 6 6 8-5
36 7 5:5 5-5 8
32 7 6 6 8-5

BY STEPHEN J. COPLAND. 77

. Specimens examined and Locality Records of Hemiergis d. talbingoensis.
1 (A.C. 1515) 4 m. W: of Marulan, N.S.W., 18.1.19438. 5
1 (A.C. 1661) 8:3 m. N. of Adaminaby, N.S.W., 27.1.1943. 6.
1 (A.C. 1687) 1-8 m. SE. of Talbingo, N.S.W., 28.1.1943. 7.
9 (A.C. 1964—6, 1973-7, 1979) 2:5 m. W. of Cullerin, N.S.W., 20-22.xi.1943. 8.
0 (A.C. 2008-20, 2030-48, 2053-60, 2080-9) within 2 m. of Talbingo, N.S.W., 28.xi.-3.xii.1943. 7.
6 (A.C. 2134-9) 2:3 m. N. of Adaminaby, N.S.W., 6.xii.1943. 6.
5 (A.C. 2159-63) 2m. N. of Collector, N.S.W., 9.xii.1943. 9. ©
2
2
3
2
2
2

isi}

(D 2459-60, Nat. Mus.) Victoria. No date.
(R 469-73, 475-81, Aust. Mus.) Mt. Kosciusko, N.S.W., 3,000 ft. (R. Helms), May, 1889. 10.
(R 530, 532-3, Aust. Mus.) Mt. Kosciusko, N.S.W., 5,000 ft. (R. Helms), May, 1889. 10.
(R 531, 534, Aust. Mus.) Mt. Kosciusko, N.S.W. (R. Helms), May, 1889. 10.
(R 2393-4, Aust. Mus.) Goulburn, N.S.W. (J. A. Thorpe), Oct., 1898. 11.
(R 12084, Aust. Mus.) Talbingo, N.S.W. (J. C. Wiburd), April, 1937. 7.
In this and the following list all specimens in the author’s collection (prefixed A.C.)
have been collected by him unless a collector’s name is given.

Variation in Auxiliotypes—Twenty-five of the 43 auxiliotypes examined agree with
the type in having the area of the rostral visible from above equal to about half that of
the frontonasal, the area in the remainder being about a third; sutures of the rostral
with the frontonasal equal a quarter that of the frontal, as in the type, in 16 cases, 4 in 15,
qo in six, and nearly 4 in six. In R 5338, A.C. 1976 and A.C. 2161 the nasal approaches
contact with the second supralabial. In R480 grooves extend across the nasal scales
trom the nostrils. In A.C. 2163 a peculiar, narrow, bow-shaped scale running between
the anterior loreals margins the entire anterior border of the frontonasal, separating
it from the rostral and nasals. In this specimen the frontonasal is considerably larger
than the frontal. The posterior part of the frontonasal in R476 is cut off to form a
large, lens-shaped scale; in A.C. 1964 the scale is partly divided on the right side.’ The
frontonasal is slightly smaller than the frontal in most specimens, but in A.C. 1966,
A.C. 1976, A.C. 1977 and A.C. 2161 it is considerably larger. Nineteen specimens have:
the frontonasal in contact with about half the upper margin of the anterior loreal,
18 with about 2, five with #, and R481 with practically the entire border. The suture
of the frontonasal with the frontal (equal to the separation of the prefrontals) is very
narrow in the nine Cullerin specimens (where four specimens have the prefrontals in
contact or nearly so, two separated by 1/20 the width of the frontal and three by ;4,
average separation 0:04). Figures for other specimens are Marulan 4, Goulburn } and 3,
Victoria two nearly in contact, Adaminaby (two in contact or nearly so, one 75, one 3,
two 3, one 4, average 0-16), Collector (one in contact, one 1/20, two 7, one 4, average
0-09), Mt. Kosciusko (one in contact, one 74, three 4, two %, two 2, five 4, two 4, average
0-20). In five specimens the prefrontals are separated from the 1st supraocular by the
meeting of the ist supraciliary and frontal. Contact of the prefrontals with the 1st
supraocular varies from a fairly wide suture to little more than a point. Normally one
and a half times as long as wide, the frontal is equidimensional in three specimens
and twice as long as wide in six. A.C. 1661 and A.C. 2136 have the frontal fused with a
small scale cut off from the left frontoparietal. The normally elongated left fronto-
parietals are practically equidimensional in a few cases. The interparietal is occasionally
as wide as long and in A.C. 1515, A.C. 2138, A.C. 2139 and A.C. 2163 it is wider than long,
larger than the frontal, and approaching twice the size of a frontoparietal. The parietals
always meet but the suture is sometimes very short, in nine cases (R472, R475, R 476,
R 479, A.C. 1966, A.C. 1975, A.C. 1979, A.C. 2134, A.C. 2160) the suture slopes backwards
towards the right. Most specimens agree with the type in having no nuchals, but about
a third have an irregular pair or a large unpaired scale on either the right or left side.
Normally the 1st supralabial is about square, but it is elongated or taller than broad in a
few lizards. It is often the third largest of the seven scales. The 6th supralabial is
occasionally larger than the 7th. R532 and A.C. 2137 have only six supralabials while
A.C. 1965 has eight, three scales replacing the normal 2nd and 3rd. On the left side
separation is not quite complete. A.C. 1966 has the primary temporal reduced in size
and roughly lens-shaped. In A.C. 1974 the scale is separated from the 4th subocular. The
right primary temporal of R534 is partly fused with the 7th supralabial. The tertiary
temporal is very variable, generally tall and lens-shaped; it is occasionally indistinguish-

L

1

78 VARIATION IN THE LIZARD HEMIERGIS DECRESIENSIS,

able from the body scales except by its position. The posterior loreal, which may taper
ventrally to a very short contact with the 2nd supralabial, is in contact with the 3rd as
well as the 2nd supralabial in five specimens and meets the scale at a point in another
three cases. The anterior loreal is divided transversely in A.C. 1964. In 20 specimens
the lower preocular meets the 2nd supralabial in a short suture. A.C. 1965 has five
suboculars on each side and A.C. 1964 five on the right only. In R534 the 2nd and
3rd suboculars are separated widely on the right side by the abnormally developed 5th
supralabial. A.C. 1661 has the fourth abnormally large and in contact with the 7th
supralabials as has R 2393 on the right side. The 2nd postocular is usually twice the
size of the 3rd and four times that of the Ist, the Ist being small and lenticular, the
2nd large and lenticular, and the 3rd practically square. A.C. 1979 has the 1st as large
as the 3rd. There are eight supraciliaries in all specimens, except A.C. 1661, which has
nine through the intercalation of an additional scale above the eye, and A.C. 2162 with
a very large scale on the right side above the eye, reducing the total number to six. The
2nd supraocular is the largest in 27 cases. Contact of the 4th supraocular with the
parietal may be at a mere point. Two specimens R532 and A.C. 2137 have five infra-
labials, and R534 and A.C. 1965 seven. The left of the first chin-shields in A.C. 2137
has fused to the postmental. The depression of the ear varies from a wide, shallow area
to a comparatively narrow slit with a deep pocket about the centre. All specimens of
H.d. talbingoensis in the table except those from Talbingo are auxiliotypes. Nine of the
18 groups are represented, two by only one specimen each. Lack of dorsal and dorso-
lateral lines is evidently characteristic of populations near Adaminably where, of seven
individuals, one is uniform in colour, five have dorsal flecks, and one has six faintly
dotted lines.

Measurements of Ten Auziliotypes of H.d. talbingoensis in mm. (All Specimens with Complete Tails Included.)

Length.
Number. Snout- Tail. Snout- Axilla- Head. Fore- Hind-
' Vent. Forelimb. Groin. Length. Width. limb. limb.
A.C.1661 a3 be 55 33 155 36 7 6 6 8
A.C.1966 Se i 53 71 15 35 8 6 6 9
A.C.1973 3 aa 54 77 15 33 7 6-5 6 9
A.C.1976 ee Be 61 73 15 43 7-5 6 OLD) 8
A.C.1979 ae ae 50 67 14:5 32 7 55 6 8
A.C.2135 ne ae 61 55+ 15 43 8 6 Byol55 7285)
A.C.2136 ae BY 54 67 Bok) 37 a 6 5:5 8
A.C.2139 ha Si 27 P30 9 16 5 4 3 4-5
A.C.2161 ae ae 47 61 13 31 7 6 08) MG
A.C.2163 fs ae 31 37 10 19 5 4°5 5 6

Table of Midbody Scale Rows and Lamellae under Median Toes of H.d. talbingoensis Itemized by Localities.

Number of Number of
Seale Rows. Lamellae. Number
Locality. = of
20 21 22 24 6 8 Specimens.

“I

Marulan an a ate ue ths sa Si — — 1 — — 1 — 1
Cullerin —_— — 9 — 1 a 1 9
Adaminaby 5 — 2 — 4 3 — 7
Collector 1 - 4 oo 1 4 os 5
Victoria — -= 2 — 1 1 — 2
Mt. Kosciusko 8 2 7 — 3 14 — 17
Goulburn He — _- 2 — — 2 — 2
Total: auxiliotypes. . sy 14 2 27 — 10 32 1 43
Talbingo: holo- and paratypes — os 51 2 1 24 28 53

14 2 78 2 11 56 29 96

Total: all specimens

Hemiergis d. talbingoensis is an upland form occurring at altitudes between 1,300
and 5,000 feet. At the lower elevation the race is found in mountain valleys. It prefers
gentle slopes where the soil under logs and stones is just moist. I have never collected
a specimen where the ground was normally dry or wet. At Talbingo, where the sub-

BY STEPHEN J. COPLAND. . 719

species is common on hillsides leading down to the Tumut River, they were absent from
the dry crests of ridges between small creeks as well as at the swampy margins of the
watercourses. On the intervening slopes they were, in places, found under practically
every log. I did not find them in grass packed against the sides of logs not yet begun
to rot, which was a favoured habitat of Leiolopisma guichenoti. They prefer half-rotted
logs under which mould has collected. They are absent under timber which has been
eaten out by insects leaving a dry, cardboard-like interior, probably because termites and
other insects on which they depend for food have moved elsewhere. Grassy slopes on
which fallen logs give cover appear to be the chosen habitat of the lizards. Of 73 specimens
collected by me, at least 65 were under logs. Cullerin was the only locality where lizards
were found under stones. Field notes show that of the nine specimens collected near
Cullerin, three were under stones, one under a log, and five under stones and logs. I
have never seen one abroad in the daytime. They seem to be strictly cryptozoic, seeking
their food in mould under logs in the darkness. Scales, especially those of the head,
in most individuals bear longitudinal scratches caused by grit and other sharp objects
encountered while burrowing. When uncovered by lifting a log, they remain perfectly
still for several seconds and are then easy to capture. After the first surprise they
move away rapidly with a wriggling, snake-like motion. Once buried in mould they are
hard to find again. I have never seen a lizard using its weak limbs to move in a direct
line with its body straight. H.d. talbingoensis is apparently fairly resistant to cold for
no specimen was too sluggish to make an effort to escape, unlike, for example, Leiolopisma
entrecasteauxii (Duméril and Bibron) and Siaphos equalis (Gray), which often remain
curled up making no attempt to escape when disturbed in cold weather.

Lucas and Frost (1894, p. 24), who probably included specimens of H.d. talbingoensis,
follow Boulenger’s description except for measurements and give notes on habits and
distribution. They say “Habits similar to Hemiergis peronii’, i.e., “found under logs
and flat stones on the hillsides and in gullies. Movements very slow’. Distribution is
given as Ferntree Gully and Beechworth in Victoria, South Australia and Kangaroo
Island.

\

VI. HEMIERGIS DECRESIENSIS DAVISI, n. subsp. PI. vi, fig. 4.

Diagnosis: Hemiergis decresiensis davisi is separated from the typical subspecies
H.d. decresiensis by the lower number of midbody scale rows (20 in holotype against 24:
average for 136 specimens examined 19-90), lower number of lamellae beneath mid-toe
(six in-the holotype against eight: average for 133 specimens 5:77). Other points of
difference in scalation, colour (especially the four-lined dorsal pattern: Group J) and
size are dealt with in the tables and descriptions.

Holotype. Author’s Collection, No. 821; Poison Swamp Creek, 2:7 miles south of
Bendemeer, N.S.W., which is 30-51 S., 151-10 E. Altitude c. 2,500 feet. 5.xii.1940.

Description of Holotype.—Rostral moderately high, area visible from above equal to
half that of the frontonasal, long concave sutures with the nasals and nearly straight,
approximately vertical ones with the 1st supralabials; the fairly straight junction with
the frontonasal is about a quarter the width of the frontal. Nasals large, not in
contact, roughly quadrilateral, sutures long and convex with the rostral and frontonasal
and shorter and about straight with 1st supralabial and anterior loreal; the round nostril
is slightly behind the centre, there is no sign of a groove running from it on the left
side, but on the right a depression runs back from the upper margin to the anterior
loreal. No supranasals. Frontonasal large, subequal in area to the frontal, also in
contact with prefrontals, nasals, rostral and anterior two-thirds of upper margin of
anterior loreal. Prefrontals large, well developed, sutures long and nearly straight with
frontonasal, convex with frontal, nearly straight with lst supraciliary, which separates
it rather widely from the Ist supraocular, slightly concave with posterior loreal, only
narrowly in contact with anterior loreal. Frontal kite-shaped, not much longer than
broad, rounded posteriorly against left frontoparietal, slightly indented anteriorly against
frontonasal and at sides against lst supraciliaries, long nearly straight postero-lateral
sides against lst and 2nd supraoculars, shorter, slightly concave, anterolateral sutures
with prefrontals. Frontoparietals paired, subequal in area with interparietal, left scale

80 VARIATION IN THE LIZARD HEMIERGIS DECRESIENSIS,

a rough crescent, twice as long as wide, long median convex border against parietal,
interparietal and right frontoparietal, irregular outer border against 2nd, 3rd and 4th
supraoculars, indented slightly against frontal; right frontoparietal in contact with the
same scales, except frontal, but in shape squat and roughly quadrilateral, pointed between
interparietal and left frontoparietal, separated from frontal by 2nd supraocular. Inter-
parietal kite-shaped, approximately similar in shape and size to the frontal, rounded
behind, pointed in front and at sides, sutures long and nearly straight with parietals,
shorter with frontoparietals, concave with left, slightly sinuous with right; a rounded
milky-coloured area in the midline one-third of the length of the scale from its posterior
end covers the pineal foramen. Parietals are the largest head shields, irregularly shaped,
at least twice as long as broad, meeting in an oblique suture behind the interparietal,
other sutures are straight and long with interparietal, shorter and concave with fronto-
parietals, very short with 4th supraocular, 8th supraciliary and 2nd postocular straight

Va

Figs. 2-3.—Head scales of H.d. davisi. 2. Dorsal view. 3. Lateral view.

A, accessory palpebrals, upper and lower; B, anterior loreal; C, chin-shields, first and third;
D, frontal; EH, frontonasal; F, frontoparietal; G, infralabials, first and sixth; H, interparietal;
I, lower secondary temporal; J, mental; K, nasal; Ll, nuchal; M, palpebral series, upper and
lower; N, parietal; O, posterior loreal; P, postmental; Q, postoculars, first, second and third;
R, prefrontal; S, preoculars, upper and lower; T, primary temporal; U, rostral; V, suboculars,
first and fourth; W, supraciliaries, first and eighth; X, supralabials, first and seventh; Y,
supraoculars, first and fourth; Z, upper secondary temporal.

BY STEPHEN J. COPLAND. 81

with upper secondary temporal, left scale in contact with two dorsal scales, right with
three. There is one unmistakable pair of nuchals. Seven supralabials, the anterior three
roughly quadrilateral with postero-dorsal points of the first two directed backwards:
the upper margins of the scales form a straight line with the nasals, loreals and lower
preocular; posterior four pentagonal, haystack-shaped, lower margins horizontal, anterior
and posterior sutures vertical, the other two sides meeting in a point dorsally. Size in
decreasing order, 7, 6, 3, 2, 1, 5, 4, 5th under centre of eye. Primary temporal a rough
square, antero- and postero-ventral sides between 6th and 7th supralabials, postero-dorsal
side against upper secondary temporal, point between posterior sides narrowly in contact
with lower secondary temporal, antero-dorsal side in contact with 2nd and 3rd _ post-
oculars and 4th subocular. There are two secondary temporals, the upper twice the
size of the lower, which is subequal to the primary temporal. Body scales begin behind
the parietals, secondary temporals and 7th supralabials. Scales behind the secondary
temporals are slightly enlarged. The two loreals are roughly oblong, slightly higher
than long, the anterior is between the nasal, frontonasal, prefrontal, posterior loreal and
two anterior supralabials, the posterior between the anterior loreal, prefrontal, 1st
supraciliary, upper and lower preoculars and 2nd supralabial. The upper and lower
palpebral series abut against the upper preocular, which is also in contact with the 1st
supraciliary, the upper accessory palpebral (a small scale intercalated between the
upper palpebral series and the ist and 2nd supraciliaries), the posterior loreal, lower
preocular and the lower accessory palpebral (a small scale between the anterior of the
lower palpebral series and the ist subocular). The lower preocular is twice the size
of the upper and lies between it, the posterior loreal, 3rd supralabial, 1st subocular and
the lower accessory palpebral. There are four suboculars, the anterior three roughly
equidimensional with downwardly-directed wedge-shaped points lying between the supra-
labials; the 4th is oblong, the 3rd is the largest and the 4th the smallest. The 4th lies
between the primary temporal, 3rd subocular, 3rd postocular, 6th supralabial and the
small postpalpebral scales. The postoculars are well developed, the 2nd being larger
than the 4th subocular and equal to the 8th supraciliary. There are eight supraciliaries,
the anterior and posterior scales largest and flexed away from the curved chain formed
by the other six. The 7th is also large and the chain could be taken as ending either
with or against it; in the former case a name would have to be given to the posterior
scale and, in the latter, names to the two posterior scales. It is simpler to regard the
whole eight scales as members of the supraciliary chain. The ist supraciliary is
irregularly quadrilateral, lying between the prefrontal, frontal, 1st supraocular, 2nd
supraciliary, upper accessory palpebral, upper preocular and just touching the posterior
loreal. The 8th lies between the 4th supraocular, parietal, 1st and 2nd postoculars and
7th supraciliary. The posterior scales of the upper and lower palpebral series and a
number of small postpalpebral scales abut against the 7th supraciliary. There are
four large supraoculars, the 2nd by far the largest; the 3rd is comparatively small and
bandlike on the right side but well developed on the left, the Ist is separated from the
prefrontal by the 1st supraciliary, the frontal is in contact with the 1st and 2nd, and
the frontoparietal with the 2nd, 38rd and 4th. The transparent disc in the lower eyelid
is undivided, convex, and lens-like, more than half the length of the eye aperture. Its
upper rim is one scale wide, and before and behind it is bounded by groups of small
scales. The large mental and postmental are followed by three pairs of chin-shields,
the Ist pair in contact, the 2nd separated by a large median scale, and the 3rd by three
small scales. There are six infralabials, the 1st is higher than long, but they gradually
become more elongated posteriorly until the 6th, which is twice as long as high.

The ear about six scales behind the mouth runs obliquely downwards and forwards
as a shallow, scale-covered, slit-like depression, deepest postero-dorsally over the actual
ear. There are about four scales along its anterior border.

Scales 20 at midbody, subequal, but slightly larger dorsally. Caudal scales larger,
subcaudal row widest, 13 rows round the tail at length of hindlimb behind vent, number
decreasing progressively to tip of tail, where the terminal scale is spine-like and at
least twice the length of nearby scales. Four preanal scales, inner pair greatly enlarged,
outer pair very small. Scales from above vent to parietals, 78. Body much elongated;

82 VARIATION IN THE LIZARD HEMIERGIS DECRESIENSIS,

the distance between the end of the snout and the forelimb is contained 3-14 times in the
distance between axilla and groin.

Limbs small and weak, especially the forelimbs, separated by nearly three times
the length of the hindlimbs when adpressed. Lamellar formula for fingers, 4, 5, 5. Nine
enlarged tubercles, the three largest across the wrist, surround three about the same
size. Lamellar formula for toes, 5, 6, 6. Seven large flattened tubercles on the sole
surround two of the same size.

Measurements are given with those of the paratypes.

Colour. Ground colour of head, body and tail is medium brown. A black dorso-
lateral line begins just behind the eye in the 1st postocular, runs through the centre of
the upper secondary temporal, and follows the third row of dorsal scales, occupying about
the middle third of each scale, to the tail, where at the length of the hindlimb behind
the vent, it switches to the second row of scales. It then continues to the end of the
tail becoming progressively more irregular. A pair of black dorsal lines about a quarter
of a scale wide, begins behind the parietals and runs to the tip of the tail. Continuous
on the body, the lines consist of discontinuous wedge-shaped dots on the tail. Slightly
interrupted black lines, about a quarter scale wide, run along the second row of dorsal
scales, originating just behind the parietals and petering out before the tail, where
their position becomes occcupied by the dorsolateral lines. Dark brown to nearly black
‘patches occur on all the head shields, being most concentrated on the supralabials.
Under high power they are seen to consist of hundreds of tiny deep brown dots,
normally scattered widely at the borders of the patches, then forming dendritic patterns
and finally a dense aggregation. Hundreds of minute dots occur on all dorsal, lateral
and dorsal caudal scales, darkening the whole colour of the specimen. Five rows of
lateral scales below those carrying the dorsolateral lines bear irregular ill-defined
longitudinal lines, formed by an aggregation of pigment about the centre of most scales.
These dark markings become more pronounced on the tail and occupy more of each
scale until those near the tip are practically all black. Ventral scales are yellow, each,
with the exception of a few midway between the limbs, with a brownish dot just behind
the centre. These dots become larger posteriorly. The throat and neck to the fore-
limbs are reticulated with brown, only the centres of the scales being free of dots or
nearly so. About half of the two large inner anal scales is occupied by crescent-shaped
markings. Scales behind the vent are about half black. The black areas have a

brownish margin of dots, and the borders of each scale are yellowish. Scales become

progressively darker posteriorly, but some yellow persists to the end of the tail. This
is the six-lined pattern of Group L.

The race is named for the late Dr. H. F. Consett Davis, whose death in a plane
accident in New Guinea on 12th December, 1944, was a loss to science and his friends.

Variation in Paratypes (A.C. 818-20, A.C. 822)—Minor variation does not affect
the essential uniformity of the Bendemeer series of five specimens. The suture between
the rostral and frontonasal in A.C. 818 is equal to a quarter the width of the frontal,
which is slightly longer than in the other three paratypes and the holotype. No paratype
shows any sign of a divided nasal. The prefrontals of A.C. 818 meet in a point, and are
separated by a distance equal to a quarter the width of the frontal in A.C. 822, otherwise
as holotype. In A.C. 818 the frontal is one and a half times as long as broad. In all
four specimens both frontoparietals touch the frontal. In A.C. 819 the ‘suture between
the parietals slopes backwards towards the right instead of the normal left. No
specimen has nuchals but A.C. 822 has a large irregular scale on the left, and A.C. 818-20
have a large irregular scale on the right. In A.C. 819 the primary temporal is antero-
dorsally in contact with only two scales—the 2nd postocular and 4th subocular, the 3rd
postocular having fused to one of these two scales. The lower preocular is in contact
with the 2nd supralabial as well as the 3rd in A.C. 818 and A.C. 819. A.C. 820 has
five suboculars on the right side. All paratypes agree with A.C. 821 in having large and
well-developed 2nd postoculars at least equal in size to the 8th supraciliary. The ist
supraocular is never in contact with the prefrontal. The distance between the tip of the
snout and the.forelimb is contained in the distance between axilla and groin 2-64 times
in A.C. 818, 2:50 times in A.C. 819 and A.C. 822, and 2-80 times in A.C. 820. Each specimen

—— ee

oe

Se ee ee

BY STEPHEN J. COPLAND. ; 83

has six lamellae beneath the middle toes. All paratypes belong to the four-lined pattern
of Group J.

Measurements of Holotype and Paratypes of H.d. davisi in mm.

. Length.

Number. Snout- Tail. Snout- Axilla- Head. Fore- Hind-

Vent. Forelimb. Groin. Length Width. limb. limb.
A.C.818 .. Ae ae 54 54+ 14 37 6°5 5 5-5 8
A.C.819 .. ee als 51 54+ 14 33 i 5 5 7
A.C.820 .. 2 ve 57 35+ 14°5 40 7 5-5 BG) 8
A.C.821 .. Fo se 60 75 14 44 8 6 6 8
6 4°5 5 7

A.C.822 .. ot 20 48 51 12-5 30

Specimens examined and Locality Records of Hemiergis d. davisi.

(A.C. 474) Armidale, N.S.W. (C. Davis), May, 1939. 12.

(AL@.- 5212-5) 5:5 m. N. of Liangothlin, N.S.W., 30.%.1939. 13.

(A.C. 729) nr. Little Hartley, N.S.W., 11.11.1940. 14.

(A.C. 748) Armidale, N.S.W. (C. Davis), 18.11.1940. 12.

(A.C. 744) 3 m. NW. of Armidale, N.S.W. (C. Davis), Feb., 1940. 12.

(A.C. 768-73, 775) 2m. W. of Hartley, N.S.W., 21.vii.1940. 14.

(A.C. 818-22) 2:7 m. S. of Bendemeer, N.S.W., 5.xii.1940. 15.

(A.C. 1136) Armidale, N.S.W: (C. Davis), Nov., 1940. 12.

(A.C. 1154) Armidale, N.S.W. (C. Davis), 23.xi1.1940. 12.

(A.C. 1162-3) Lett River, nr. Hartley, N.S.W., 19.i1.1941. 14:

(A.C. 1200-1) 10 m. from Jenolan Caves on Hampton Rd., N.S.W., 20.11.1941. 16.

(A.C. 1373) Armidale, N.S.W. (C. Davis), 26.x.1941. 12.

(A.C. 1695) 2:3 m. BE. of Laggan, N.S.W., 30.1.1943. 17.

(A.C. 1701-3) 8-8 m. N. of Abercrombie River on Taralga-Oberon Rd., N.S.W., 31.1.1943. 18.

(A.C. 1704-7) 11:2 m. N. of Abercrombie River on Taralga-Oberon Rd., N.S.W., 31.11.1943.
18.

Po ePreby we HH mya rRRe PE

8 (A.C. 1710-7) 3 m. N. of Curraweela, N.S.W., 31.13.1943. 18.
10 (A.C. 1718-27) 5:2 m. N. of Abercrombie River on Taralga-Oberon Rd., N.S.W., 31.11.1943.
18.
CAC 1732) 7 m. S) of Black Springs; N:.SOW.; 111.1943: 19:
(A.C. 17386) (1-6 m. NE. of Black Springs, N.S.W., 1.i11.1943. 19.
(A.C. 1748-51) 5m. S. of “Rorter’s Retreat, N.S.W., 1.111943. 18.
(A.C. 1752) 0-7 m. from Duckmaloi River towards Oberon, N.S.W., 1.ii1.1943. 20.

(A.C. 1758-9, 1762-3) Duckmaloi River nr. Oberon, N.S.W., 2.i11.1943. 20.

(A.C. 1764-72) 3 m. from Oberon on Jenolan Caves Rd., N.S.W., 2.11.1943.. 20.

(A.C. 1781) 4 m. from Hampton on Oberon Rd., N.S.W., 4.i1.1943. 16.

(A.C. 1799-803, 1812-7) 4 m. from Hampton on Oberon Rd., N.S.W., 23-24.iii1.1943. 16.
(A.C. 1818) nr. Little Hartley, N.S.W. (D. Ross), 19.iv.1943. 14.

(A.C, 1820-2) 1 m. S. of Rydal, N.S.W. (D. Ross), 19.iv.1943. 14.

(A.C. 1834-42) 2 m. S. of Black Springs, N.S.W. (D. Ross), 20.iv.1943. 19.

(A.C. 2228) 5 m. SE. of Armidale, N.S.W. (C. Davis), 13.iv.1944. 12.

(R.994, Aust. Mus.) Forest Reefs, N.S.W. (H. J. McCooey), Feb., 1891. 21. }
(R.1348-9, 1351, 1353, Aust. Mus.) Hartley, Blue Mts., N.S.W. (R. Grant), Nov., 1892. 14.
(R.2514, Aust. Mus.) Tarana, N.S.W. (W. Hawken), June, 1899. 14.

(R.2919, 2922-3, 2925, Aust. Mus.) Salisbury, N.S.W. (D. A. Porter), June, 1900. 22.
(R.3189-94, Aust. Mus.) Tarana, N.S.W. (W. Hawken), June, 1901. 14.

(R.3613-8, Aust. Mus.) Southern Australia. No date.

(R.3982, Aust. Mus.) Hartley Vale, N.S.W. (A. H. S. Lucas), no date. 14.

(R.3983, Aust. Mus.) Capertee, N.S.W. (A. H. S. Lucas), no date. 23.

(R.3984, Aust. Mus.) Bundanoon, N.S.W. (A. H. S. Lucas), no date. 24.

(R.3985, Aust. Mus.) Liverpool Plains, N.S.W. (A. H. S. Lucas), no date. 25.
(R.10925, Aust. Mus.) Lithgow, N.S.W. (H. E. P. Bracey), May, 1933. 14.

(R.12268, Aust. Mus.) Oberon, N.S.W. (C. Davis), 13.x.1937. 20.

Variation in Auxiliotypes (131 Specimens examined).—Length of the suture between
rostral and frontonasal to the width of the frontal varies from one-half to one-tenth
(4, 4; 27, 4; 35, 4; 23, &: 21, 4; 12, 4; 8, 4%; 1 abnormal). This character has little
geographic significance although the suture tends to be wider in northern specimens.
A.C. 1136 has the frontonasal divided on the right side so that the main portion is
separated from the anterior loreal. A.C.1803 has the scale divided symmetrically on each
side so that the main part remains shield-shaped, longer than wide, and the cut-off
portions lie between it, the prefrontal, anterior loreal and nasal. In A.C.1154 a similar

PRE H EP HE HE H ODP RH HOW HrPHEP Ho RH BH

ee)
ee

VARIATION IN THE LIZARD HEMIERGIS DECRESIENSIS,

pair of scales is incompletely separated. In A.C.1801 the prefrontals are enormously
developed and occupy most of the area normally taken by the frontal, which is missing;

contact between the two prefrontals is longer than the length of the frontonasal. The
suture between the frontal and frontonasal is slightly wider in Northern Tableland

specimens (including holotype and paratypes) than in those from the Central Tableland,

the average distance compared to the width of the frontal being 0-236 (1, in contact;

eae (2p a ee Ge. lo Ge Os ae Ay BS Gy fs) Bysewnagi, Oemall (GG sha jayauayeis 4h, <i52 IB}, Zhe

13, 4; 15, 4; 28, 4;- 238, 4; 1, 3; 2, abnormal). The frontal, normally one and a half

times as long as wide, varies from as broad as long to twice as long as broad. The

frontal is in contact with the right frontoparietal in all except four cases. A symmetrical
scale has formed from portions of the right and left frontoparietals in R.3193, so that

three kite-shaped scales—the frontal, the abnormal scale and the interparietal—follow

each other down the midline; the remaining sections of the frontoparietals are laterally

in contact with only the 3rd and 4th supraoculars. A.C.770 has the posterior third of
the left scale divided off; and the anterior portion of the left frontoparietal has fused
with the right in R.3189. The interparietal is occasionally squat but normally one and

a half times as long as wide. In A.C.1770 it is malformed into an irregular rounded

square; in one of the three R.2919 specimens it is reduced in size to one-third the area
of a frontoparietal; and in A.C.1749 and A.C.1836 it is fused with the left parietal. The
suture between the parietals runs backwards towards the right in 27 individuals, towards

the left, as in the type, in 104. About half the specimens have no nuchals, about 15 per

cent. one pair, about 10 per cent. two pairs, about 7 per cent. one large scale (in nearly
every case on the left side), while the remainder have combinations such as two or
three large scales on the left or right; one specimen has three irregular pairs of nuchals.
Of 15 specimens with six supralabials, the 4th under the eye, Hampton and Black Springs
are each represented by seven and Bundanoon by one. The posterior loreal is rarely in
contact with the 3rd supralabial. The lower preocular, which is often three times larger
than the upper, is nearly always in contact with the 2nd and 3rd supralabials, but in
about one-fifth cases it touches only one of these scales. The 4th subocular is divided
on the right side of A.C.1751. There are seven supraciliaries on each side in A.C.1836,
seven on the right side only in A.C.1835, R.3190, R.3613 and R.3614. <A.C.1801 has
immense, abnormal 2nd supraoculars meeting in a wide suture anterior to the fronto-
parietals. The 2nd supraocular is divided on the right side in A.C. 1799 and R.3616.

There are three supraoculars on the right side of A.C.1800, the anterior two having
fused, and three on each side in one of the R.2919 specimens, the anterior two on the
left side being incompletely separated. The depression of the external ear varies consider-
ably in length and depth.

All specimens of H.d. davisi in the colour table, except those from Bendemeer, are
auxiliotypes. Sixty of 131 individuals have the four-lined pattern of Group J, two connect
this group through Group K to the six-lined pattern L, to which five auxiliotypes belong.
Twenty-seven of the 29 specimens from the Northern Tableland belong to Group J
(including the four paratypes), and the remaining two (including the holotype) to
Group L. Thirty-six Central Tableland specimens belong to the closely allied, exclusively
H.d. davisi, Groups N, O and P. Lack or searcity of dorsal pigment is evidently less
common in H.d. davisi than in H.d. talbingoensis. The northern race is represented by
only eight cases in Groups Q and R against 20 of the southern although 136 individuals
were examined against 96. R.12268 in the Australian Museum, collected at Oberon on
13th October, 1937, by Dr. H. F. C. Davis, is a representative lizard from the Central
Tableland. With 20 midbody scale rows and lamellae beneath the three toes, 5, 6 and 6,
the colour description is: the four distinet black lines of Group J; head flecked more
or less longitudinally with black; sides greyish-black with lighter and darker spots;
venter yellow: throat to forelimbs brown-flecked, posterior third of most scales being
brown; underside of tail increasingly dark caudad.

Undersides of specimens of H.d. davisi (also H.d. talbingoensis and probably also
the other two races) are very rarely white, more frequently pale lemon-yellow, but in
the great majority vary between chrome-yellow and orange.

BY STEPHEN J. COPLAND. 1 85

Measurements of Eighteen Auxiliotypes of H.d. davisiin mm. (Only Specimens with Complete Tails and generally only
One from Each Locality Included.)

Numbers Al@:522) " JAlC:729) “ArC2773 AVC 1054. A-C21695 ASCl1705 ~A-C@.1713 A/€:1'736 A. C11762
Snout-vent 55 60 47 45 5S 67 53 61 62
Tail ae 63 66 58 55 64 71 70 72 87
Axilla-groin 35 41 30 29 35 46 333 40 38
Snout-forelimb 15 15 13 12 14 15 15 15 16
Number .. A.C.1821 A.C.1839 R.1353 R.2514 R.2923 R.2925 R.3189 R.3193 R.10925
Snout-vent 63 65 49 56 63 54 60 60 60
Tail ae 72 68 57 65 7) 70 80 74 78
Axilla-groin 43 45 32 39 45 38 38 42 39
Snout-forelimb yy 15 13 14 14 13 16 15 16

Table of Midbody Scale Rows and Lamellae under Median Toes of H.d. davisi iemized hy Localities.

Number of Scale Rows. Number of Lamellae.
Locality.

18 20 22 4 5 6 7
Liangothlin .. a. ee as _- 4 — oo — 3 _-
Armidale Me ae ae st — 7 — — — 7 —
Liverpool Plains ae ae we — 1 —- — — — 1
Salisbury He a se ae 1 10 1 — — li 1
Capertee be ens ne ae _— —_ 1 — 1 — —
Forest Reefs .. 8% =. ee — —_— il — — — 1
Lithgow so ee te fa — 1 — — — 1 —
Rydal .. 1 72 — =~ 3 —
Tarana ie 2 — 6 1 — —_— 7 —
Hartley Vale oe ab — 1 — —— — — a=
Victoria Pass and Hartley.. 1 12 2 — 1 14 —
Hampton 8 6 2 9 3 —_—
Oberon a are Be 1 13 1 — 4 iil —
Black Springs Sie 24; st 5 6 2 6 3 —
Porter’s Retreat — 21 — —_— 4 15 2
Curraweela -- 8 — —_ 1 5 1
Laggan — 1 — = = 1 =
Bundanoon a — 1 — — 1 —
Southern Australia .. ein — 7 — — — o —
Total: Northern Tableland il 27 i — — 26 2
Total: Cental Tableland 5 84 8 4 27 70 4
Total: auxiliotypes. . ate its 16 106 9 4 27 91 6
Bendemeer : holotype and paratypes —_— 5 — — = 5
Total: all specimens a a 16 111 9 4 27 96 6

Three specimens with mutilated or abnormal toes have been excluded from the lamellar counts, making the total
examined 133 against 136 examined for midbody scale counts.

Tails in only 45 of the 136 specimens of H.d. davisi are complete. Others are in all
stages of regeneration, ranging in length from 5 mm. to 76 mm. Some closely simulate
undamaged tails, others being blunt and a few ending in short spines.

Loveridge (1934, p. 370) apparently included specimens of H.d. davisi under Siaphos
equalis (Gray). Under the heading of Siaphos equalis he makes the following remarks
and gives a key to separate Siaphos equalis and Hemiergis decresiensis.

“Numbers 10189-10191 were received as Hemiergis decresiense, a species which they
closely resemble. Apart from the scaly lower eyelid, a character which is often some-
what obscured, the two may be distinguished as follows:

Midbody scale rows 18-22, average 20; lamellae beneath median toe 3-6. Total

iMesalencl ah alae aaa gals is lhe meee Neen eee Cent aaalery chin itee Ss. CreNCRO GIS ret rusia em cececaene area equalis.
Midbody scale rows 24-26, average 24; lamellae beneath median toe 7-9. Total
Nemo AO Smee epee ee mer Ate | rel Gira Phyecstinceain crea an cote sus faewawe Geen eter d decresiense.”

The key is valid when only H.d. decresiensis and H.d. continentis are considered
but places specimens ‘of H.d. davisi and H.d. talbingoensis under Siaphos equalis.

86 VARIATION IN THE LIZARD HEMIERGIS DECRESIENSIS,

Loveridge apparently did this with his Salisbury (M.C.Z. 10190-1) and Hartley Vale
(M.C.Z. 10189) specimens. His three lizards were received from the Australian Museum.
Those from Salisbury are certainly part of a large series collected in that locality by
D. A. Porter in June, 1900, an entry noting the exchange occurring in the museum
register. The 12 specimens of the series remaining in the Australian Museum are
undoubtedly H.d. davisi. Salisbury individuals differ from any others that I have
examined in having very small transparent discs. The discs, though small, are distinct.
The lizards are typical in colouration, pattern and scalation.

eg
It may be mentioned here that the lowering and widening of the Cassilis Gap may
be expected to split H.d. davisi into Central Tableland and Northern Tableland races.

Identification of Siaphos equalis (Gray) with Hemiergis decresiensis is of long
standing. Duméril and Bibron (1839, p. 766) include Seps aequalis, ? Siaphos aequalis
and Peromeles aequalis in the synonymy of H. decresiensis, which species they attribute
to themselves.

Steindachner (1867, p. 50) also synonymizes Siaphos equalis with H. decresiensis.
His five specimens from Sydney were almost certainly all S. equalis.

Giinther (1867, p. 48) (see reference under H.d. decresiensis) may have made his
comparison either with H.d. davisi or Siaphos equalis. Both agree with the two diag-
nostic points he noted—18 or 20 midbody scale rows and toes less developed than in
Kangaroo Island specimens. On the grounds that individuals of H.d. davisi with 18
midbody scale rows are comparatively uncommon and that collections would more
probably have been made near Sydney (where S. equalis is abundant) than on the
tablelands, I am inclined to think that the comparison was made with 8S. equalis.

In a reference to Hemiergis decresiensis, H. Claire Weekes (1929, p. 43) says:
“Although the oviparous species appear to be more or less restricted to lower levels,
many of the viviparous lizards flourish at all altitudes, Tiliqua scincoides, Hgernia
whitei, E. cunninghami, E. striolata, Lygosoma (Hemiergis) decresiensis and L. quoyi
having been collected in large numbers in coastal districts little above sea-level.”

Three criteria—‘large numbers, coastal districts and little above sea-level’”—suggest
that the lizard noted as Hemiergis decresiensis is Siaphos equalis.

Seven specimens of H.d. davisi in the Australian Museum (R.3613-8; R.3618, two
specimens) are labelled Southern Australia. As the locality is indefinite and Boulenger

(1887, p. 327), by whose description and key the lizards were probably identified, gives |

the range as “Southern Australia’, it is probable that the locality was only written in
at the time of identification for the sake of completeness.

Distribution.—The northern range of H.d. davisi may be expected to extend only
slightly beyond Llangothlin, the present extreme record in the author’s collection,
although the same ecological conditions persist to the Macpherson Ranges on the Queens-
land border. As far as I am aware no specimen of Hemiergis has been found north of
Llangothlin although fairly extensive collecting has been carried out in this area,
especially the Macpherson Ranges. Searches made by me at Bungulla, 82 miles north
of Llangothlin and Wilson’s Peak at more than 3,000 feet on the Queensland border
have been unsuccessful. The low-lying Lake George senkungsfeld forms the southern
boundary of the subspecies. The definitive characters of H.d. davisi and H.d. talbin-
goensis become more marked as we move away from this zone, particularly so north-
ward for H.d. davisi. No exact line can be drawn to separate the two forms, but this
may follow in time if the populations on each side of this belt of dilution in numbers
breed true. It is more probable that a zone of hybridization will persist. The present
condition is apparently a stage in which neither of two gene-complexes has assumed
complete dominance.

VII. RELATIONSHIP OF THE FOUR SUBSPECIES.
The two following tables illustrate the gradual reduction in number of midbody
scale rows and mid-toe lamellae from the closely allied H.d. decresiensis and H.d.
continentis through H.d. talbingoensis to H.d. davisi.

——_— a

rw

.

BY STEPHEN J. COPLAND. 87

Table of Midbody Scale Rows.

Scale Rows x0 a0 Bs b0 ais 18 20 22 24 26 Average.
H.d. decresiensis (6 specimens) a fee = — — 6 ess 24-00
H.d. continentis (20 specimens) be nc = = 19 1 24-10
H.d. talbinyoensis (94 specimens)* ae —— 14 73 2 a 21-74
H.d, davisit (136 specimens) a ~ 16 111 9 —s = 19-90

* Two specimens with 21 rows not included.

Scale rows should be counted exactly at midbody for the sake of uniformity. There
is little chance of making a mistake in most specimens, but in some, additional rows
from before or behind approach midbody.

SONLE IRONS
hese Lye Lbs 6

00. Paw OE CRESIENSIS
Lavo ALL SPECIMENS
2 ;
FZ
LU
=
©
Li)
OL
Y)
Le
S
LU
©
<I
ae
Ee,
LU
@
(Oe
LU
OQ
LAMELLAE
CONTINENTIS = i
KEV ELDITADBINGORNSIG <= 2 eas ae
DAVIS| > L=LAMELLAE

Fig. 4.—Graph showing the close agreement between midbody scale and mid-toe lamellae
characters in each race; also the concentration of individuals at three modal centres, illustrating
close relationship between H.d. decresiensis and H.d. continentis and separation of the other two
subspecies. The graph gives an exaggerated idea of overlap.

88 VARIATION IN THE LIZARD HEMIERGIS DECRESIENSIS,

Table of Mid-toe Lamellae.

Number of Lamellae.

4 5 6 i 8 9 Average.
H.d. decresiensis (6 specimens) — — — —_— 6 — 8-00
H.d. continentis (20 specimens) —_— —- — 1 18 1 8-00
H.d. talbinaoensis (96 specimens) — a= 12 55 29 — 7-18
H.d. davisi (133 specimens)* .. 4 27 96 6 = 5:77

* Three specimens, R.3982, toes mutilated ; A.C.524, 7 on right, 6 on Jeft ; A.C.1717, 6 on right, 7 on left ; are
not included.

The following table gives the number of specimens against body lengths, which
have been tabulated in classes, in the four subspecies.

Body Lengths in mm.
23-27 28-32 33-37 38-42 43-47 48-52 53-57 58-62 63-67 68-72

HI.d. decresiensis a 1 — 1 2 2 —
H.4. continentis hs — — 1 1 10 6 |

H.d. talbingoensis =e 1 5 4 — 4 oases 32 20 5) —
A.d. davisi ae ae 1 1 — 4 10 20 36 28 27 9

Body lengths gradually increase from H.d. decresiensis to H.d. davisi. No specimen
of H.d. decresiensis (average length 37:83 mm.) exceeds 45 mm. Forty per cent. of
H.d. continentis (average length 47-00 mm.) are in excess of 47 mm., the largest being
54 mm., but none enters the three largest groups. Twenty-five specimens of H.d.
talbingoensis (average length 52:40 mm.) exceed 57 mm., five being in the 63-67 group
(longest 64 mm.). Twenty-seven individuals of H.d. davisi (average length 56:69 mm.)
enter the 63-67 group and nine extend into the 68-72 group, being the only one of the
four subspecies to do so; the longest specimens are A.C. 1799 and A.C.1800 from near
Hampton, which each measure 72 mm.

The average separation of the prefrontals compared to the width of the frontals
gives a measure to a diagnostic character of the genus Hemiergis—well-developed
prefrontals. In H.d. decresiensis the average separation is 0-108 (no specimens with
prefrontals in contact), in H.d. continentis 0:155 (2), in H.d. talbingoensis 0-139 (15),
and in H.d. davisi 0-218 (9).

Lizards of the four races are found at greater heights above sea-level as we proceed
northwards. This is more probably due to the fact that suitable habitats are only
found in New South Wales at higher altitudes than in southern states than to any
direct response of the animals to the effects of altitude.

Key to the four races of Hemiergis decresiensis.
Midbody scale rows 24 or 26, lamellae under mid-toe 8 (few 7 or 9).

Body short (average less than 40 mm.), habitus slender .............. H.d. decresiensis
Body longer (average more than 45 mm.), habitus more robust .......... H.d. continentis
Midbody scale rows 22 (exceptionally 24, few 20, average 21:74), lamellae under mid-toe 7
CLEW6 OLAS AVES ST LS ey css nee a ee cere Te eR a TI ee el i ERS UE H.d. talbingoensis
Midbody scale rows 20 (few 22 or 18, average 19:90), lamellae under mid-toe 5 or 6 (few
AOFS hs ANCLALS! DAUM) Sicthcsyeleucleis leis Gum a ceusktkecomnee Al eee Wate In Le Cae H.d. davisi

VIII. CoNCLUSIONS.

The question of the evolution of the four races cannot be decided finally, but a
satisfactory hypothesis is that H.d. continentis most closely approaches to the parent
form and that the genetic trend is towards reduction in the number of midbody scale
rows and subdigital lamellae, to mention two important characters, H.d. continentis with
its high number of scale rows and uniformity in other characters appears to be the most
conservative race. Isolated from the mainland, H.d. decresiensis remained essentially
unchanged but became smaller and slimmer in habitus. H.d. talbingoensis developed into
the widespread race of the Southern Tableland of New South Wales by the reduction of
midbody scale rows to 22. The population of the type locality, Talbingo, is most uniform,

BY STEPHEN J. COPLAND. 89

2
WY
D
FZ
LU
| D
(ey
@e
5)
a
10
2 D
m7 =
= Zz
ES) —
LW S
ae * S
Tp)
3k
ce
ae =
4
17 =
=
GS 2
mom 3 a
Dm
a) Pp
Z a7
IS &
S
Ze
an
8 n>.
3 al ee ‘ ee
215 2630 3185 36L0 LHS 650 5155 5660 6165 6670 7175 MM, :

BODY LENGTHS (IN CLASSES )

Fig. 5.—Graph, based on slightly different classes to those in the foregoing table, showing
body lengths shortest in the insular race, an increase in the nearest continental population, and
then two further increases by way of H.d. talbingoensis to the most northern form H.d. davist.

yet two specimens of the series of 53 still have the relict number of 24 midbody scale
rows. From this viewpoint specimens with 20 rows must be taken as members of the
advance guard of H.d. davisi and probably indicate that there will in time be a general
reduction to this number. H.d. davisi must be taken as a form in which rows have been
reduced to 20. The few individuals with 22 show that absolute uniformity has not been
reached. In some centres reduction to 18 rows is proceeding and it is impossible to
judge to what extent this change will be carried. Northern Tableland populations have
been practically unaffected by the trend towards 18 midbody scale rows, but those in
parts of the Central Tableland are altering rapidly. Rare examples of H.d. davisi with
22 rows may be regarded as living in comparatively isolated centres. Migration, with
its interchange of genes and consequent establishment of new characters, must be very
slow for the weak limbed, cryptozoic H. decresiensis. EXven more mobile lizards, which
have apparently excellent means of distribution, have little tendency to migrate—see
G. K. Noble’s studies as noted by Dobzhansky (1937, p. 145)—and voluntary movements

90 VARIATION IN THE LIZARD HEMIERGIS DECRESIENSIS,

of individuals in sufficient numbers to allow establishment in new territory are prob-
ably only brought about by gross overcrowding. No evidence of this state of affairs has
yet been adduced for the species. Carriage in logs during floods is probably the only
accidental means of dispersal, yet this must be an unimportant factor because the
animals generally lie in the mould under logs and not in them, the slopes they frequent
are unlikely to be flooded sufficiently to move logs of any size, turbulent mountain streams
would be an unsuitable medium for the prolonged carriage of passengers, and the odd
individual carried downstream would probably most often find itself in unfavourable
surroundings. Establishment of characters to the order of a new race would be always
slow and, as many factors are involved, more complex than say the rapid change from
red to white head colour in the African Barbet (Lybius torquatus) in southern Nyasaland
(Mayr, 1942, p. 77). The odd individual of H.d. decresiensis with 26 midbody scale rows
may indicate that the parent stock was originally uniform at this high number. All
four races agree with the condition of Mayr (1942, p. 16) that ‘‘in the case of subspecies,
it is a good convention that at least 75 per cent. of the individuals in one subspecies (or
of the available specimens) should be separable, on the basis of their diagnostic
characters, from the specimens of the most similar subspecies’. Where one subspecies
has practically replaced another and in its turn is being replaced by a third, as in
H.d. talbingoensis, the subspecies must be defined (inter alia) as a group of individuals,
the great proportion of which have attained distinctive genetic stability (as indicated
by diagnostic taxonomic characters) but whieh contains a small percentage of specimens
of the older subspecies which it is replacing as well as a small percentage of the newer
subspecies which is replacing it. The alternative to accepting these qualifications, with
their essential simplicity and which give a true picture of the fluid state of the species—
the distinctive races of which are being modified—is to treat each colony or group of
similar colonies in each area as a unit. Only homogeneous, pure-bred populations could
be considered as belonging to a rigidly defined subspecies. With this static concept there
would be many pockets of one subspecies within the range of another and wide zones of
hybridization.

IX. WESTERN AUSTRALIAN RECORDS.

Hemiergis decresiensis has been listed several times as occurring in Western
Australia. Records for this State should almost certainly be attributed to Hemiergis
tridactylum (Boulenger), an exclusively Western Australian form.

One reason for misidentification is probably that H. tridactylum was not described
until 1915, Boulenger (1915, p. 65), whereas the key in Boulenger’s catalogue, which
would be consulted for identification in most cases, was printed in 1887 and would
result in H. tridactylum being identified as H. decresiensis (1887, p. 223). The two
species are similar in being tridactyle, but in H. tridactylum midbody scale rows are
18 or 20 and the third toe is much longer than the second against 24 or 26 rows for
South Australian H. decresiensis, which have the second toe slightly longer than the
third. The only specimen of H. tridactylum I have seen, Australian Museum R.2454,
collected at Perth, is, among other characters, also separated from H. decresiensis by
colouration, incomplete differentiation of the transparent disc, and by the number of
subdigital lamellae, left, 6, 9, 11; right, 6, 9, 12. H. tridactylum is evidently extremely
common in some Western Australian localities. The Harvard Museum of Comparative
Zoology has 60 specimens collected at Augusta in 1927 by W. S. Brooks (Loveridge,
1934, p. 368).

Waite (1929, p. 161), who gives the range of H. decresiensis as ‘‘Western and
South Australia, Victoria and New South Wales”, may have followed Zietz (1920,
p. 216), who gives the same range.

Werner (1910, p. 481) possibly was the authority for these two authors. He had
five lizards from Lunenberg and Donnybrook, which he identified as H. decresiensis.
The specimens had 20 midbody scale rows, which suggest H. tridactylum. Lunenberg
and Donnybrook are in the same area as Yallingup, the type locality of H. tridactylum,
and Augusta, Margaret River, Wallcliffe and Manjimup, where it has been collected.

BY STEPHEN J. COPLAND. 91

Giinther (1867, p. 48), who separated his H. polylepis from H. decresiensis mainly
on the grounds that it had 26 rows of scales at midbody against 18 or 20 (see under
H.d. decresiensis) could not have had H. tridactylum as his material because he says
the toes of his specimens with 26 rows were “more developed” than in his type with
18 or 20.

It is a coincidence that Gray (1845, p. 87) originally noted the single British
Museum specimen of H. decresiensis as from the Swan River. The locality was corrected
in the same volume (p. 272) to Kangaroo Island.

Gtinther (1875, p. 18) evidently follows Gray’s earlier entry in giving the range of
H. decresiensis, which he attributes to Péron, as Swan River and Adelaide. Giinther’s
succeeding entry gives his H. polylepis as from South Australia (Kangaroo Island).

Mr. L. Glauert, Curator of the Western Australian Museum, wrote to me on
ist December, 1943: ‘“Hemiergis decresiensis is not represented in our collection, nor
does it occur in this State as far as I am aware, in spite of what certain lists say.
We have, however, a species Lygosoma (Hemiergis) tridactylum Boulenger, which was
originally described as a variety of peronii.”

XxX. ACKNOWLEDGEMENTS.

I wish to acknowledge help and advice from Professor W. J. Dakin and Professor
E. A. Briggs, of the University of Sydney; Dr. A. B. Walkom, Mr. J. R. Kinghorn and
Mr. T. Iredale, of the Australian Museum; and Mr. L. Glauert, of the Western
Australian Museum. Mr. Kinghorn, Mr. H. M. Hale, of the South Australian Museum,
and the late Mr. D. J. Mahony, formerly of the National Museum, kindly lent me
specimens. J have to thank Miss A. G. Burns, of the Department of Zoology, University
of Sydney, for the photography. Besides providing specimens, the late Dr. H. F. Consett
Davis was a source of ungrudging and invaluable encouragement.

XI. BIBLIOGRAPHY.

ALLEN, GLOVER, M., 1938.—The Mammals of China and Mongolia. Natural History of Central
Asia. New York, Vol. 1.

————,, 1940.—Id., Vol. 2.

BouLencer, G. A., 1887.—Catalogue of the Lizards in the British Museum. 2nd Ed., London.
Vol. 3.

——_——,, 1915.—-Descriptions of Two New Lizards from Australia. Ann. Mag. Nat. Hist..
(8) 16: 64-66. .

CuviER, BARON, 1829.—Le Régne Animal distribué d’aprés son organisation . .. Paris, Vol. 2.

9)
Davis, CONSETT, and LEE, D. J., 1944.—The Type Concept in Taxonomy. Aust. J. Sci., 7 (1):
16-18.

DoBZHANSKY, THEODOSIUS, 1937.—Genetics and the Origin of Species. New York.

DuMERIL. A. M. C., and Brpron, G., 1839.—Erpétologie générale, ou Histoire naturelle compléte
des Reptiles. Paris. Vol. 5.

FITzIncer, L. J. F. R., 1826.—Neue Classification der Reptilien nach ihren natiirlichen

Verwandtschaften ... Wien.

GRAY, JOHN Epwarp, 1831.—A Synopsis of the Species of the Class Reptilia in the Animal
Kingdom ... by the Baron Cuvier ... with Additional Descriptions... by E. Griffith...
and Others. London. Vol. 9.

a , 1839.—Catalogue of the Slender-tongued Saurians. ... Ann. Nat. Hist., London, 2:
287-293.

—, 1845.—Catalogue of the Specimens of Lizards in the Collection of the British Museum,
London.

GUNTHER, ALBERT, 1867.—Additions to the Knowledge of Australian Reptiles and Fishes. Anz.
Mag. Nat. Hist., (3) 20: 45-68.

————, 1875.—A List of the Saurians of Australia and New Zealand. The Zoology of the
Voyage of H.M.S. Erebus and Terror ... . during... 1839-43. London.

LONNBERG, EINAR, and ANDERSSON, L. G., 1913.—Results of Dr. E. Mjéberg’s Swedish Scientific
Expeditions to Australia, Part 8, Reptiles. K. svenska VetenskAkad. Handl., Stockholm,
yz) (33) 8 alii

LovERIDGE, ARTHUR, 1934.—Australian Reptiles in the Museum of Comparative Zodlogy,
Cambridge, Massachusetts. Bull. Mus. Comp. Zool., 77 (6): 243-283.

Lucas, A. H. S., and Frost, C., 1894.—The Lizards indigenous to Victoria. Proc. Roy. Soc.
Vict., 6 (mew series): 24-92.

————— and Le Sourr, W. H. DupLey, 1909.—The Animals of Australia (Mammals, Reptiles,
and Amphibians). Christchurch.

$2 VARIATION IN THE LIZARD HEMIERGIS DECRESIENSIS,

Mayr, Ernst, 1942.—Systematics and the Origin of Species. New York.

MERTENS, ROBERT, 1931.—Ablepharus boutonii (Desjardin) und seine geographische Variation.
Zool. Jb., 61: 63-210.

PARKER, H. W., 1926.—A New Lizard from South Australia. Ann. Mag. Nat. Hist., (9) 18:
203-205.

Pope, CLirrorD H., 1935.—The Reptiles of China: American Museum of Natural History, New
York: Vol. 10 of Natural History of Central Asia.

Proctor, JOAN B., 1923.—The Flora and Fauna of Nuyts Archipelago and the Investigator Group,
No. 5, The Lizards. Trans. Roy. Soc. S. Aust., 47: 79-81.

SmitH, Mautcotm A., 1937.—A Review of the Genus Lygosoma and its Allies. Rec. Ind. Mus.,
39 (3) 2 213-234.

STEINDACHNER, FRANZ, 1867.—Reptilien, in Reise der Osterreichischen Fregatte Novara um die
Erde in den Jahren 1857, 1858, 1859, Wien, Vol. 1.

Waite, E. R., 1927.—The Fauna of Kangaroo Island, South Australia, No. 3, the Reptiles and
Amphibians. Trans. Roy. Soc. 8S. Awst., 51: 326-329.

———, 1929.—-The Reptiles and Amphibians of South Australia. Adelaide.

WEEKES, H. CLairn, 1929.—On Placentation in Reptiles, No. 1. Proc. LINN. Soc. N.S.W., 54 (2):
34-60.

WERNER, FRANZ, 1910.—Reptilia (Geckonidae und Scincidae) in Die Fauna Stdwest-Australiens ;
Ergebnisse der Hamburger stidwest-australischen Forschungsreise 1905 herausgegeben von
... W. Michaelsen und... R. Hartmeyer, Jena, Vol. 2.

Zintz, F. R., 1920.—Catalogue of Australian Lizards. Rec. S. Aust. Mus., 1 (3): 181-228.

EXPLANATION OF PLATE VI.

Figs. 1-4.—Dorsal views of the four races of -Hemiergis decresiensis.
Fig. 1.—H.d. decresiensis, topotype, No. R.2191.

Fig. 2.—H.d. continentis, holotype, No. R.2190.

Fig. 3.—H.d. talbingoensis, holotype, No. A.C.2081.

Fig. 4.—H.d. davisi, holotype, No. A.C.821.

Body lengths of specimens are 45, 52, 60 and 60 mm. respectively.

Photos.—Miss A. G. Burns.

93

REVISION OF AUSTRALIAN LEPIDOPTERA. OECOPHORIDAE. XIII.

By A. JEFFERIS TURNER, M.D., F.R.E.S.

[Read 25th July, 1945.]

Two species were omitted in Part xii.

1875.

PHILOBOTA MICRANEPSIA Turn., Proc. Roy. Soc. Tasm., 1926, p. 153.

(Cradle

Mt., Lake Fenton.)

complex consisting in a main stem with numerous branches.

1876.7 PHiLogora Tropica Meyr., Hxot. Micro., i, p. 250.

( Darwin.)

Most of the Australian Oecophoridae are contained in the great Hulechria-Philobota

The present instalment

consists of a group of genera outside this complex, to which their relationship, though

close, appears to be collateral rather than direct.
Machimia complex.

They may be conveniently termed the
Generally speaking they have a common facies, are often more or

less rosy or red in their colouring, and frequently have vein 5 of the hindwings curved
and approximated to 4 at its origin.

bo

vo

=

M

Key to Genera.

A DiMWAL Ha DICAlMInTehiOon tute mom SCCONG joint 62... «qcsiore eres is cio eo cle ele wis sleveaias 1
eM Oem © tam ELI EC Clamlep swe ste feats eek teat cL is, oi Gsista, opis eeiate, clnyer ails eyisi opisneim tsicbris, Senta’ oc arta ss edeyautebeuesevistelie leh a) Seaqemeens 10
al pimwi Unwcerimun alm OlMt Stouts cayenne creierancheusiaacke te love) Scie wie ees crichisrey suede cust siete ye aus aielene Alderete 3
AVA mayVpGlmeKc CITIT allies] OUN ta SIEM Cr 7 aie, sie sheconsyegeeic is, avers sects oot a cayeasreue auie lcberlabiedersite sbapansbapéaewevrewtas 4
Palpi with second joint very long and thickly rough-scaled throughout ........ Polyzeucta
Palpi with second joint moderate, in basal half slender ................... Arachnographa
Pindwineswithaoeand, ” approximated or stalked (0.5.0.2 602c5-+ee ees os Heteroptolis
nin dwyine Smile OnanGauieseparatessperalleliseepeieiecis cieicrers aicicherciciecosuctelcuecestencns ciel s shel aia tues 5
PAO Balen UOeaMieNnl gooe HE toe He Gael sscosaeapenouusuceoucneuboooodndubuesuoo.s 6
Pall pimwikheverminaleyoint iGhoGgeerniwthannSeCONG: os si eis ce ches sc oe eiel cele el ensie le els se = = clejepens rt
AMITERIOMMUI OT VemUNICKEMEGE suazals ses lteii a Bh lol cece otoreual atau eiysl,c cits utesie coeugue) wlede teice ues ehaisle ocerehele Hybocrossa
ANTICC IT OGM UD LAC wert O tarbh COM CGE arcs a. sisi cdecaisdene reticle och ens) suche isl cr(ero. sus) eherswecleyccaysstlele\euelallens Tortricopsis
PAMIECTINGA emule OU I CCLETI casein os torcnsnete) mistsyec et cle merensy op ale .cesojieyeyeu sie nos oitessehie) sence sen sei Gaeyetevesce) 6 Phyllophanes
PANGCTIMNGA CMR WAC e CLE TION meristci cote Seat)! auClea set auen crane eis le ei eras lat rar tale uelnipve rete eek een eccl ican onde seus tela ts tte 8
alpiewathsecond joint not reaching base of antennae .........-2 02.55... -5-... Clonitica
Palpimwathasecond= joint reaching, basevofeantennae 4.4.5... +5 e020 cue gee tie sem oo seen 9
I2Allon, Walldn | Wwersrovioeyl afore GoOSeYbe Voye WES: Sab oadegcogeodnodedoceu Lucu OB eoCBUe Zelotechna
alpimwALmcernain ales OINteamoremthany One-half ss cies ccs ele pee is cuctersie oie eiels cere «alec Wingia
HORE w ine Siawi thier om UOULCEIMNGM cscs, 3 cicl-. scsieuaererettarraencutienocevep ais. eersist rl ehevcUsucxevenouels, ote) oom 6 sire Callizyga
LORE WANE Smawil Glee Saat OM COSA marie sete rcs aic, cilai.cte Stemi Noon ore rslic tie anlar ueelOs, emerLee obama wele tine ast auaradinaaseapegts ilal
Eimnchwin Simp LOACe re sth ane hORe wil 2S) b-esrieetronsiebeielcme cies cassie creee easter sie cima erelse sels osciennenclints 12
ISHMCHIMOES MOE leo Kee WENO toyAen Abo So5odbavcoduboouuububosodeudeudodooouoEuoUdUS IZ
Palpiawirhetermin alin Olntnextremely ish Onteeeeees ec | tao cicero cleie ceteris e eens tiers Thyromorpha
al pimwathmrermin ala) OlmntmModerate on 1Onewemmniacias ocd cle cules) a cicicbessnewenees) e cocci ealoieemoieners 3
ATCT ACM Wil eppeCUCIin sewaratarcn ate ieso.cach ay deh eCMe oes Naten a iecelicc oie emelolaiatana acre con cum sqhltnre corepeivel a: eye! Anthocoma
PANITTCNIN ACR WAU OME De GLEN) eustetla tein ear eth ol arate ns emcee t bale tite op tie ohalie ch aobevsuahianale \dveiterave ireoa eateries tara melane 14
NMI MNAE) yer JooEVI NEnsonU CENA tame miBI EL eIs tet cl accliG- th sinececrcne eaakt es olcicie ercnein a oan ence eo aI Huprionocera
ANGennaewOtunNaletn ote Tami Mate ta: eis cd aga aecast sha eee ea seal oilamatatton ene tel ah calavesa cen shauen suai gubuie vets le. haibaikenals 15
Palpi with second joint rough anteriorly beyond middle ...................... Ancistrodes
pian Gh SECON Gas ON tsi OOthey tistics attains oes Hue cer cits rede) nga yatale juste utc osy ot. etic meet pete 16
ANTAL SROEREYE (ONE TOON Wey. Str 0 aT ON VES eal Sse ee ae sie i Sy Tok AR Oe aA a ee Nea Gymnoceros
ANTUCEIDIDENS Ce TOME IEY TAKONEN Spiro a OV eV ee eee cucecl ito ey oer oer intaro. Geko Een Mic ees cee Roe Platyphanes
ANIME OTE. MONEDA SCE OK EXd La yelp ace a UEC aes 1 OP RRO ORES SORE CTE Cais anc arn DE er tse AF 18
ANTTUCETPHG HO: TEN OMENS). -TOKONE 10) WK) Sy VEG ly ene Gore eRe ic On, Linear Geena ere otter ORCI ETE Oa Ercan a aie ee eee 19
TOGO Wil “ovr Ore ieeVIeaul Been: Ae ocoooopduoeedeupobodoosodoBbodoUobouDS Habroscopa
HORE VINES Six. @ O tliereds pees hisoceaien eeu pa tapa tlic rapes Sey stave nel ate cies oh ayhieyewatinn ucla s “scoucerfonreriawellayonls tahicts Lepidotarsa
HOnewiANne Sevan mall due SieC OLNCIO SME oiz-c5/- panels oteehe) sneleier eh sili siaten-)ie cl peieeers cle ete ees = Atelosticha
ORE WAN 2 Siew t iene tevn) Cie SiS Le allk CClmtewaie ate wt eicetse ayn. sa seer easton, Satcten aulcuray ens kara taba Bicone seieie aueainen rand! 20
ORE INES Alea WOU Oe TENE) Sea Gebio ooscansoob6hb0coouhbebcoesoudaouEs Lophopepla
FSOLEW ATS Sy SIN OO Ckteertwey cae Uae ey eee lenses ia cel nave ua alias wah ATA GTA & fos acchenedeney bs Reakel aba tein terete out veus/aicbeecreeet 21

94 REVISION OF AUSTRALIAN LEPIDOPTERA. OECOPHORIDAE. XIII,

210 Pal pisswithmtenminales) ointone-hallisoneleSSmrarsrtelelenieteaaie n-ne iee- noel aei mete ietal te enema nee ae 22
Palpiewithuerminal joint mMonremchanrone—lelliterer-aiieaenee- lenis it-seraiene lanai teen net a 25
22. Palpi with second joint not reaching base of antennae .......:............... Pyricausta
Pallipignwitheseconds)ointeneachinegb aSe moi ea miUemi ae mrenewetei aimee iene te eet etree ae eer eee 23
PE) LUMO AEM AMAT NOM PA sd OLGA KOLOR IVAN Go moomoo Oe oudGoda sada oidaandnooucens sé 005 Aglaodes
Morewinges) with2 front mean ‘angeles: 2 sceva age wer tuetele dersistairs ellen epecetattepene operons: Pienceete laine Aronson 24
24). Antennae: of Mmalerisimple sac sees avs soles eee eee eration ei dacene NeoPa enemen Hehe eal oameEn es Callithauma
Antennae: Of MAleveiiat Sd! es ecw ele lea, Ses eee eee ee ro eee eRe dercir eeeee ie nena eu one Tisobarica
25. sDhOrax swith POSteriOr rCRESUs caacwy-- he ono tie cee een yen ee te ieee RCI Re ae nen eee Eclecta
Litt Xo) ar: bo ames) cLOY Oo) el « uae rae tener ne Lerner a ee URE Rtas Pen acc tS olan GO biG Oo Bd oldb laa @bics 26
26. Palpi loose-scaled anteriorly with median thickening; ..-.5.....3....4.+-.6-5-:++-+s000- 27
f t2 1 8) ati a (011 Sh 0 ee a rane ra Perea at eRe IMaA LICR E Wace cau So eaIOIe ona RRO IO OOO ole OUlord om O D OI0.0.0 66 28
2s BOrewinesy Wwithy 1) vO COSCA cree cui. s eas erateneene ea ci eue Nomen ec lic rene) aac RR cnCa nme Ema ae eee ELuphiltra
Morewinges: wath tor termr em sachs cencye ancien op aie oo Wena cian eee ete eta RAPS en ci cena Zonopetala
28. Palpi with second joint not reaching base of antennae ....................... Parocystola
Palpisawith) second joints reachines base) ot antennae a eae ene 29
29% sPalpiewithwsecond joint Lurrowedsanternlorlivaaeuenierlste eet een ein nee nen Archaereta
L242) 9 op ile 0X0) Cans lo Vaan eirrast nen eae Rei teen ieee eA tener An) Si rats inanbo nin F Dench miata cata) ord Gace Ord. oO DS OlOd- Gb o-6lc 30
308 SMorewinges: with) ((r8i2 GeStal Keays 5 tes. cy Ware oes OMe error est cen UES es ce ee en 3
MOLE WANES) i withie 9) SCWAT AEC gs Pees ot nccieclecoreosh te tsaee i oe erties ere ic ees end genre leer sos mA aoe cli Pa 32
ol. —Palpi with second joint) notereachines basco vamvennaley yaaa) acus aieeeeinien renee Macrophara
Palpi with second joint reaching base of antennae .........................- Epicharactis
324 eosterion tarsis withe basaleyoint eloneateree acetate ae oie ee eee ee Leistarcha
Posterior: tarsivnonmeall (25 ic dts fuel ole oi MO ae lens le tah hel cRORSA SR aie aed Ree ALU AeA): ER IRE ee 3
Bey Jebiach bass waldo by Enaorropabnmeycercl: ie 4) Ene Grensibn Godooacecasccob0dconodaubogosoanoboOSUS 3
Tabhoohynbayers: Walle. i) TOE Eyojocopahonencol Wor 2) So colwoddgoocobonouncaonducs ado un ooeubooobOUS 38
34, Horewines with tvermen! Sinwaen ers cisiisencntenieiereia aces pugee Mee es eeeucint aes te eare ote ake Colpoloma
Morewines: with termen not simulate: feria cele choses ve tenee eee eS eS Cee ee net 35
i, IEEWho Valin Weoley! Folie AS Noms AS SCCM! GecasbuccnsosucucccooononLe boos Compsotropha
Palpiwitheterninalliey oiniteshOncersth<inmSe COM Cimersiiaelcieieee rience asic men ne neieme nein nie a ene 36
365 sealpiiswithy terminals j omit iStoutees cia crso cence nene ce ee nee a inn erenene Pycnozancla
Palpi-with terminal: jointsslen@er. i: 02..7..02. 2a do nee rte hee TERE SOL ee Ie eee BT
37. Forewings with a fan-shaped tuft of scales on dorsum ....................-- Ptycholoma
Morewings! without GOrsal HCUEt | is he psic ce ee ahceamere een eal! elt oe ae eT ERE IE aimee Machimia
38) Palpiiwithe second joint roughy anterlionrivs Saar re eee eee saan eee Lepidozancla
iPalpiti witht: second) Joint: Smooth 2a Ca revo deer niche isos ee ae ERE NG ona aN Seema rah orem ele aie 39
392) Palpiwith: terminal joint ‘stout. ssa Aes soa che lettre eect Pelee Pica ee Usa anes em Urs Eochrois
Palpiawith terminals jointsslend eres qc sic eee ee eee eae ere Hoplomorpha

134. Gen. PoLyzeucTa Turn.

Trans. Roy. Soc. S. Aust., 1917, p. 104.

Palpi long ascending, somewhat recurved; second joint very long, greatly thickened
by dense hairs on lower surface throughout, ending in a rounded apical inferior tuft;
terminal joint stout, obtusely pointed. Antennae with strong pecten. Forewings with
7 and 8 stalked, 7 to termen. Hindwings normal. Monotypical.

1877. PoLyzEUCTA CALLIMORPHA Low., Trans. Roy. Soc. 8S. Aust., 1894, p. 93. (Cairns.)

135. Gen. ARACHNOGRAPHA Meyr.

IHGO, MOCO, Mg 1s ABZ

Palpi moderate, ascending, recurved; second joint reaching base of antennae,
gradually expanding from base to form a rounded apical inferior tuft; terminal joint
stout, pointed. Antennae with strong pecten. Hindwings with 5 curved and approxi-
mated to 4 at origin. Monotypical.

1878. ARACHNOGRAPHA MICRASTELLA Meyr., Proc. Linn. Soc. N.S.W., 1882, p. 433.
(Sydney, Melbourne, Beaconsfield, Gisborne, Tasmania, Mt. Lofty.) Larvae on Hxocarpus
cupressiformis (according to Meyrick) and Juniperus hibernicus (according to N. Geary).
A bred series shows that the white markings are inconstant.

136. Gen. Hrerrroprotis Meyr.
HOt MaACnOn le Ds 2 aie
Palpi moderate, ascending, recurved; second joint reaching base of antennae, with
a long triangular apical inferior tuft; terminal joint less than one-half, very slender,
acute. Antennae with basal pecten. Forewings with 7 and 8 stalked, 7 to termen.
Hindwings with 6 and 7 stalked or closely approximated. Monotypical.
1879. Hererrroprovis LrEUcosTA Low., Trans. Roy. Soc. S. Aust., 1892, p. 10. (Mt. Lofty.)

BY A. JEFFERIS TURNER. 95

137. Gen. Hyspocrossa Turn.

Trans. Roy. Soc. S. Aust., 1917, p. 105.

Palpi moderate, ascending, recurved; second joint reaching base of antennae, with
acute apical inferior tuft; terminal joint as long as second, slender, acute. Antennae
without pecten. Anterior tibiae and tarsi thickened with dense scales. Forewings with
7 and 8 stalked, 7 to termen. Hindwings normal. Monotypical.

1880. HyBocrossa PARATYPA Turn., Trans. Roy. Soc. S. Aust., 1917, p. 105. (Macpher-
son Rge., Uki, Sydney.)

138. Gen. Torrricopsis Newm.

Trans. Ent. Soc., 1855, p. 293. Meyr., Gen. Ins., Oecoph., p. 142.

Palpi moderate, ascending, recurved; second joint reaching base of antennae, with
an apical inferior tuft; terminal joint as long as second, slender, acute. Antennae with
pecten. Hindwings with 7 and 8 stalked, 7 to termen. Hindwings normal. Type,
T. uncinata. . Hleven species.

1881. TortTRICOPSIS AULACOIS Meyr., Proc. Linn. Soc. N.S.W., 1882, p. 438. Gen. Ins.,
Oecoph., Pl. 5, f. 92. (Stanthorpe, Toowoomba, Murrurundi, Bathurst, Mittagong,
Sydney, Narracan, Geelong.)

1882. TORTRICOPSIS UNCINELLA Zel., Lin. Hnt., ix, p. 355. Meyr., Proc. Linn. Soc.
N.S.W., 1882, p. 437. (Brisbane to Tasmania, Mt. Lofty, Pt. Lincoln, Kangaroo I.)

1883. TorRTRICOPSIS ERYTHRURA Meyr., Hxot. Micro., i, p. 228. (Cairns.)

1884. ToRTRICOPSIS SEMIJUNCTELLA WIk., xxix, p. 777. Meyr., Proc. Linn. Soc. N.S.W.,
1882, p. 436. (Atherton, Duaringa to Melbourne, Milmerran, Bathurst, Mt. Lofty,
Ardrossan, Waroona, Perth.) ;

1885. TorTRICOPSIS PYROPTIS Meyr., Trans. Roy. Soc. S. Aust., 1902, p. 143. (Yeppoon
to Melbourne, Stanthorpe, Scone.)

1886. ToRTRICOPSIS EURYPHANELLA Meyr., Proc. Linn. Soc. N.S.W., 1882, p. 435.
(Victoria, Tasmania. )

1887.7 ToRTRICOPSIS HESYCHAEA Meyr., ibid., 1885, p. 826. (Mt. Kosciusko.)

1888. ToRTRICOPSIS MESOPHTHORA Meyr., ibid., 1885, p. 825. (Tasmania. )

1889. ToRTRICHOPSIS TARACHODES Turn., Proc. Roy. Soc. Qd., 1919, D. 171. (Hidsvold.)

1890. ToRTRICOPSIS CROCOPEPLA, N. Sp. (xpokomem)os, clothed in saffron.)

9. 22-23 mm. Head and thorax pale orange. Palpi with tuft as long as second
joint; grey, upper edge ochreocus-whitish. Antennae whitish annulated with pale
fuscous. Abdomen grey; apices of segments and tuft ochreous-whitish. Legs dull
reddish; posterior pair whitish. Forewings dilated posteriorly, costa arched to middle,
thence straight, apex rounded-rectangular, termen sinuate, not oblique; pale orange;
markings brownish; stigmata minute, first discal at one-fourth, plical slightly beyond it,
second discal slightly beyond middle; three fine transverse lines, first from one-fourth
costa to two-fifths dorsum, second from midcosta to three-fourths dorsum, third oblique
from three-fourths costa, soon bent to become subterminal to before tornus; cilia grey,
extreme apices whitish. Hindwings and cilia grey-whitish.

West Australia: Albany in November; two specimens.

1891. TorRTRICOPSIS ERYTHROPEPLA, 1. Sp. (€pv@pomemdos, in reddish clothing.)

9. 22 mm. Head and thorax dull reddish. Palpi with tuft as long as second joint;
dull reddish, upper edge whitish. Antennae whitish with narrow fuscous annulations.
Abdomen whitish-grey. Legs dull reddish; posterior tarsi whitish. Forewings slightly
dilated, costa straight except near base, apex rounded-rectangular, termen slightly
sinuate, not oblique; dull reddish; costa narrowly whitish-ochreous with two minute
fuscous dots, first at middle, second at three-fourths, slightly oblique; cilia reddish-
fuscous, extreme apices grey-whitish. Hindwings and cilia whitish.

West Australia: Nornalup in November; one specimen.

139. Gen. PHYLLOPHANES Turn.
Trans. Roy. Soc. 8. Aust., 1896, p. 21.
Palpi long, ascending, recurved; second joint very long, more than three times
length of face with pointed apical inferior tuft one-third length of joint; terminal joint

96 REVISION OF AUSTRALIAN LEPIDOPTERA. OECOPHORIDAE. XIII,

one-half, slender, acute. Antennae without pecten. Forewings with tuft of long scales
on costa shortly before middle; 7 and 8 stalked, 7 to termen. Hindwings with long
projecting cilia at apex; 5 approximated to 4 at origin. Monotypical.

1892. PHYLLOPHANES DYSEURETA Turn., Trans. Roy. Soc. S. Aust., 1896, p. 21.
(Brisbane. )

140. Gen. Cronitica Meyr.

Hxot. Micro., i, p. 2338.

Palpi with second joint not reaching base of antennae, thickened with dense scales,
rough and projecting, in a very short tuft anteriorly towards apex beneath; terminal
joint less than half second, slender acute. Antennae with pecten. Forewings with 7 and
8 stalked, 7 to termen. Hindwings with 5 from middle. Monotypical.

1893.7 CLoNITICA EUSARCA Meyr., Trans. Roy. Soc. S. Aust., 1902, p. 144. (Quorn.)

141. Gen. ZeLorecHNA Meyr.

Hots MUCiOn mien pse2 22.

Palpi long, obliquely porrect: second joint very long, expanded, with hairs above
and beneath and with apical inferior tuft; terminal joint very slender, short
(one-fourth to one-half), acute. Antennae with pecten. Forewings with 7 and 8 stalked,
7 to termen. Hindwings normal. Type, Z. falcifera. Seven species.

1894. ZELOTECHNA LITHOCOSMA Meyr., Proc. Linn. Soc. N.S.W., 1885, p. 827 (Sydney,
Melbourne, Grampians, Dimboola, Perth, Mogumber.)

1895.7 ZELOTECHNA SIGMASTROPHA Low., ibid., 1898, p. 51. (Kangaroo I.)

1896. ZELOTECHNA CALLICHROA Meyr., Trans. Roy. Soc. S. Aust., 1902, p. 144.
(S. Aust.: Sheringa, Pt. Lincoln.)

1897. ZELOTECIINA FALCIFERA Meyr., Proc. Linn. Soc. N.S.W., 1882, p. 440. Gen. Ins.,
Oecoph., Pl. 5, f. 90. (Sydney, St. Helens, Yanchep.)

1898.7 ZELOTECHNA WIRAX Meyr., ibid., 1882, p. 441. (Sydney.)

1899. ZELOTECHNA PSITTACODES Turn., Trans. Roy. Soc. S. Aust., 1917, p. 103. (Tweed
Hds. )

: 1900. ZeLorecHNA sSTICHOPTIS Low., Proc. Linn. Soc. N.S.W., 1915, p. 481. (Broken
Hill.)

142. Gen. Winera Wlsm.

Cat. Oxf. Mus., ii, p. 552.

Palpi moderate, ascending, recurved; second joint reaching base of antennae, with
an apical inferior tuft; terminal joint shorter than second, slender, acute. Antennae
with pecten. Forewings with 7 and 8 stalked, 7 to termen. Hindwings normal. Type,
W. lambertella. Ten species:

1901. WiINGIA AURATA WIK., xxix, p. 775, Meyr., Proc. Linn. Soc. N.-S:W., 1882,
p. 427. (Atherton, Brisbane to Tasmania, Macpherson Rge., Katoomba, Cunnamulla,
Mt. Lofty, Ardrossan, Mt. Barker, Waroona, Lake Grace.) ;

1902. WINGIA LAMBERTELLA Wing., Proc. Roy. Soc., 1849, p. 105. Meyr., Proc. Linn.
Soc. N.S.W., 1882, p. 428. Gen. Ins., Oecoph., Pl. 5, f. 91. (Scone, Neweastile, Sydney,
Gisborne, Dunkeld, Bendigo, Lake Grace, Waroona.)

1903. WINGIA RECTIORELLA WIK., xxix, p. 775. Meyr., Proc. Linn. Soc. N.S.W., 1882,
p. 430. = aurigena WIk., xxix, p. 775. = confectella Wlk., xxix, p. 776. = euryptera Turn.,
Trans. Roy. Soc. S. Aust., 1896, p. 3. (Noosa to Moe, Macpherson Rge.)

1904. WINGIA HESPERIDELLA Meyr., Proc. Linn. Soc. N.S.W., 1882, p. 429. (Sydney,
Katoomba, Melbourne, Sale, Tasmania.)

1905. WINGIA THEOPHILA Meyr., ibid., 1885, p. 825. (Deloraine.)

1906. WINGIA LECHRIOZONA, n. Sp. (Aexptogwvos, Obliquely banded.)

2. 22 mm. Head whitish. Palpi with second joint reaching base of antennae,
terminal joint one-half; pale fuscous, terminal joint and apex of second whitish. Antennae
whitish. Thorax pale brownish. Abdomen pale grey. Legs ochreous-whitish; anterior
pair except tarsi pale fuscous. Forewings arched to one-fourth, thence straight, apex
pointed, termen obliquely rounded; pale brownish; markings fuscous; a narrow oblique
fascia from one-third costa to near mid-dorsum; several minute costal dots beyond

BY A. JEFFERIS TURNER. 97

middle; a series of minute dots on termen; cilia whitish, beneath apex fuscous. Hind-
wings and cilia white.

Tasmania: Cradle Mt. (3,000 ft.) in January (W. B. Barnard); one specimen.

1907.; WINGIA DOROPHANES Meyr., Hxot. Micro., i, p. 194. (Zeehan.)

1908. WINGIA THALAMIA Meyr., Proc. Linn. Soc. N.S.W., 1882, p. 482 = synnephela
Turn., Trans. Roy. S. Aust., 1917, p. 108. (Stanthorpe, Barrington Tops, Katoomba,
Wangaratta, Glenrowan.)

1909. WINGIA SUBROSEA Turn., Trans. Roy. Soc. S. Aust., 1894, p. 134. (Strad-
broke I.)

1910. WINGIA EONEPHELA Meyr., Proc. Linn. Soc. N.S.W., 1882, p. 432. (Nambour
to Sydney, Murrurundi, Bathurst.)

143. Gen. CaLLizyea Turn.

Trans. Roy. Soc. S. Aust., 1894, p. 132. Proc. Linn. Soc. N:S.W., 1913, p. 208.

Palpi ascending, recurved; second joint reaching base of antennae, much thickened
with appressed scales; terminal joint shorter, but nearly as stout as second, tolerably
pointed. Antennae without pecten. Forewings with 7 and 8 stalked, 8 to termen.
Hindwings broader than forewings; neuration normal. Monotypical.

1911. CaLtizyea DISPAR Turn., Trans.. Roy. Soc. S. Aust., 1894; p. 132. (Cairns,
Atherton, Brisbane, Tweed Hds., Lismore.)

144. Gen. THyromMorRPHA Turn.

Trans. Roy. Soc. S. Aust., 1917, p. 108.

Palpi ascending, recurved; second joint reaching base of antennae, much thickened
with evenly appressed scales; terminal joint less than half second, tolerably slender,
acute. Antennae without pecten. Forewings with 7 and 8 stalked, 7 to termen. Hind-
wings broader than forewings; 3 and 4 stalked, 5 slightly curved and approximated to
4 at origin. Monotypical.

1912. THYROMORPHA STIBAROPIS Turn., Trans. Roy. Soc. S. Aust., 1917, p. 108.
(Brisbane, Macpherson Rge., Lismore.)

145. Gen. ANTHOCOMA, n.g. (dvdoxouos, flowery.)

Palpi long, ascending, recurved; second joint reaching base of antennae, thickened
and roughened anteriorly towards apex; terminal joint as long as second, slender, acute.
Antennae with pecten. Forewings with 2 and 3 stalked, 7 and 8 stalked, 7 to termen.
Hindwings broader than forewings; 5 curved and approximated to 4 at origin.

1913. ANTHOCOMA EUTERPNES, Nl. Sp. (evrepmrvyns, charming.)

6. 30 mm. Head brown. Palpi whitish, outer surface of second joint except
extreme apex blackish. Antennae ochreous-whitish with pinkish-grey annulations;
ciliations in male 5. Thorax brownish faintly tinged with pink. Abdomen ochreous.
Legs ochreous; tarsal rings blackish mixed with rosy; anterior tibiae and tarsi blackish.
Forewings oval, costa strongly arched, apex rounded, termen very obliquely rounded;
pale yellow; basal fourth of costal edge blackish; a broad rosy stripe on costa from
near base to middle, thence narrowing and receding from costa, ending beneath three-
fourths costa; a similar dorsal stripe from base to middle; an elongate rosy suffusion
beneath middle from two-thirds to near termen; cilia pale yellow, beneath apex rosy-
fuscous. Hindwings pale yellow; a broad rosy median suffusion not reaching base;
cilia pale yellow. Underside of forewings bright rosy surrounded by a marginal ring
of pale yellow.

Queensland: Mt. Tamborine in November; one specimen.

146. Gen. EupRIONOCERA Turn.

Trans. Roy. Soc. S. Aust., 1896, p. 6. Proc. Linn. Soc: .N:S:W., 1917, p..101.

Palpi ascending, recurved; second joint reaching base of antennae, moderately
thickened, smooth; terminal joint shorter than second, slender, acute. Antennae without
pecten; in male thickened, shortly laminate, with very short ciliations. Forewings with
2 and 3 stalked, 7 and 8 stalked, 7 to termen. -Hindwings broader than forewings;
neuration normal. Monotypical.

98 REVISION OF AUSTRALIAN LEPIDOPTERA. OECOPHORIDAE. XIII,

1914. EupRIONOCERA GEMINIPUNCTA Turn., Trans. Roy. Soc. S. Aust., 1896, p. §.
(Brisbane, Milmerran, Scone, Sydney.)

147. Gen. ANCISTRODES, N.g. (ayKioTtpwins, hooked.)

Palpi long, ascending, recurved; second joint exceeding base of antennae, thickened
and shortly rough-sealed anteriorly beyond middle; terminal joint shorter than second,
slender, acute. Antennae without pecten. Forewings with apex produced and faleate;
7 and 8 stalked, 7 to termen. Hindwings broader than forewings; 5 approximated to 4
at origin.

1915. ANCISTRODES PHRYGANOPHANES, Nl. SP. (puvyavodharys, like a dried stick.)

3, °. 30-34 mm. Head whitish-ochreous. Palpi whitish-ochreous; external surface
of second joint except apex dark brown. Antennae whitish-ochreous; ciliations in male
1. Abdomen brown; apices of segments and tuft whitish-ochreous; underside red. Legs
red; anterior pair dark fuscous. Forewings dilated posteriorly, costa strongly arched,
apex produced, faleate, termen sinuate, not oblique; pale brown, stigmata whitish out-
lined with fuscous; first discal at one-third, plical beyond it, second discal before two-
thirds, larger; a dark fuscous terminal line; cilia white, a median line and apices
fuscous. Hindwings and cilia pale ochreous.

Queensland: Macpherson Rge. (3,000 ft.) in October; four specimens.

148. Gen. GYMNOCEROS, D.g. (yuLvoKEpws, smooth-horned. )

Palpi long, ascending, recurved; second joint exceeding base of antennae, moderately
thickened, smooth; terminal joint shorter than second, slender, acute. Antennae without
pecten; in male thickened, simple. Forewings with 7 and 8 stalked, 7 to termen. MHind-
wings broader than forewings; 5 curved towards 4 at origin. In spite of the absence
of antennal ciliations this genus should be referred here. Callithauma is an analogous
case.

1916. GYMNOCEROS PALLIDULA, n. Sp. (pallidulus, pale.)

3. 28-30 mm. Head and thorax pale pinkish-grey. Palpi with terminal joint three-
fourths; grey-whitish. Antennae grey-whitish. Abdomen whitish. Legs whitish;
anterior pair grey. Forewings moderately dilated, costa moderately arched, apex
rectangular, termen slightly rounded, not oblique; pale pinkish-grey with numerous
scattered minute fuscous dots; costa pale pinkish with extreme edge whitish; a small
fuscous median discal spot at three-fifths; cilia whitish. Hindwings and cilia whitish.

Queensland: Macpherson Rge. in October; two specimens.

149. Gen. PLATYPHYLLA, n.g. (mAatrvdvAdos, broad-winged.)

Palpi ascending, recurved; second joint exceeding base of antennae, evenly smooth-
sealed; terminal joint as long as or shorter than second, slender, acute. Antennae
without pecten. Forewings with 7 and 8 stalked, 7 to termen. Hindwings broader
than forewings, 5 approximated to 4 at origin. The male may show additional characters.
Type, P. zophosphena.

1917. PLATYPHYLLA ZOPHOSPHENA, N. Sp. (odoodynvoes, with dark wedge.)

9. 30 mm. Head white. Palpi with terminal joint 1; white, base of outer surface
of second joint blackish. Antennae fuscous. Thorax rosy-fuscous; patagia white.
Abdomen grey. Legs fuscous; posterior pair grey-whitish. Forewings with costa
strongly arched, apex rounded, termen rounded, not oblique; blackish dots on bases of
costa and dorsum; a large blackish wedge edged whitish, extending on costa from one-
fifth to near middle, oblique, posterior edge excavated, ending in a rounded apex
shortly above mid-dorsum; two blackish whitish-edged dots placed transversely in middle
of disc; a slender outwardly curved slightly wavy fuscous line from two-thirds costa
to three-fourths dorsum; minute blackish dots on termen and apical fourth of costa;
cilia whitish-grey. Hindwings and cilia grey.

Queensland: Macpherson Rge. in January; one specimen.

1918. PLATYPHYLLA NIPHOLEUCA, nN. Sp. (vidodevxKos, SNOW-white.)

9. 32mm. Head and thorax pale grey. Palpi, abdomen, and legs white. Antennae
whitish. Forewings suboblong, costa moderately arched, apex rectangular, termen
slightly rounded, not oblique; white; cilia white. Hindwings and cilia white.

BY A. JEFFERIS TURNER. 99

New South Wales: Maryland, near Stanthorpe, Queensland, in November (W. B.
Barnard); two specimens. Type in Queensland Museum.

150. Gen. DIAPHORODES, D.g. (dtadopwons, unlike.)

Palpi ascending, recurved; second joint reaching base of antennae, thickened with
smoothly appressed scales; terminal joint shorter, slender, acute. Antennae without
pecten; in male ciliated. Forewings with 2 and 3 stalked, 7 and 8 coincident. MHind-
wings with 5 from middle of cell. An isolated genus of uncertain affinity.

1919. DIAPHORODES EURYSCIA, N. Sp. (evpvaxios, broadly shaded.)

6. 15 mm. Head and thorax white. Palpi with terminal joint two-thirds; white.
Antennae grey, near base whitish; ciliations in male 1. Abdomen grey-whitish. Legs
white; anterior pair grey. Forewings oblong, costa strongly arched at base, thence
straight, apex rectangular, termen straight, not oblique; white densely suffused with
ochreous-fuscous except near base and along costa; four minute costal dots from shortly
before middle to near apex; a suffused dark fuscous tornal spot; cilia white, bases grey,
on tornus fuscous. Hindwings whitish with a fuscous terminal line; cilia whitish.

North Queensland: Cape York in April (W. B. Barnard); one specimen.

151. Gen. Hasproscopa Meyr.

Haot. Micro., i, p. 223.

Palpi slender, ascending, recurved; second joint reaching base of antennae, some-
times rough-scaled near apex anteriorly; terminal joint as long as or shorter than second,
slender, acute. Antennae with pecten. Anterior tibiae and tarsi dilated with scales.
Forewings with tufts of raised scales; 7 and 8 stalked, 7 to termen. Hindwings with
5 from middle of cell.

1920. HaBroscopa IRIODES Meyr., Proc. Linn. Soc. N.S.W., 1883, p. 365. (Brisbane,
Gosford, Sydney.)

1921. HaABROSCOPA DICTYOSEMA, N. Sp. (OdektTvogonuos, Marked with a network.)

3, 2. 14-18 mm. Head grey; face white. Palpi with second joint reaching base
of antennae, terminal joint 1; grey, anterior edge except a subterminal ring in second
joint white. Thorax grey. Abdomen whitish. Legs white; inner surface of anterior
pair grey. Forewings with costa incurved from middle to apex, apex pointed, termen
straight, slightly oblique; brownish-grey; a broad basal fascia, white with patchy fuscous
irroration; edged by a fuscous line from one-third costa to mid-dorsum; a median costal
erest; a fine white line from midcosta to tornus, angled in middle; a similar line from
just beyond this to midtermen; this is crossed by a line from angle of first line to
five-sixths costa; a fine white line on apical half of costa and all termen, leaving costal
and terminal edge fuscous; cilia fuscous barred with white. Hindwings and cilia pale
grey.

North Queensland: Cape York in November (W. B. Barnard); three specimens. Type
in Queensland Museum.

152. Gen. Lrepmporarsa Meyr.

Proc. Linn. Soc. N.S.W., 1882, p. 446.

Palpi ascending, recurved; second joint reaching base of antennae; terminal joint
shorter than second, slender, acute. Antennae with pecten. Anterior tibiae and tarsi
thickened with scales. Forewings with 7 and 8 stalked, 7 to termen. Hindwings with 5
from middle of cell. Type, L. chrysopoca Meyr. Eleven species.

1922. LEPIDOTARSA NICETES, N. sp. (vexnrys, victorious. )

dg. 23-24 mm. Head orange-red. Palpi with terminal joint one-third; orange-red.
Antennae whitish-grey; ciliations in male 8. Thorax yellow reticulated with bright red.
Abdomen whitish-ochreous; bases of segments ochreous-brown. Legs red; anterior
femora fuscous; posterior pair whitish-ochreous. Forewings elongate, costa gently arched
to middle, thence straight, apex acute, termen sinuate, oblique; yellow coarsely reticulated
with bright red; an oblique fuscous streak from two-fifths costa to dorsum near base;
this is connected with base of costa by a streak along fold; an oblique fuscous streak
from three-fifths costa joining a fuscous tornal blotch, which extends to apex, and is

100 REVISION OF AUSTRALIAN LEPIDOPTERA. OECOPHORIDAE. XIII,

confluent with a large spot on mid-dorsum; cilia red, apices yellow, on apex and tornus
fuscous. Hindwings whitish; rosy-tinged at apex; cilia whitish.

Queensland: Macpherson Rge. (Springbrook) in October (W. B. Barnard); two
specimens. Type in Queensland Museum.

1923. LEPIDOTARSA CHRYSERYTHRA Turn., Trans. Roy. Soc. S. Aust., 1894, p. 134.
(Brisbane. )

1924. LEPIDOTARSA ATYPA, 1. Sp. (adrumos, unmarked.)

dg. 16 mm. Head white. Palpi with second joint reaching base of. antennae,
terminal joint one-half; pinkish-white finely sprinkled with fuscous, inner surface
whitish. Antennae ochreous-grey-whitish, towards base pink; ciliations in male 23.
Thorax white; tegulae pale ochreous. (Abdomen missing.) Legs white; anterior pair
crimson-pink. Forewings with costa moderately arched, apex pointed, termen straight,
oblique; pale ochreous; apex fuscous-tinged; cilia pale ochreous, on apex fuscous. Hind-
wings and cilia ochreous-whitish.

North Queensland: Cape York in June (W. B. Barnard); one specimen.

1925. LEPIDOTARSA RHODOPECHYS, N. Sp. (f0d0rnxXus, rosy-armed. )

9°. 16 mm. Head and thorax whitish; face faintly rosy. Palpi with terminal joint
one-half; whitish, apex of second joint faintly rosy. Antennae whitish with some
fuscous annulations. (Abdomen missing.) Legs whitish; anterior tibiae and tarsi
rosy with fuscous bars. Forewings narrow, costa gently arched, apex acute, termen
straight, strongly oblique; shining white; fuscous spots on base of dorsum, above fold
near its termination, and on tornus; cilia rosy, on tornus fuscous. Hindwings and cilia
pale grey. :

Queensland: Injune in August (W. B. Barnard); one specimen.

1926.7 LEPIDOTARSA ARGYROPiIS Meyr., Trans. Roy. Soc. S. Aust., 1902, p. 145. (Sydney. )

1927. LEPIDOTARSA IDIOcCOSMA Turn., ibid., 1898, p. 210. (Mt. Tamborine.)

1928. LrEPIDOTARSA PENTASCIA Turn., ibid., 1917, p. 106. (Tweed Hds., Macpherson
Rege.)

1929.; LEPIDOTARSA TRITOXANTHA Meyr., Proc. Linn. Soc. N.S.W., 1885, p. 828.
(Deloraine. )

1930. LrEpIpOTARSA CHRySOPOCA Meyr., ibid., 1882, p. 447. (Stanthorpe to Tasmania,
Ebor, Mittagong. )

1931. LeEPIDOTARSA ALPHITELLA Meyr., ibid., 1882, p. 447. = leucella Turn., Trans.
Roy. Soc. S. Aust., 1894, p. 135. (Brisbane, Newcastle, Sydney.)

1932. LEPIDOTARSA PROTEIS Meyr., Proc. Linn. Soc. N.S.W., 1887, p. 981. (Albany.)

1933.; LEPIDOTARSA LEUCODETIS Meyr., ibid., 1887, p. 931. (Geraldton.)

153. Gen. ATELOSTICHA Meyr.

Proc. Linn. Soc. N.S.W., 1887, p. 490.

Palpi with second joint reaching base of antennae, shortly rough-scaled; terminal
joint as long as second. Antennae without basal pecten. Forewings with 7 and 8
coincident. Type, A. phaedrella Meyr.

1934. ATELOSTICHA PHAEDRELLA Meyr., Proc. Linn. Soc. N.S.W., 1887, p. 491. (Bris-
bane, Tweed Hds., Newcastle, Sydney.)

1935. ATELOSTICHA CHRYSIAS Low., Trans. Roy. Soc. S. Aust., 1901, p. 65.
(Duaringa.)

154. Gen. LopHopEPLA Turn.

Trans. Roy. Soc. 8S. Aust., 1896, p. 10.

Palpi with second joint reaching base of antennae, thickened and dilated towards
apex; terminal joint stout, moderately long, shorter than second. Thorax with a small
double posterior crest. Forewings with tufts of raised scales; 7 and § stalked, 7 to termen.
Type, Lophopepla igniferella W1k.

1936.7 LoPHOPEPLA ARGYROCARPA Meyr., Haxot. Micro., i, p. 220. (Queensland.)

1937. LOPHOPEPLA TRISELENA Low., Trans. Roy. Soc. S. Aust., 1902, p. 240. (Cook-.

town, Townsville.) .

BY A. JEFFERIS TURNER. 101

1938. LopHOPEPLA ASTEROPA Low., Proc. Linn. Soc. N.S.W., 1900, p. 410. (Towns-
ville, Tweed Hds.)

1939. LoPHOPEPLA IGNIFERELLA WIK., xxix, p. 670. Turn., Trans. Roy. Soc. Aust., 1896,
p. 10. (Darwin, Thursday I. to Lismore, Macpherson Rge.)

155. Gen. AgraopEs Turn.

Trans. Roy. Soc. S. Aust., 1898, p. 205.

Palpi smooth, slender, ascending, recurved; second joint reaching base of antennae;
terminal joint one-third. Antennae without pecten. Forewings with 2 curved from
four-fifths, 7 and 8 stalked, 7 to apex. Hindwings with 5 from middle. Monotypical.

1940. AGLAODES CHIONOMA Turn., Trans. Roy. Soc. S. Aust., 1898, p. 205. (Brisbane,
Macpherson Rge.)

156. Gen. CALLITHAUMA Turn.

Trans. Roy. Soc. S. Aust., 1900, p. 79.

Palpi with second joint exceeding base of antennae; terminal joint one-half to three-
fifths. Antennae without basal pecten; in male simple. Forewings with 7 and 8 stalked,
7 to apex. Type, C. basilica Turn. Six species.

1941. CALLITHAUMA LEPTODOMA Turn., Proc. Linn. Soc. N.S.W., 1916, p. 366. (Tweed
Hds., Mt. Tamborine, Macpherson Rge., Killarney.)

1942. CALLITHAUMA GLYCERA Turn., ibid., 1916, p. 365. (Mt. Tamborine, Macpherson
Rge., Killarney.)

1943. CALLITHAUMA PYRITES Turn., Trans. Roy. Soc. S. Aust., 1896, p. 9. (Atherton,
Nambour, Brisbane, Stradbroke I., Mt. Tamborine, Killarney.)

1944. CALLITHAUMA BASILICA Turn., ibid., 1900, p. 15. (Mt. Tamborine, Macpherson
Rge., Toowoomba, Killarney.)

1945. CALLITHAUMA CALLIANTHES Meyr., Proc. Linn. Soc. N.S.W., 1888, p. 1595.
(Macpherson Rge., Stanthorpe, Robertson, Sydney, Mt. Wilson, Gisborne, Fernshaw,
Melbourne. )

1946. CALLITHAUMA MINIATULA, n. sp. ,(miniatulus, tinged with red.)

‘gS, 9. 14-15 mm. Head whitish-ochreous. Palpi with second joint reaching base
of antennae, terminal joint one-third; pale reddish. Antennae whitish. with fuscous
annulations; in male simple. Thorax. whitish-ochreous tinged with red. Abdomen
grey. Legs whitish; anterior pair reddish. Forewings narrow, costa slightly arched,
apex pointed, termen very oblique; grey: markings whitish-ochreous edged with red;
a dot on base of costa; another near base of dorsum; an oblique elongate spot from
one-fifth costa to fold, almost touching a similar spot from above fold to two-thirds
dorsum; a discal spot preceding an irregular curved interrupted line from midcosta
to tornus; a crenulate terminal line; cilia whitish-ochreous, on tornus grey. MHind-
wings whitish with some grey suffusion at apex; cilia grey.

Queensland: Toowoomba in September (W. B. Barnard); four specimens. Type in
Queensland Museum.

157. Gen. Tisoparica WIE.

WIK., xxix, p. 812. Meyr., Proc. Linn. Soc. N.S.W., 18838, p. 363.

Palpi with second joint very long, much exceeding base of antennae; terminal joint
one-fourth to one-half. Antennae without pecten; in male moderately ciliated. Fore-
wings with 7 and 8 stalked, 7 to apex. Type, T. jucundella Wlk. Ten species.

1947.; TISOBARICA ANCYROTA Meyr., Proc. Linn. Soc. N.S.W., 1883, p. 343. (Deloraine.)

1948. TIsoBARICA HEDANOPA Turn., ibid., 1916, p. 369. (Mt. Tamborine.)

1949. TisopaRICA HABROMORPHA Low., Trans. Roy. Soc. S. Aust., 1923, p. 54.
(Dorrigo.)

1950. TrsoBARICA LAROTYPA Turn., Proc. Linn. Soc. N.S.W., 1916, p. 368. (Toowoomba,
Sydney.)

1951. TIsoBARICA ERANNA Turn., ibid., 1916, p. 367. = jucundella Meyr., ibid., 1883,
p 364, nec Wlk. (Gympie, Brisbane, Macpherson Rge., Toowoomba, Newcastle, Sydney.)

102 REVISION OF AUSTRALIAN LEPIDOPTERA. OECOPHORIDAE. XIII,

1952. TISOBARICA JUCUNDELLA WIk., xxix, p. 813. Turn., Proc. Linn. Soc. N.S.W.,
1916, p. 367. (Brisbane, Sydney.)

1953. TISOBARICA PYRRHELLA Turn., Trans. Roy. Soc. S. Aust., 1896, p. 8. (Brisbane,
Toowoomba.)

1954. TISOBARICA HEMIGENES Meyr., Proc. Linn. Soc. N.S.W., 1888, p. 1596. (Albany,
Busselton. )

1955. TIsoBARICA THYTERIA Meyr., ibid., 1888, p. 1596. (Glen Innes, Ebor, Murru-
rundi, Mittagong, Gisborne, Melbourne.)

1956. TISOBARICA EXQUISITA, Nn. sp. (exquisitus, admirable.)

6, 9. 14-15 mm. Head crimson. Palpi with second joint reaching base of antennae,
terminal joint one-third; apex of second joint and terminal joint except base and apex
dark fuscous. Antennae whitish with blackish annulations; ciliations in male 1.
Thorax crimson; bases of tegulae and a posterior spot whitish. Abdomen grey. Legs
whitish with fuscous rings. Forewings narrow, costa slightly arched, apex pointed,
termen oblique; crimson; markings pale yellow slenderly edged with blackish; an
irregularly margined fascia from base of costa to one-fourth dorsum; a fascia from
one-fourth costa to mid-dorsum, interrupted above dorsum by a blackish bar; an inter-
rupted line from midcosta to tornus; an elongate spot from three-fourths costa not
reaching tornus; a blackish terminal line; cilia pale orange, on tornus fuscous. Hind-
wings and cilia whitish-ochreous.

New South Wales: Uki in October and November (W. B. Barnard); two specimens.
Type in Queensland Museum.

158. Gen. PyRICAUSTA, n.g. (vpixavstos, scorched.)

Palpi short; second joint not nearly reaching base of antenna, terminal joint one-
half. Antennae without basal pecten; ciliations in male long. Type, P. phaeopyra Turn.

1957. PyRICAUSTA PHAKOPYRA Turn., Proc. Roy. Soc. Tasm., 1926, p. 146. (Mt.
Wellington. )

159. Gen. EupHintra Meyr.

Proc. Linn. Soc. N.S.W.,.1882, p. 458.

Palpi with second joint just reaching base of antennae, thickened in middle and
loosely rough-scaled anteriorly; terminal joint as long as or shorter than second,
slender, acute. Antennae without pecten. Forewings with 7 and 8 stalked, 7 to costa.
Hindwings with 5 from middle of cell. Type, EZ. eroticella Meyr. Eleven species.

1958., EUPHILTRA CHRYSORRHODA Meyr., Trans. Roy. Soc. 8S. Auwst., 1902, p. 145.
(Sydney. )

1959. EupHiitrra Eprrecta Turn., ibid., 1917, p. 107. (Brisbane, Toowoomba. )

1960. EUPHILTRA CELETERTA Turn., ibid., 1917, p. 107. (Warwick, Killarney, Bunya
Mts., Lismore.)

1961. Eupninttra orrtHozona Low., ibid., 1920, p. 64. (Dalby, Injune. )

1962. EUPHILTRA HEMILEUCA Turn., ibid., 1890, p. 31. (Brisbane, Bunya Mts.)

1963. EUpHILTRA GEMMARIA Meyr., Exot. Micro., ii, p. 386. (Dalby.)

1964.; EUPHILTRA TRICENSA Meyr., ibid., ii, p. 219. (Murrurundi.)

1965. HuPHILTRA ANGUSTIOR’ Turn., Trans. Roy. Soc. 8. Aust., 1894, p. 187. (Brisbane
to Murrurundi, Macpherson Rge., Mittagong.)

1966. HUPHILTRA EROTICELLA Meyr., Proc. Linn. Soc. N.S.W., 1882, p. 458. = thermo-
zona Turn., Trans. Roy. Soc. S. Aust., 1894, p. 137. (Brisbane to Melbourne, Macpherson
Rge., Milmerran, Mt. Wilson, Kewell.)

1967. EUPHILTRA FUSIPLAGA Turn., ibid., 1917, p. 106. (Brisbane, Macpherson Rge.,
Toowoomba, Bunya Mts.)

1968. EUPHILTRA PHANEROZONA, N. Sp. (davepogwvos, plainly girdled.)

3, 2. 14-16 mm. Head white. Palpi with second joint two-thirds; white, basal half
of outer surface of second joint dark fuscous. Antennae whitish with fuscous annula-
tions; ciliations of male 1 and a half. Thorax white; tegulae and posterior edge blackish.
Abdomen fuscous; tuft grey-whitish. Legs dark fuscous; posterior pair whitish-ochreous.
Forewings with costa gently arched, apex rounded, termen obliquely rounded; white

BY A. JEFFERIS TURNER. 103

with fuscous markings; a very narrow basal fascia; a transverse fascia at one-third,
expanded on margins; a subterminal fascia from apex of costa to tornus, expanded on
eosta, where it contains a white spot; a terminal line thickened into a median spot;
cilia pale ochreous. Hindwings and cilia fuscous.

Queensland: Milmerran in March and April; three specimens.

160. Gen. ZoONOPETALA Meyr.

Proc. Linn. Soc. N.S.W., 1882, p. 459.

Palpi with second joint reaching base of antennae, thickened in middle and usually
loosely rough-haired anteriorly; terminal joint as long as or shorter than second, slender,
acute. Antennae without pecten. Forewings with 7 and 8 stalked, 7 to termen.
Hindwings with 5 approximated to 4 at origin. Type, Z. clerota Meyr. Fifteen species.

1969. ZONOPETALA DIVISELLA WIK., xxix, p. 677. Meyr., Proc. Linn. Soc. N.S.W., 1882,
p. 460. (Duaringa to Castlemaine, Milmerran, Bathurst, Mittagong, Birchip.)

1970. ZONOPETALA CLEROTA Meyr., ibid., 1882, p. 461. (Herberton, Nambour to
Melbourne, Glen Innes, Mittagong, West Australia.)

1971. ZONOPETALA CORRECTA Meyr., Haxot. Micro., i, p. 304. (Macpherson Rge.,
Warwick, Sydney.)

1972. ZONOPETALA SYNARTHRA Meyr., Proc. Linn. Soc. N.S.W., 1885, p. 929. (Mt.
Kosciusko, Gisborne, Quorn, Broken Hill.)

1973. ZONOPETALA GLAUCONEPHELA Meyr., ibid., 1882, p. 462. (Brisbane, Toowoomba,
Sydney, Trafalgar, Adelaide. )

1974. ZONOPETALA DECISANA WI1k., xxviii, p. 367. Meyr., Proc. Linn. Soc. N.S.W.,
1882, p. 4638. = mediella WI1k., xxx, p. 1033. = ustella WIk., xxix, p. 673. (Sydney,
Murrurundi, Gosford, Melbourne, Birchip, Adelaide.)

1975. ZONOPETALA DIDYMOSTICHA, nN. Sp. (dcdvuoorrxos, twin-streaked.)

6, .. 14-17 mm. Head white. Palpi with terminal joint two-thirds; white, base of
second joint fuscous. Antennae whitish with fuscous annulations; ciliations in male 1
and a half. Thorax dark fuscous; apices of tegulae and anterior and posterior spots
white. Abdomen whitish-ochreous. Legs fuscous; posterior pair whitish-ochreous.
Forewings with costa gently arched, apex rounded, termen obliquely rounded; white;
markings dark fuscous; a very narrow basal fascia; a median fascia, broader towards
dorsum, anterior edge incurved from one-third costa to two-fifths dorsum, posterior
nearly straight from two-fifths costa to two-thirds dorsum; a short slender vertical
streak connecting these two fasciae; two short closely parallel streaks running to termen
beneath apex, sometimes partly fused; cilia white, on apex fuscous. Hindwings whitish-
ochreous suffused with fuscous towards apex; cilia whitish-ochreous, on apex fuscous.
Very like Z. decisana; distinguished by the apical twin streaks on forewings.

Queensland: Nambour in November; Brisbane in October; Tweed Hds. in November.
New South Wales: Gosford in November. Ten specimens.

1976.4 ZONOPETALA ZYGOPHORA Low., Trans. Roy. Soc. 8S. Aust., 1894, p. 95. (Duaringa.)

1977. ZONOPETALA VISCATA Meyr., Hxot. Micro., i, p. 194. (Cairns to Lismore.)

1978. ZONOPETALA PROPRIA, N. Sp. (proprius, peculiar.)

3. 19-22 mm. Head pale grey; face white. Thorax pale grey. Palpi with second
joint exceeding base of antennae, terminal joint two-thirds; white, base of second joint
fuscous. Antennae whitish-grey, base fuscous; ciliations in male 2. Abdomen ochreous.
Legs pale reddish; posterior pair ochreous. Forewings with costa strongly arched, apex
subrectangular, termen straight, not oblique, rounded beneath; pale grey; markings
dark fuscous sometimes partly edged with reddish; a semi-circular spot on base of
costa; a spot on one-third costa connected by a short oblique streak with a spot in disc
at three-fifths; a line of minute fuscous dots from three-fifths costa, angled before
termen, thence curved to dorsum before tornus; a series of minute fuscous terminal
dots; cilia pale grey. Hindwings ochreous; cilia pale ochreous-grey.

Queensland: Emerald; Milmerran in November; Injune in December; three
specimens. ;

104 REVISION OF AUSTRALIAN LEPIDOPTERA. OECOPHORIDAE. XIII,

1979. ZONOPETALA PAROOSPILA, Nl. Sp. (mapwoarXos, brown-spotted. )

dg. 18-20 mm. Head, thorax and abdomen pale grey. Palpi with terminal joint
two-thirds; pale grey, outer surface of second joint except apex fuscous. Antennae pale
grey; ciliations in male 4. Legs fuscous; posterior pair whitish. Forewings triangular,
costa strongly arched, apex rectangular, termen sinuate, not oblique; pale grey sparsely
and unequally dotted with fuscous-brown; a rather large suffused fuscous-brown
subcostal spot at four-fifths; four dark fuscous costal spots beyond middle; a terminal
series of dark fuscous dots; cilia pale grey, bases fuscous except on apex and tornus.
Hindwings with termen sinuate; pale grey; cilia pale grey. :

Queensland: Noosa in October; Macpherson Rge. (3,000 ft.) in March. New South
Wales: Lismore in October. Three specimens.

1980.. ZONOPETALA TEPHRASTIS Turn., Trans. Roy. Soc. 8S. Aust., 1917, p. 107.
(Innisfail. )

1981. ZONOPETALA ERYTHROSEMA Meyr., Proc. Linn. Soc. N.S.W., 1885, p. 829.
(Victoria, Tasmania. )

1982. ZONOPETALA QUADRIPUSTULELLA WIK., xxix, p. 761. Meyr., Proc. Linn. -Soc.
N.S.W., 1882, p. 466. (Atherton, Brisbane to Tasmania.)

1983. ZONOPETALA MELANOMA Meyr., ibid., 1882, p. 465. (Macpherson Rge., Sydney.)

161. Gen. ParocystoLa Turn.

Trans. Roy. Soc. S. Aust., 1896, p. 30.

Palpi ascending, recurved; second joint reaching base of antennae, smooth, slender;
terminal joint shorter than second, slender, acute. Antennae without pecten or rarely
with a few scales only. Forewings with termen sinuate; 7 and 8 stalked, 7 to termen.
Hindwings with 5 approximated to 4 at origin. Monotypical. It shows collateral
relationship to Zonopetala viscata.

1984. PAROCYSTOLA LEUCOSPORA Turn., Trans. Roy. Soc. S. Aust., 1896, p. 30.
(Brisbane, Toowoomba. )

162. Gen. ArcHAERETA Meyr.

Exot. Micro., i, p. 223.

Palpi long, ascending, recurved; second joint exceeding base of antennae, somewhat
thickened with smoothly appressed scales, with a median anterior furrow; terminal
joint shorter than second, slender, acute. Forewings with 7 and 8 stalked, 7 to
termen; termen sinuate. Hindwings with 5 approximated to 4 at origin. Monotypical.

1985. ARCHAERETA DORSIVITTELLA WIk., xxviii, p. 538- Meyr., Trans. Roy. Soc. 8S.
Aust., 1902, p. 146. (Toowoomba, Barrington Tops, Mt. Macedon, Hobart, Mt. Lofty.)

163. Gen. COLPOLOMA, n.g. (koArwohwuos, With sinuate margins.)

Palpi ascending, recurved; second joint reaching base of antennae, much thickened
especially towards apex, rough anteriorly; terminal joint much shorter than second,
stout, obtuse. Antennae with strong pecten. Forewings with costa and termen sinuate;
7 and 8 stalked, 7 to termen. Hindwings with 5 approximated to 4 at origin.

1986. COLPOLOMA FRAXINEA, n. sp. (fraxineus, ashen.)

°. 32 mm. Head, thorax, palpi and antennae pale grey. Abdomen grey. Legs
fuscous with whitish rings; posterior pair grey-whitish. Forewings elongate, costa
strongly arched at base, apex pointed, slightly produced; termen not oblique; pale grey;
a small tuft of fuscous scales on one-fifth dorsum; costal edge from one-third to apex
whitish interrupted by minute fuscous dots; first discal at one-third, plical beyond it,
both fuscous, second discal at three-fifths, snow-white; traces of a curved subterminal
line; cilia pale grey mixed with fuscous. Hindwings and cilia grey.

New South Wales: Katoomba (Undercliff Falls) in August (W. B. Barnard); one

specimen.

164. Gen. MAcropHasRA, n.g. (paxpopapos, long-cloaked.)

Palpi short; second joint slender, not reaching beyond middle of face; terminal
joint as long as second, slender, acute, scarcely reaching vertex. Antennae with pecten.

——

BY A, JEFVYERIS TURNER. 105

Forewings with 2 and 3 stalked, 7, 8, 9 stalked, 7 to apex, 10 arising nearer to 11 than
to end of cell. Hindwings with 5 curved and approximated to 4 at origin.

1987. MaAcropHARA ANEURETA, 1. Sp. (dvevperos, hard to find.)

do. 24-28 mm. Head and palpi grey. Antennae fuscous; ciliations in male 2.
Thorax dark fuscous; tegulae grey-whitish. Abdomen fuscous; tuft grey-whitish. Legs
fuscous; posterior tibiae and tarsi grey-whitish. Forewings narrow, costa nearly
straight; apex rounded, termen obliquely rounded; grey-whitish lightly sprinkled with
fuscous; a cloudy fuscous basal fascia, produced on dorsum to middle; a fuscous dot
on tornus; stigmata dark fuscous, minute, first discal at one-third, plical beneath it,
second discal before two-thirds; cilia grey-whitish. Hindwings and cilia grey-whitish.

Queensland: Mt. Tamborine in October; Bunya Mts. in November; two specimens.

165. Gen. EPICHARACTIS, n.g. (éem-xapaxris, furrowed.)

I propose this name for Paracharactis Meyr. (H«wxot. Micro., ii, 1918, p. 215) which
is preoccupied by Meyrick and Lower (Trans. Roy. Soc. S. Awst., 1907, p. 205).

Palpi recurved, ascending; second joint reaching base of antennae; terminal joint
shorter than second, slender, acute. Antennae without pecten. Forewings with 7, 8,
9 stalked, 7 to apex. Hindwings with 5 curved and approximated to 4 at origin.

1988. HPICHARACTIS MITOSEMA Turn., Proc. Linn. Soc. N.S.W., 1916, p. 373.
(Brisbane, Toowoomba. )

166. Gen. LeristTarcHA Meyr.

Proc. Linn. Soc. N.S.W., 1883, p. 422. = Tigava WI1k., preoccupied.

Palpi long, ascending, recurved; second joint exceeding base of antennae, moderately
thickened, smooth; terminal joint shorter than second, slender, acute. Antennae without
pecten. Posterior tarsi in male with basal joint much elongated. Forewings with 7 and
8 stalked, 7 to termen. Hindwings with 5 curved and approximated to 4 at origin.
Monotypical.

1989. LEISTARCHA SCITISSIMELLA WIk., xxix, p. 807. = iobola Meyr., Proc. Linn. Soc.
N.S.W., 1883, p. 326. (Cape York, Brisbane, Toowoomba, Stanthorpe, Murrurundi, Picton,
Gisborne, Beaconsfield. )

167. Gen. PycNnozANcLA Turn.

Trans. Roy. Soc. 8S. Aust., 1917, p. 109.

Palpi ascending, recurved, second joint reaching base of antennae, smooth; terminal
joint stout as compared with second and shorter acute. Antennae without pecten.
Forewings with 7 and 8 stalked, 7 to termen. Hindwings with 5 curved and approxi-
mated to 4 at origin. Type, P. acribes.

1990. PycNOZANCLA ERYTHRODES: Turn., Trans. Roy. Soc. S. Auwst., 1917, p. 109.
(Nambour, Mt. Tamborine, Bunya Mts., Allyn R.).

1991. PycNOZANCLA EPIPREPES Turn., ibid., 1894, p. 136. = plinthomicta Meyr.,
Exot. Micro., i, p. 193. (Cairns, Atherton, Townsville, Gympie.)

1992. PycnozANCLA ACRIBES Turn., Trans. Roy. Soc. S. Aust., 1894, p. 135. (Gympie,
Nambour, Brisbane, Toowoomba, Stanthorpe. )

168. Gen. CompsorropHa Meyr.

Proc. Linn. Soc.. N.S.W., 1883, p. 511.

Palpi ascending, recurved; second joint reaching base of antennae, slender, smooth;
terminal joint as long as second, slender, acute. Antennae without pecten. Forewings
with 7 and 8 stalked, 7 to termen. Hindwings with 5 curved and approximated to 4 at
origin. Type, C. selenias.

1993. ComPpsopoTiIS CHARIDOTIS Meyr., Proc. Linn. Soc. N.S.W., 1883, p. 513. (Quilpie,
Melbourne, Wirrabara. )

1994. ComMPSOTROPHA STROPHIELLA Meyr., ibid., 1883, p. 513. (Atherton, Brisbane to
Tasmania, Glen Innes, West Australia.)

1995. CoMPSOTROPHA SELENIAS Meyr., ibid., 1883, p. 512. (Stanthorpe to Hobart.)

106 REVISION OF AUSTRALIAN LEPIDOPTERA. OECOPHORIDAE. XIII,

169. Gen. PryopTiLa, n.g. (atvomridos, fan-winged.)

Palpi ascending, recurved; second joint reaching base of antennae, moderately
slender, smooth; terminal joint shorter than second, slender, acute. Antennae without
pecten. Forewings with fan-shaped tuft of scales on one-fourth dorsum; 7 and 8 stalked,
7 to termen. Hindwings with 5 curved and approximated to 4 at origin.

1996. PryorpriILA MATUTINELLA WIk., xxix, p. 698. Meyr., Proc. Linn. Soc. N.S.W.,
1882, p. 453. = marginella Wlk., xxix, p. 761. (Gayndah to Wangaratta, Stanthorpe,
Ebor.)

170. Gen. MacHimi1a Clemens.

Proc. Acad. Nat. Sci. Philad., 1860, p. 211.

Palpi ascending, recurved; second joint reaching base of antennae, moderately
slender, smooth, or rarely with some rough scales anteriorly; terminal joint shorter
than second, slender, acute. Antennae without pecten, or with a few scales only.
Forewings with 7 and 8 stalked, 7 to apex or termen. Hindwings with 5 curved and
approximated to 4 at origin. Type, M. tentiforella Clem. from North America. Nothing
can be gained by dividing this genus according to the termination of 7 of the forewings,
which varies sometimes in the same species. The antennal ciliations, which vary from
minute to very long, are often helpful in distinguishing species. I have therefore given
them, where known, immediately after the name of each species.

Of this large genus I have listed 141 Australian species, and more will be discovered.
Meyrick records 84 from South America, ten from North America, and one from Japan.
He infers that the genus originated in South America and spread to Australia across the
Antarctic, but it seems to me equally probable that it travelled in the opposite direction.

1997. MACHIMIA THAUMASTICA, N. Sp. (@avuaorixos, surprising. )

°. 38 mm. Head and thorax grey. Palpi with second joint exceeding base of
antennae, terminal joint two-thirds; pale grey. Antennae whitish annulated with
fuscous. (Abdomen missing.) Legs grey-whitish. Forewings elongate, costa gently
arched, apex pointed, termen very obliquely rounded; 7 and 8 long-stalked, 7 to apex;
grey-whitish lightly sprinkled with fuscous; a slender median fuscous line from
two-fifths nearly to termen with a short parallel line on its costal side; three short
streaks running to costa, one to apex, several to termen, and one to tornus; cilia grey;
apices whitish. Hindwings with 5 nearly approximated to 4 at origin; ochreous-
yellow; a pale fuscous suffusion around apex, termen and tornus; cilia pale ochreous,
bases pale fuscous. Though superficially very distinct, this is a true Machimia suggesting
some relationship to Epicharactis.

New South Wales: Mittagong in November (G. M. Goldfinch); one specimen. Type
in Australian Museum.

1998. MACHIMIA CONSPERSA, Nn. sp. (conspersus, sprinkled.)

d. 24 mm. Head and thorax orange. (Palpi broken.) Antennae orange;
ciliations in male two-thirds. Abdomen pale ochreous. Forewings with costa strongly
arched, apex rounded-rectangular, termen straight, slightly oblique; orange; markings
purple-fuscous, suffused; a sub-basal costal spot; two broad lines from one-third costa,
one transverse to one-third dorsum, the other oblique to middle of disc, there bent and
ending on mid-dorsum; a discal spot at two-thirds connected with a dot on two-thirds
costa, and with a small subapical costal blotch, from which proceeds a slender line to
tornus; a dotted terminal line; cilia pale ochreous. Hindwings and cilia pale ochreous.

Victoria: Macedon in October; one specimen. Type in National Museum.

1999. MAcHIMIA HYPERTRICHA Turn., Proc. Roy. Soc. Tasm., 1926, p. 147. (Mt.
Wellington.)

2000. MACHIMIA CALLIANASSA Meyr., Proc. Linn. Soc. N.S.W., 1882, p..450. (Victoria,
Tasmania, Mt. Lofty.)

2001. MAcHIMIA SARCOXANTHA Low., Trans. Roy. Soc. S. Aust., 1893, p. 176. = zelota

Turn., Proc. Linn. Soc. N.S.W., 1916, p. 370. (Stanthorpe, Glen Innes, Sydney, Victoria.)

BY A. JEFFERIS TURNER. 107

2002. MACHIMIA LAETIFERANA WIK., xxviii, p. 336. Meyr., Proc. Linn. Soc. N.S.W.,
1882, p. 449. = semifusella W1k., xxix, p. 696. = pudorinella W1k., xxix, p. 760. (Yeppoon
to Melbourne, Stanthorpe. )

2003. MACHIMIA MELANOPLECTA Turn., Trans. Roy. Soc. S. Aust., 1917, p. 110.
(Gympie, Nambour, Bunya Mts.)

2004. MAcHIMIA DIPHRACTA Low., ibid., 1920, p. 63. (Cairns.)

2005. MACHIMIA LEIOCHROA Low., ibid., 1907, p. 114. = sporeta Turn., ibid., 1917,
p. 75 (Cape York, Townsville. )

2006. MACHIMIA PHOENOPIS Turn. (two-thirds). Proc. Linn. Soc. N.S.W., 1916, p. 371
(Darwin, Claudie R. to Sydney, Stanthorpe, Milmerran.) :

2007. MacHIMIA ABSUMPTELLA WIk. (one-half). xxix, p. 567. Meyr., Proc. Linn. Soc.
N.S.W., 1882, p. 506. (Brisbane to Launceston, Macpherson Rge., Ebor, Mt. Lofty, Perth.)

2008.7 Macuimia ALMA Meyr. (1 and two-thirds). Hxot. Micro., ii, p. 375. (Gisborne. )

2009. MACHIMIA MICROMITA, nN. Sp. (ptxpourros, With small threads.)

20 mm. Head and thorax grey faintly reddish-tinged. Palpi with second joint ©
exceeding base of antennde, terminal joint almost 1; second joint whitish with terminal
and subterminal fuscous rings, terminal joint grey. Antennae whitish with narrow
fuscous rings; ciliations in male one-third. Abdomen grey; tuft whitish-ochreous. Legs
whitish with fuscous tarsal rings. Forewings rather strongly arched, apex rounded-
rectangular, termen obliquely rounded; whitish-ochreous with grey irroration forming
obscure lines and streaks on and between veins; stigmata minute, first discal at
one-third, plical beneath it, second discal at two-thirds; cilia whitish. Hindwings and
cilia whitish. Very like M. mitescens, but differing in its short antennal ciliations and
long terminal joint of palpi. .

North Queensland: Cape York in October (W. B. Barnard); one specimen.

2010. MacHIMIA MITESCENS Meyr. (2 and a half). Haot. Micro., i, p. 174. (Townsville,
Injune. )

2011. MAcHIMIA MILTOPSARA Turn. (one-half). Proc. Linn. Soc. N.S.W., 1914, p. 560.
= biseriata Meyr., Exot. Micro., ii, p. 375. (Brisbane, Mt. Tamborine, Ebor, Allyn R.,
Lorne, Crade Mt., Mt. Wellington.)

2012. Macuimia sErvA Meyr. (1). Hxot. Micro., ii, p. 375. (Murrurundi, Canberra,
Birchip. )

2013.4 MACHIMIA PRAEPEDITA Meyr., ibid., ii, p. 375.. (Hoyleton, Pinnaroo.)

2014. MACHIMIA DELOSTICTA, n. Sp. (dndAooTLKTOs, With distinct dots.)

6. 24-25 mm. Head and thorax pale ochreous-grey. Palpi with second joint
reaching base of antennae, terminal joint three-fifths; ochreous-whitish. Antennae pale
grey; ciliations in male 4. Abdomen grey; apices of segments and tuft ochreous-
whitish. Legs grey; tarsi fuscous with whitish rings. Forewings with costa moderately
arched, apex obtuse, termen obliquely rounded; pale ochreous-grey with sharply
distinct dark fuscous dots and scanty irroration; first discal at one-fourth, plical slightly
beyond, second discal at three-fifths; a dot beneath second; a few dots towards base;
a very slender fuscous line from mid-costa to four-fifths, bent beneath costa and ending
on tornus, slightly rippled or divided into dots; a terminal series of dots continued
on apical fourth of costa; cilia concolorous. Hindwings ochreous-whitish, towards apex
slightly greyish; cilia ochreous-whitish.

Victoria: Moe in February; four specimens.

2015. MACHIMIA CHOLODELLA Meyr. (1 and a half). Proc. Linn. Soc. N.S.W., 1882,
p. 507. (Katoomba).

2016. MAcHiIMIA SERICATA Meyr. (2). Ibid., 1882, p. 497. (Brisbane, Mt. Tamborine,
Sydney).

2017. MAcHIMIA COMPLANULA Turn. (2). Trans. Roy. Soc. S. Aust., 1896, p. 4.
(Brisbane, Tweed Hds., Stanthorpe, Murrurundi.)

2018. MACHIMIA STENORRHODA, Nn. Sp. (orevoggod0s, narrowly rosy.)

6, 9. 22 mm. Head and thorax grey. Palpi with second joint just reaching base
of antennae, terminal joint two-thirds; grey. Antennae whitish with fuscous annula-
tions; ciliations in male 1 and a half. Abdomen pale grey; apices of segments and tuft

108 REVISION OF AUSTRALIAN LEPIDOPTERA. OECOPHORIDAE. XIII,

whitish. Legs whitish; anterior pair grey. Forewings suboblong, costa gently arched,
apex rounded-rectangular, termen slightly rounded, slightly oblique; pale grey; costa
narrowly rosy with extreme edge whitish; stigmata fuscous, first discal at one-third,
minute, plical beyond it, minute, second discal before two-thirds, larger, slightly elongate
and outwardly oblique; subterminal line absent or very faintly indicated; cilia grey-
whitish. Hindwings ochreous-whitish with faint grey suffusion on apex and termen;
cilia whitish.

Queensland: Brisbane and Tweed Hds. in September; four specimens.

2019. MACHIMIA SUBMISSA, nN. sp. (submissus, humble.)

3, 2. 16-18 mm. Head and thorax dull rosy. Palpi with second joint reaching
base of antennae, terminal joint one-half; rosy. Antennae grey; ciliations in male 1 and
a half. Abdomen grey; sides, apices of segments, and tuft whitish. Legs rosy; posterior
tibiae whitish. Forewings narrow, costa gently arched, apex subrectangular, termen
obliquely rounded; dull rosy with a rosy costal line; stigmata minute, obscure, pale grey,
' first discal at two-fifths, plical beyond it, second discal at three-fifths; cilia grey-
whitish. Hindwings ochreous-whitish; cilia whitish.

North Queensland: Cape York in October (W. B. Barnard); two specimens. Type
in Queensland Museum.

2020. MACHIMIA RUFESCENS, nh. sp. (rufescens, reddish.)

d. 20 mm. Head and thorax reddish. Palpi with second joint reaching base of
antennae, terminal joint one-half; whitish, terminal joint fuscous anteriorly. Antennae
whitish with dark fuscous annulations; ciliations in male one-fourth. Abdomen
ferruginous-fuscous. Legs grey; anterior pair reddish-tinged; tarsi fuscous with whitish
rings. Forewings suboblong, costa moderately arched, apex rounded, termen obliquely
rounded; reddish; markings fuscous, obscure; stigmata minute, first discal at one-third,
plical beyond it, second discal at three-fifths; a basal dorsal spot, and another median
near base; obscure slender irregular transverse lines at one-third and two-thirds; a
slender slightly dentate line from midcosta, subcostal to near termen, thence bent to three-
fourths dorsum; a terminal series of minute dots; cilia reddish, apices whitish.
Hindwings grey; cilia whitish. The male should be easily recognized by its minute
antennal ciliations.

Tasmania: Launceston (Gravelly Beach) in February (W. B. Barnard); one
specimen. ;

2021. MACHIMIA HABROPTERA Low., Proc. Linn. Soc. N.S.W., 1900, p. 411. = coccinea
Turn., Trans. Roy. Soc. S. Aust., 1917, p. 59. (Cairns, Brisbane, Carnarvon Rge.)

2022. MACHIMIA RUFA Meyr. (1). Proc. Linn. Soc. N.S.W., 1882, p. 504. (Stradbroke
I., to Melbourne, Stanthorpe, Mt. Lofty.)

2023. MACHIMIA LOXOMITA, n. sp. (XAogoucTos, With an oblique thread.)

3, 2. 20-22 mm. Head and thorax pale reddish. Palpi with second joint reaching
base of antennae, terminal joint two-thirds; whitish. Antennae whitish annulated with
dark fuscous; ciliations in male 1. Abdomen grey; apices of segments and tuit
ochreous-whitish. Legs ochreous-whitish; anterior pair pale rosy. Forewings with costa
gently arched, apex rounded-rectangular, termen obliquely rounded; whitish-ochreous;
costa more or less rosy; markings pale rosy or pale fuscous; stigmata minute, first
discal at one-third, plical slightly beyond it, second discal before two-thirds; an
oblique line from three-fourths costa to mid-dorsum, sometimes only faintly developed,
running through second discal; a very fine outwardly oblique line from -three-fourths
costa, bent beneath costa, ending on dorsum before tornus; an interrupted or suffused
terminal line; cilia ochreous-whitish. Hindwings and cilia ochreous-whitish.

North Queensland: Atherton in November. Queensland: Caloundra in February;
Mt. Tamborine in September; Toowoomba in November and December. Six specimens.

2024. MACHIMIA UMBRATICA, n. sp. (umbraticus, shady.)

3, 2. 20-22 mm. Head and thorax dusky reddish. Palpi with second joint reaching
base of antennae, terminal joint three-fifths; rosy, towards base whitish. Antennae
whitish annulated with fuscous; ciliations in male 1. Abdomen greyish-brown; apices
of segments and tuft whitish-ochreous. Legs whitish-ochreous; anterior pair rosy.

ee Nine

BY A, JEFFERIS TURNER. 109

Forewings with costa gently arched, apex rectangular, termen obliquely rounded;
dusky reddish, on costa rosy; stigmata usually obsolete; cilia grey-whitish. Hindwings
and cilia grey.

Queensland: Injune in April; Milmerran in March; five specimens.

2025. MACHIMIA ONCOSPILA, Nn. Sp. (dyKoomA0s, With swollen spot.)

9. 20-25 mm. Head ochreous-grey. Palpi wih second joint exceeding base of
antennae, terminal joint two-thirds; rosy. Thorax rosy wih central fuscous spot.
Abdomen grey; apices of segments and tuft whitish-ochreous. Legs whitish; anterior
pair rosy. Forewings elongate, costa gently arched, apex rectangular; termen slightly
rounded, scarcely oblique; pale rosy, brighter on costa; extreme costal edge whitish;
markings fuscous; a median sub-basal dot; first discal at one-third, plical well beyond it,
second discal at two-thirds, swollen, circular; short incomplete subterminal and terminal
lines; cilia ochreous-whitish. Hindwings grey; cilia whitish with a grey sub-basal
line.

West Australia: Kalamunda, near Perth, in December (W. B. Barnard); two
specimens. Type in Queensland Museum.

2026. MACHIMIA SINCERELLA WIK., xxxv, p. 1835. = hepatitis Meyr., Trans. Roy. Soc.
S. Aust., 1902, p. 147. (Gisborne.)

2027. MACHIMIA RHODOXANTHA Meyr. (1 and a half). Proc. Linn. Soc. N.S.W., 1887,
p. 933. (Brisbane to Melbourne.)

2028. MACHIMIA COSTIMACULA Meyr. (2). Ibid., 1882, p. 502. (Cape York to
Sydney. )

2029. MACHIMIA HABROSCHEMA, nN. Sp. (aB8pooxnuos, softly patterned.)

6. 24 mm. Head and thorax dull rosy. (Palpi and antennae missing.) Abdomen
brownish. Forewings elongate, costa slightly arched, apex rounded-rectangular, termen
slightly rounded, scarcely oblique; rosy-grey; a bright rosy costal line, extreme costal
edge ochreous; a triangular rosy-edged spot on three-fifths costa, connected by rosy and
ochreous suffusion with two-thirds dorsum; stigmata small, purple-fuscous; first discal
at one-third, plical before it, second discal before two-thirds; a suffused grey terminal
line; cilia rosy, apices pale ochreous. Hindwings and cilia whitish.

South Australia: Mt. Lofty (Blackwood). Type in South Australian Museum.

2030. Macwuimra pupica Zel., Lin. Hnt., x, p. 152.. Meyr., Proc. Linn. Soc. N.S.W.,
1882, p. 500. (Yeppoon to Hobart, Millmerran, Macpherson Rge., Ebor, Katoomba,
Mt. Lofty, West Australia.) .

2031. MAcHIMIA SPATIOSA Meyr., Exot. Micro., ii, p. 387. (Mt. Wilson, Melbourne,
Beaconsfield. )

2032. MACHIMIA OCHRA, nN. sp. (xpos, pale.)

3, 2. 28-32 mm. Head and thorax pale greyish-ochreous. Palpi with second joint
exceeding base of antennae, terminal joint three-fifths; pale rosy. Antennae grey-
whitish; ciliations in male 4. Abdomen grey; apices of segments and tuft whitish. Legs
rosy; posterior pair white. Forewings not dilated, costa slightly arched, apex rect-
angular, termen obliquely rounded; pale greyish-ochreous faintly tinged reddish; a
rosy costal line leaving extreme edge whitish; markings pale fuscous, often faint or
absent; a dentate line from one-fourth costa to mid-dorsum; a line of dots from costa
beyond middle to four-fifths, there bent to end on dorsum before tornus; a terminal
series of dots; cilia concolorous. Hindwings and cilia whitish.

West Australia: Waroona in November; Perth in September; seven specimens.

2033. MACHIMIA LEUCERYTHRA Meyr. (1). Proc. Linn. Soc. N.S.W., 1882, p. 501.
= rufimaculella Turn., Trans. Roy. Soc. S. Aust., 1896, p. 7. = acosmeta Turn., ibid., 1896,
= rhodopepla Turn., ibid., 1903, p. 221. (Cape York to Tasmania, Mt. Kosciusko, Mt.
Lofty, West Australia.)

2034. MACHIMIA HAPLOCEROS, nN. Sp. (amAoxepws, with simple antennae.)

3, 2. 22-26 mm. Head and thorax dull rosy. Palpi with second joint three times
length of face, terminal joint one-fifth; dull rosy. Antenna grey; ciliations in male
extremely minute. Abdomen whitish. Legs whitish; anterior pair rosy. Forewings
with costa strongly arched to middle, thence straight, apex rectangular, termen slightly

N

110 REVISION OF AUSTRALIAN LEPIDOPTERA. OECOPHORIDAE. XIII,

rounded, sligbtly oblique; dull rosy, brighter on costa; markings fuscous; first discal
at one-third, plical slightly beyond it, second discal at three-fifths; a line of minute
dots from midcosta, subcostal to near apex, there angled, and curved to before tornus;
a marginal series of dots on termen and apical fourth of costa; ciliations pale rosy.
Hindwings and cilia whitish.

Queensland: Macpherson Rge. (3,000 ft.) in October and November (W. B.
Barnard); two specimens. Type in Queensland Museum.

2035. MAcHIMIA AMATA Meyr. (1). Hot. Micro., i, p. 175. (Waroona.)

2036. MACHIMIA LiIMBATA Meyr., Proc. Linn. Soc. N.S.W., 1882, p. 471.

This and the next two species are so similar that, if it had’ not been for the
differences in their antennal ciliations, I should not have distinguished them.

6, 2. 20-28 mm. Antennal ciliations in male 1. Forewings of male with oblique
line from two-fifths costa to one-fifth dorsum, sometimes suffused on dorsum, occasionally
absent; a second oblique line from two-thirds costa to mid-dorsum, more or less broadly
suffused; in female these markings are rarely developed. Hindwings yellow, apex and
sometimes termen suffused with fuscous; cilia fuscous. (Toowoomba, Dalby, Stanthorpe,
Glen Innes, Ebor, Melbourne, Beaconsfield.) Seventeen specimens.

2037. MACHIMIA SIMILIS, n. sp. (similis, similar.)

3, @. 22-25 mm. Antennal ciliations of male 2. Forewings without oblique lines.
Hindwings pale yellow without any grey suffusion; cilia grey.

New South Wales: Glen Innes; Katoomba. Victoria: Castlemaine.

2038. MACHIMIA BREVICILIA, nh. sp. (brevicilius, with short ciliations. )

do. 22-24 mm. Antennal ciliations of male extremely minute. Forewings with post-
median suffused oblique line. Hindwings pale yellow with slight grey suffusion at
apex; cilia grey.

New South Wales: Ben Lomond, Mt. Kosciusko; three specimens.

20389. MAcHIMIA EOXANTHA Turn., Trans. Roy. Soc. 8. Aust., 1896, p. 7. (Cape York
to Sydney, Stanthorpe, Katoomba. )

2040.7 MAcHIMIA ATOECHA Meyr., Proc. Linn. Soc. N.S.W., 1885, p. 830. (Sydney.)

2041.4 MACHIMIA METRIOPIS Meyr., ibid., 1887, p. 941. (Bathurst.)

2042. MACHIMIA REPANDULA Zel. (1). Lin. Hnt., x, Pl. 150, f. 3. Meyr., Proc. Linn.
Soc. N.S.W., 1882, p. 499. (Brisbane to Tasmania, Mt. Lofty.)

2043. MACHIMIA CARNEA Zel. (1). Lin. Ent., x, Pl. 148. = pseudota Low., Trans.
Roy. Soc. S. Aust., 1901, p. 85. = insana Meyr., Hxot. Micro., ii, p. 387. (Atherton,
Brisbane to Tasmania. )

2044. MACHIMIA MODERATELLA WIk. (one-half), xxix, p. 566. = liosarca Meyr., Proc.
Linn. Soc. N.S.W., 1887, p. 941. = rubella Turn., Proc. Roy. Soc. TFasm.; 1938; p: 92-

(Vict.: Beaconsfield. Tasm.: Derwent Bridge, Triabunna. )

2045. MACHIMIA ATRIPUNCTATELLA Turn. (two-thirds). Trans. Roy. Soc. S. Aust.,
1896, p. 7. (Gayndah, Brisbane, Macpherson Rge., Milmerran.)

2046. MACHIMIA BRACHYTRICHA Turn. (one-half). Proc. Roy. Soc. Tasm., 1926, p. 147.
(Cradle Mt., Railton, Lake Fenton.)

2047. MACHIMIA HOLOCLERA Meyr. (2). Proc. Linn. Soc. N.S.W., 1887, p. 940.
(Brisbane to Victoria, Stanthorpe, Mt. Lofty, Ardrossan.)

2048. MAcHIMIA 10zZONA Low., Trans. Roy. Soc. S. Aust., 1893, p. 174. = episarca Low.,
ibid., 1903, p. 220. (Birchip, Melbourne, Mt. Lofty, Perth.)

2049.4 MACHIMIA UNGUENTARIA Meyr. (4). Hxot. Micro. ii, p. 386. (W. Aust.:
Dundas. )

2050. MACHIMIA METAXANTHA, nN. Sp. (etaéavdos, yellow posteriorly.)

d, 9. 25-35 mm. Head and thorax pale rosy-grey or dull rosy. Palpi with
second joint reaching base of antennae, terminal joint two-thirds to three-fourths;
rosy-whitish. Antennae grey-whitish; ciliations in male 3. Abdomen grey; apices of
segments and tuft whitish-ochreous. Legs whitish-ochreous; anterior pair rosy.
Forewings elongate, costa strongly arched, apex rectangular, termen obliquely rounded;
rosy-grey or dull dosy; a narrow bright rosy costal line, extreme edge whitish except

BY A. JEFFERIS TURNER. ial

towards base; stigmata reddish, usually minute or more often absent; cilia whitish.
Hindwings pale yellow, cilia paler.

New South Wales: Ben Lomond in February. Victoria: Gisborne in February.
West Australia: Perth in December. Eleven specimens.

2051.7; MACHIMIA CRATEROMBRA Meyr., Haxot. Micro., ii, p. 314. (Brisbane.)

2052. MACHIMIA OCELLIFERA Meyr. (1). Proc. Linn. Soc. N.S.W., 1882, p. 488.
Gen. Ins., Oecoph., Pl. 5, f. 86. (Noosa to Hobart, Stanthorpe, Glen Innes.)

2053. MACHIMIA IDIOSEMA Turn. (1). Trans. Royal Soc. S. Aust., 1917, p. 113.
(Duaringa, Brisbane, Charleville, Cunnamulla, Bollon, Castlemaine, Perth.)

2054. MACHIMIA PLAYPORPHYRA, Nl. Sp. (aAatutopdupos, broadly purple.)

6d, °. 17-19 mm. Head and thorax grey. Palpi with second joint reaching base
of antennae, terminal joint one-half; grey-whitish. Antennae grey-whitish with blackish
annulations; ciliations in male two-thirds. Abdomen fuscous; apices of segments pale
ochreous. Legs whitish-ochreous; anterior tibiae rosy; anterior tarsi rosy with pale
ochreous rings, terminal joint fuscous. Forewings with costa gently arched, more
strongly towards base, apex rounded, termen obliquely rounded; pale greyish-ochreous;
a slender rosy line on costa with extreme costal edge whitish; two large purple-fuscous
blotches, one basal with a straight margin at one-third, but not quite reaching costa;
the other apical, its anterior margin rounded from two-thirds costa to three-fourths
_ dorsum; two small purple-fuscous spots, one on midcosta, another beneath middle
of disc; cilia fuscous-purple, apices whitish. Hindwings and cilia fuscous.

Queensland: Brisbane in September; Toowoomba in October; two specimens.

2055. MACHIMIA NOTOPORPHYRA, Nl. SP. (vwromropdupos, purple-backed. )

©. 26 mm. Head and thorax pale ochreous-grey. Palpi with second joint exceeding
base of antennae, terminal joint two-thirds; grey-whitish. Abdomen fuscous; apices of
segments and tuft grey-whitish. Legs ochreous-whitish; tarsi grey; anterior pair rosy.
Forewings moderately dilated, costa strongly arched to middle, thence straight, apex
rectangular, termen sinuate, not oblique; a bright rosy line on costa with extreme costal
edge whitish; a small fuscous spot on base of dorsum; median area with pale reddish
suffusion; markings purple-fuscous; an oblique line from one-third costa expanding in
mid-dise to a large oblong spot extending on margin from mid-dorsum to near tornus; a
discal dot at two-thirds; a finely serrate line from two-thirds costa obliquely to four-,
fifths, thence vertical to tornus; a terminal line; cilia pale rosy, around tornus pale
grey. Hindwings and cilia grey.

South Australia: Mt. Lofty (Blackwood) in November (F. M. Angel); one
specimen.

2056. MACHIMIA CYLICOTYPA, N. Sp. (kvAcKorumos, marked with a circle.)

2. 24 mm. Head and thorax whitish-ochreous. Palpi with second joint exceeding
base of antennae, terminal joint one-half; pale rosy. Antennae pale grey. Abdomen pale
ochreous. Legs whitish; anterior pair pale rosy. Forewings with costa strongly
arched, apex rectangular, termen slightly rounded, slightly oblique; whitish-ochreous;
costal edge faintly rosy; markings fuscous; a broad dorsal stripe from near base to near
tornus; resting on this three spots, two small at one-fourth and middle, one large and
circular at three-fourths; a discal dot at one-third; a few scattered scales in posterior
part of disc; minute terminal dots; cilia whitish-ochreous. Hindwings and cilia pale
ochreous.

Queensland: Clermont in October (E. J. Dumigan); one specimen.

2057. MACHIMIA PICTURATA, nh. Sp. (picturatus, embroidered.)

3, 9. 20-21 mm. Head pale grey. Palpi with second joint reaching base of
antennae, terminal joint 1; white, apical half of second joint dark fuscous. Antennae
white annulated with blackish; ciliations in male 1. Thorax pale grey with dark
fuscous posterior spot. Abdomen pale ochreous. Legs pale ochreous; anterior pair
fuscous with whitish-ochreous tarsal rings. Forewings with costa slightly arched near
base, thence straight, apex round-pointed, termen straight, not oblique; pale grey;
costal edge whitish faintly tinged rosy; markings dark fuscous; a suboblong spot on
base of dorsum narrowly separated from a broader spot, extending to two-thirds; discal

ily, REVISION OF AUSTRALIAN LEPIDOPTERA. OECOPHORIDAE. XIII,

dots at one-third and two-thirds, united by a white line resting on posterior dorsal spot,
with a minute dot on its upper margin; a slender line from beneath two-thirds costa to
upper angle of posterior dorsal spot; a broad crenulate subterminal line; cilia whitish
faintly rosy-tinged. Hindwings pale ochreous suffused with grey except towards base
and costa; cilia pale grey.

Queensland: Roma; two examples bred from larvae feeding on spun-together broad
leaves of sapling Hucalyptus; emerging in Brisbane in July.

2058. MAcHIMIA PHANOZONA Turn. (1). Trans. Roy. Soc. S. Aust., 1896; p. 5.
(Brisbane, Toowoomba.) .

2059. MACHIMIA COCCOSCELA, N. Sp. (KoxxooxedXos, crimson-legged. )

9. 25-28 mm. Head and thorax pale grey. Palpi with second joint reaching base
of antennae, terminal joint three-fifths; rosy. Antennae white, annulated with rosy on
basal fourth, with fuscous on apical three-fourths. Abdomen grey-brownish; sides,
apices of segments and tuft grey-whitish. Legs white; anterior pair bright crimson.
Forewings rather broad, slightly dilated, strongly arched to middle, thence straight,
apex rectangular, termen straight, not oblique; pale grey with a crimson costal line;
a fuscous-reddish basal dot on dorsum, with another shortly above; an outwardly curved
fuscous-purple line from one-fourth costa to two-fifths dorsum; a similar less distinct
line from two-thirds costa to tornus; area between these lines suffused with reddish;
an irregular dark fuscous dentate line from two-thirds costa, incised beneath costa, to
tornus; a terminal line; cilia pale rosy, towards tornus pale grey. Hindwings and cilia
pale grey.

West Australia: Merredin in September; Tammin in October; three specimens.

2060. MAcCHIMIA RUFOGRISEA Meyr. (2). Proc. LInN. Soc. N.S.W., 1882, p. 483.
(Toowoomba, Sydney, Beaconsfield, Birchip, Albany, Denmark.)

2061. MAcHIMIA CRYPTORRHODA, . Sp. (KputToepod0s, with hidden rose-colour. )

°. 36 mm. Head and thorax pale ochreous-grey. Palpi with second joint exceeding
base of antennae, terminal joint one-half; pale ochreous-grey. Antennae whitish annu-
lated with fuscous. Abdomen grey; apices of segments and tuft whitish. Forewings
rather strongly arched, apex rectangular, termen straight, not oblique; pale ochreous-
grey lightly sprinkled with fuscous; costal edge rosy from base to two-thirds; markings
fuscous; stigmata small, distinct, first discal at one-third, plical beyond it, second discal
at two-thirds; a curved line of dots from beneath two-thirds costa to tornus; some
terminal dots; cilia whitish. Hindwings and cilia whitish.

Queensland: Toowoomba in September. New South Wales: Murrurundi in October.
Two specimens.

2062. MAcHIMIA GypsopyeA Meyr. (1). Haxot. Micro., i, p. 220. (Stanthorpe, Ebor,
Mt. Kosciusko, Gisborne, Wilmot, St. Mary’s.)

2063. MACHIMIA TRIPHAENATELLA WIk. (2). xxix, p. 753. Meyr., Proc. Linn. Soc.
INES Wag TESS, 1s A = protoxzantha Meyr., ibid., 1882, p. 477. = phylacopis Meyr., ibid.,
1887, p. 935. = sarcodes Turn., Trans. Roy. Soc. 8S. Aust., 1917, p. 113. = floridula Meyr.,
‘Exot. Micro., i, p. 128. (Yeppoon to Melbourne, Bothwell, Renmark, Waroona, York.)

2064.7 MACHIMIA EUGRAMMA Low., Trans. Roy. Soc. S. Aust., 1894, p. 93. (Duaringa.)

2065.7 MACHIMIA SARCOPTERA Low., Proc. Linn. Soc. N.S.W., 1897, p. 266. (Gisborne. )

2066. MACHIMIA OECOPHORELLA WIK., xxix, p. 760. = paralyrgis Meyr., Proc. Linn.
Soc. N.S.W., 1882, p. 479. (Melbourne, Stawell, Mt. Lofty.)

2067.7 MACHIMIA CROCOXANTHA Meyr., ibid., 1887, p. 934. (Albany.)

2068. MaAcnHimMia EUDOXA Meyr. (1). Ibid., 1885, p. 831. (Melbourne, Mornington,
Mt. Lofty, Quorn.)

2069. MACHIMIA INCLUSELLA Meyr. WIk., xxix, p. 767. Meyr:, Proc. Linn. Soc.
N.S.W., 1882, p. 451. (Sydney.)

2070. MAcHIM1A PERICOSMA Low. (1). Trans. Roy. Soc. S. Aust., 1903, p. 220. (Bris-
bane, Kewell, Adelaide, Renmark.)

2071. MACHIMIA CYPHOPLEURA, N. Sp. (Kudomdevpos, with curved costa.)

°. 25 mm. Head and thorax dull rosy. Palpi with second joint reaching base of
antennae, terminal jcint three-fifths; pale rosy. Antennae whitish with fuscous rings.

BY A. JEFFERIS TURNER. 113

Abdomen ochreous. Legs rosy-ochreous; posterior tibiae whitish-ochreous. Forewings
with costa strongly arched, apex rounded, termen obliquely rounded; ochreous sprinkled
with brownish; a rosy costal line; a purple-fuscous basal patch; pale circular spots on
dorsum at one-third and two-thirds, separated and followed by purple-fuscous suffusion;
a curved line of brownish dots from two-thirds costa to before tornus; a brownish
terminal line; cilia purple-grey. Hindwings and cilia ochreous.

North Queensland: Kuranda (F. P. Dodd); one specimen.

2072. MACHIMIA EUSELMA Meyr. (2). Proc. Linn. Soc. N.S.W., 1882, p. 482. (Stan-
thorpe, Ben Lomond, Armidale, Victoria, Tasmania, Mt. Lofty.)

2073. MACHIMIA PARTHENOPA Meyr. Ibid., 1882, p. 481. (Stanthorpe, Ben Lomond,
Barrington Tops, Vietoria, Tasmania, Mt. Lofty.)

2074. MACHIMIA I0sPILA Meyr. Ibid., 1887, p. 938. (Sydney, Adelaide.)

2075. MACHIMIA METALLOTA Meyr. (2 and a half). Ibid., 1882, p. 486. (Brisbane,
Toowoomba, Murrurundi, Victoria.)

2076. MACHIMIA HABROCOSMA. (93). Ibid., 1882, p. 464. (Macpherson Rge.,
Toowoomba, Injune, Sydney, Stawell.)

2077. MACHIMIA POLYDESMA Low., Trans. Roy. Soc. 8. Aust., 1894, p. 94. (Sydney. )

2078.7 MACHIMIA AMBROSIAS Meyr., Arkiv f. Zool., xiv (15), p. 9. (Qd.: Cedar Creek.)

2079. MAcHIMIA RHODOPIS Meyr., Proc. Linn. Soc. N.S.W., 1887, p. 932. (Albany,
Denmark, Perth.)

2080. MAcCHIMIA INCARNATELLA WIk. (2 and a half.) xxix, p. 754. Meyr., Proc.
Linn. Soc. N.S.W., 1882, p. 468. = inceptella W1k., xxix, p. 759. Meyr., Proc. Linn. Soc.
N.S.W., 1882, p. 469. = crypsichroa Low., Trans. Roy. Soc. S. Aust., 1893, p. 175. (Bris-
bane to Victoria and South Australia.)

2081. MACHIMIA PANDORA Turn. (3). Trans. Roy. Soc. S. Aust., 1917, p. 84. (Cradle
Mt., Mt. Wellington, Lake Fenton.)

2082. MACHIMIA RHIZOBOLA Meyr. (2). Proc. Linn. Soc. N.S.W., 1887, p. 937.
(Brisbane, Toowoomba, Sydney, Victoria. )

2083. MACHIMIA PHYLARCHA Meyr. Ibid., 1882, p. 476. (Stanthorpe, Sydney,
Katoomba, Gisborne, Melbourne.)

2084. MACHIMIA ACMAEA Meyr. Ibid., 1887, p. 936. = charodes Low., Trans. Roy.
Soc. S. Aust., 1920, p. 58. (Nambour, Brisbane, Dalby, Stanthorpe, Mt. Lofty.)

2085. MACHIMIA PLAUSIBILIS Meyr. (2). Hxot. Micro., i, p. 128. (Cape York, Ather-
ton, Yeppoon, Victoria.)

2086. MACHIMIA SIMPLEX Turn. (2 and a half). Trans. Roy. Soc. 8S. Aust., 1896,
p. 5. = sarcophaea Meyr., Exot. Micro., ii, p. 386. (Cape York to Melbourne, Glen Innes,
Injune. )

2087. MACHIMIA MOLLIS, n. sp. (mollis, gentle.)

6, 2. 19-20 mm. Head and thorax dull rosy sprinkled with grey. Palpi with second
joint reaching base of antennae, terminal joint two-thirds; pale rosy. Antennae grey;
ciliations in male 2. Abdomen grey; sides and tuft ochreous-whitish. Legs whitish;
anterior pair pale rosy. Forewings with costa strongly arched, apex rounded-rectangular,
termen slightly rounded, slightly oblique; pale grey; a rosy costal line with whitish
edge; stigmata obscure, fuscous, first discal at one-third, plical beyond it, second discal
before two-thirds; a postmedian line of minute dots, sometimes almost obsolete, from
beneath three-fifths costa rounded in disc, to before tornus; a terminal series of dots;
cilia rosy-grey. Hindwings pale ochreous, sometimes with grey suffusion towards termen;
cilia whitish-ochreous or grey.

New South Wales: Scone in October (H. T. Nicholas); two specimens.

2088. MACHIMIA MILTOSTICHA, D. Sp. (ptATooTrvos, With red lines.)

dG. 24 mm. Head pale grey; face white. Palpi with second joint reaching base of
antennae, terminal joint one-half; pale grey, terminal joint and apex of second. white.
Antennae whitish with fuscous rings, towards base wholly whitish; ciliations in male 1.
Thorax whitish; lateral margins reddish. Abdomen grey; apices of segments and tuft
ochreous-whitish. Legs whitish; anterior pair grey. Forewings with costa moderately
arched, apex obtusely pointed, termen slightly’ rounded, oblique; white with red
markings; a red line closely beneath costa from base to three-fourths, leaving costa

114 REVISION OF AUSTRALIAN LEPIDOPTERA. OECOPHORIDAE. XIII,

white; stigmata minute, fuscous, first discal at one-third, plical absent, second discal
before two-thirds; a red line between discals; a slender line on fold, its base confiuent
with subcostal line; slight reddish suffusion on dorsum; apical fourth of wing suffused
with ochreous; cilia pale grey. Hindwings pale ochreous; cilia whitish.

Queensland: Stanthorpe in December (W. B. Barnard); one specimen.

2089. MACHIMIA HEMITELES Meyr. (1 and a half). Proc. Linn. Soc. N.S.W., 1882,
p. 475. (Stanthorpe to Tasmania.)

2090. MacHIMIA ELAEODES Meyr. (2). Ibid., 1882, p. 474. (Gisborne, Wilson’s
Promontory, Tasmania, Mt. Lofty.)

2091. Macurmta PELoSTIcTA Meyr. (2). Ibid., 1882, p. 473. Turn., Trans. Roy. Soc.
S. Aust., 1917, p. 112. (Brisbane, Tweed Hds., Toowoomba, Murrurundi.)

2092.; MacHIMIA HEMISCIA Meyr., Proc. Linn. Soc. N.S.W., 1882, p. 472. (Sydney.)

2093. MACHIMIA BUBROCHA, n. Sp. (evSpoxos, well knit.)

9. 15 mm. Head and thorax whitish-grey. Palpi with second joint reaching base
of antennae, terminal joint three-fifths; grey-whitish. Antennae pale grey with fuscous
annulations. Abdomen pale grey; tuft whitish. Legs whitish; anterior pair fuscous
with whitish rings. Forewings with costa gently arched, apex pointed, termen slightly
rounded, oblique; whitish-grey faintly purple-tinged, with dark fuscous dots; extreme
costal edge whitish; first discal at one-third, plical beyond it, second discal before
two-thirds; a slender line from plical dot angled upwards, returning to fold, where it
joins a curved series of dots from second discal; a curved series of dots from beneath
two-thirds costa to tornus, indented in middle; a terminal series of dots; cilia whitish-
grey. Hindwings grey-whitish; cilia whitish.

Queensland: Milmerran in September (J. Macqueen); one specimen.

2094. MAcuimia Mimica Meyr. (2). Proc. Linn. Soc. N.S.W., 1887, p. 934. (Ather-
ton, Hidsvold to Tasmania, Mt. Lofty, Margaret R.)

2095. MACHIMIA SEVERA Meyr. (2). Ibid., 1882, p. 470. (Brisbane to Tasmania,
Stanthorpe, Talwood, Mt. Lofty, York.)

2096.; MAcHIMIA COENOSA Meyr. (2). Ibid., 1882, p. 506. (Katoomba.)

2097. MACHIMIA RHODOPLEURA Turn. (1). (Brisbane to Gisborne, Stanthorpe.)

2098. MACHIMIA PYRRHOPASTA, N. Sp. (mvgpomac7os, sprinkled with reddish. )

3d, 9. 20-24 mm. Head and thorax pale ochreous sprinkled with reddish. Palpi
with second joint exceeding base of antennae, terminal joint three-fifths; ochreous-
whitish, terminal joint fuscous. Antennae ochreous-whitish annulated with dark fuscous;
ciliations in male 1. Abdomen grey; apices of segments and tuft whitish. Legs whitish;
anterior pair rosy. Forewings with costa gently arched, apex pointed, termen sinuate,
slightly oblique; pale ochreous with patchy suffusion of reddish and sometimes also
fuscous scales; sometimes a darker suffusion on dorsum from before middle to tornus;
stigmata sometimes fuscous and distinct, but usually obscure or obsolete: first discal at
one-third, plical beyond it, second discal before two-thirds; reddish irroration more
evident in terminal area; cilia ochreous-whitish. Hindwings and cilia ochreous-whitish.

Queensland: Mt. Tamborine in September; Macpherson Rge. (3,000 ft.) in November;
three specimens.

2099. MACHIMIA STENOMORPHA, 0. Sp. (oTevouoppos, Narrow.)

do. 15 mm. Head pale ochreous. Palpi with second joint reaching base of antennae,
terminal] joint one-half; pale grey. Antennae whitish with fuscous annulations; ciliations
in male 2 and a half. Thorax ochreous-brown. Abdomen grey; apices of segments and
tuft whitish. Legs whitish; anterior pair pale grey. Forewings narrow, costa gently
arched, apex rounded, termen obliquely rounded; pale ochreous; markings fuscous;
first discal at one-third, plical slightly beyond it, both minute, second dorsal before two-
thirds, larger; a slender gently curved line from four-fifths costa to tornus; cilia
whitish-ochreous. Hindwings pale grey; cilia whitish.

Queensland: Humundi, near Nambour, in November; Bunya Mts. in October; two
specimens.

2100. MACHIMIA HEBES, n. sp. (hebes, dull.)

6. 22 mm. Head and thorax ‘grey. Palpi with second joint reaching base of
antennae, terminal joint one-half; grey. Antennae grey; ciliations in male 1 and a half.

BY A. JEFFERIS TURNER. 115

Abdomen grey; tuft ochreous-whitish. Legs ochreous-whitish; anterior pair grey. Fore-
wings with costa gently arched, apex pointed, termen slightly rounded, strongly oblique;
whitish with patchy grey and fuscous suffusion; a suffused triangle on dorsum, its apex
reaching above fold; some undefined suffusion about base and apex; stigmata fuscous,
first discal at one-third, second at three-fifths, plical lost in apex of dorsal triangle; cilia
grey-whitish. Hindwings pale ochreous with slight grey suffusion at apex; cilia white
at apex, pale grey elsewhere.

South Australia: Adelaide (Glenelg) in December (D. O. Wilson); one specimen.

2101. MACHIMIA NEPHOSPILA, N. Sp. (vedoomdos, with cloudy spots.)

6. 20 mm. Head and thorax grey. Palpi with second joint exceeding base of
antennae, terminal joint two-thirds; whitish. Antennae whitish with fuscous annula-
tions; ciliations in male 1. (Abdomen missing.) Legs pale grey; posterior pair whitish.
Forewings with costa gently arched, apex rounded, termen obliquely rounded; ochreous-
whitish; costa very faintly tinged with pink; markings fuscous; a median sub-basal dot;
first diseal at one-third, plical well beyond it, second discal at two-thirds, enlarged into
a circular spot; an ill-defined spot or suffusion on dorsum at two-thirds, touching plical
and second discal; a short curved series of subterminal dots; a small terminal spot
beneath apex; cilia whitish. Hindwings grey-whitish; cilia whitish.

West Australia: Kalamunda, near Perth, in January (W. B. Barnard); one specimen.

2102. MACHIMIA PHAEOPORPHYRA Turn. (1). Proc. Roy. Soc. Tasm., 1938, p. 92.
(Derwent Bridge.)

2103. MacHIMIA ARRHODEA Turn. (3). Trans. Roy. Soc. S. Aust., 1917, p. 113. (Miles,
Injune, Birchip, Pinnaroo, Mt. Liebig.)

2104. Macuimia pEMoTICA Meyr. (1). Proc. Linn. Soc. N.S.W., 1882, p. 489. (Ben
Lomond, Ebor, Katoomba, Mt. Kosciusko, Victoria, Tasmania. )

2105.7 MACHIMIA SARCOPHANES Meyr., ibid., 1887, p. 936. (W.A.: Northampton.)

2106.7 Macuimia poLiarcHa Meyr., ibid., 1887, p. 939. (Mt. Lofty.)

2107. MAcHIMIA ENDOLEUCA Meyr. (2). Ibid., 1887, p. 938. = euspilomela Low.,
Trans. Roy. Soc. S. Aust., 1893, p. 175. = agglomerata Meyr., Hxot. Micro., ii, p. 375.
(Birchip, Adelaide, Ardrossan, York, Tammin.)

2108.7 MAcCHIMIA MESODESMA Low., Proc. Linn. Soc. N.S.W., 1898, p. 52. (Vict.:
Leongatha. )

2109. MAOCHIMIA BALIOSTICHA, N. Sp. (Padtoorcxos, with dotted lines.)

9. 30 mm. Head and thorax whitish-grey. Palpi with second joint exceeding base
of antennae, terminal joint three-fifths: whitish. Antennae pale grey. Abdomen grey;
apices of segments whitish. Legs whitish; anterior pair rosy; anterior and middle tarsi
fuscous with whitish rings. Forewings elongate, costa strongly arched to middle,
thence straight, apex rectangular, termen straight, scarcely oblique; whitish-grey with
sharply defined dark fuscous dots; a dot on base of dorsum; an imperfect slender sharply
dentate transverse line at two-fifths, sometimes represented by dots only; a similar line
of dots from one-fifth costa to near mid-dorsum; a curved line of dots from beneath two-
thirds costa to near mid-dorsum; a similar line from three-fifths costa obliquely out-
wards, bent in middle, ending on tornus; a terminal line of dots extending on apical
fourth of costa; costal edge and cilia whitish. Hindwings whitish with slight grey
suffusion towards apex; cilia whitish.

Queensland: Charleville in August; one specimen. I have seen another example
from Milmerran.

2110. MACHIMIA RHODOCHILA, N. Sp. (Go0d0xerAos, rosy-edged. )

6, 2. 32-35 mm. Head and thorax whitish-grey. Palpi short, second joint reaching
base of antennae, terminal joint one-third; whitish. Antennae grey; ciliations in male 2.
Abdomen grey-brown; tuft white. Legs whitish; anterior pair rosy. Forewings elongate-
oblong, costa moderately arched, apex rectangular, termen straight, not oblique; whitish-
grey lightly sprinkled with fuscous; costal edge bright rosy; dorsal edge sometimes dull
reddish; stigmata fuscous, first discal at one-third, plical beyond it, both minute, second
discal at three-fifths larger; cilia whitish-grey. Hindwings and cilia grey.

New South Wales: Murrurundi in September (Dr. B. L. Middleton); two specimens.

2111.7 MAcHIMIA EREBOCROSSA Meyr., Exot. Micro., iii, p. 574. (Burdekin R.)

116 REVISION OF AUSTRALIAN LEPIDOPTERA. OECOPHORIDAE. XIII,

2112. MaAcHIMIA DEFESSA Meyr., ibid., ii, p. 376. (Duaringa.)

2113. Macuimia MyopES Meyr. (One-half). Proc. Linn. Soc. N.S.W., 1882, p. 496.
= orphnophanes Turn., ibid., 1916, p. 371. (Toowoomba, Warwick, Chinchilla, Milmerran,
Stanthorpe, Glen Innes, Murrurundi.)

2114. MACHIMIA INTERJECTA, 1. Sp. (interjectus, placed between, interposed.)

6, 9. 22-25 mm. Head and thorax pale grey. Palpi with second joint exceeding
base of antennae, terminal joint two-thirds to three-fourths; whitish. Antennae whitish
with fuscous annulations; ciliations in male 1. Abdomen pale grey; apices of segments
and tuft whitish. Legs whitish; anterior pair fuscous. Forewings with costa rather
strongly arched, apex rounded-rectangular, termen slightly rounded, slightly oblique;
grey-whitish or pale grey; stigmata fuscous, minute, first discal at one-third, plical
beyond it, second discal before two-thirds; a subterminal line of minute fuscous dots
curved from beneath two-thirds costa to half across disc at four-fifths, thence more
sharply curved to two-thirds dorsum; cilia whitish. Hindwings and cilia grey-whitish
or grey. Greyer than M. achroa, especially in female, without fuscous band on second
joint of palpi as in M. myodes, differing from both in antennal ciliations of male.

Queensland: Bundaberg in September; Stradbroke I. in December; Mt. Tamborine in
November; Stanthorpe in November; six specimens.

2115. MacHIMIA ACHROA Turn. (3). Trans. Roy. Soc. S. Aust., 1896, p. 4. (Cape
York, Townsville, St. Lawrence, Brisbane, Stradbroke I.)

2116. MACHIMIA STYGNODES, N. Sp. (aTvyvwdys, gloomy.)

3, 2. 20-24 mm. Head and thorax grey. Palpi with second joint scarcely reaching
base of antennae, terminal joint two-thirds; second joint whitish or grey with subapical
dark fuscous ring, terminal joint grey. Antennae grey; ciliations in male 1. Abdomen
pale grey; apices of segments and tuft whitish. Forewings with costa rather strongly
arched, apex rounded-rectangular, termen obliquely rounded; grey with dark fuscous
dots; a dot on base of costa; first discal at one-third, plical well beyond it, second discal
before two-thirds, a dot beneath and beyond second; a subterminal line of dots from
well beneath three-fourths costa curved to tornus, not always distinct; a series of
terminal dots extending around apex; cilia grey. Hindwings and cilia darker grey.
Very similar to M. sobriella, best distinguished by the shorter palpi. In M. sobriella the
second joint extends well above the base of the antennae.

West Australia: Merredin in September; three specimens.

2117. MACHIMIA HOMOPOLIA, N. Sp. (6so70XL0s, Uniformly grey.)

dg. 22-25 mm. Head and thorax brownish. Palpi with second joint exceeding base
of antennae, terminal joint three-fifths; fuscous, terminal joint and apex of second
whitish. Antennae grey with fuscous rings; ciliations in male 1. Abdomen fuscous;
apices of segments and tuft whitish. Legs whitish; anterior pair fuscous. Forewings
elongate, costa gently arched, apex rounded, termen very obliquely rounded; pale
brownish-grey; costal edge whitish; stigmata minute, fuscous, first discal at one-fourth,
plical beyond it, second discal at three-fifths; cilia grey-whitish. Hindwings and cilia
greyish.

New South Wales: Adaminaby in October; three specimens.

2118. MAcCHIMIA EBENOSTICTA, N. Sp. (€Bevoorixros, black-spotted. )

do. 15 mm. Head and thorax pale grey. Palpi with second joint reaching base of
antennae, terminal joint three-fourths; whitish, second joint with terminal and sub-
terminal blackish rings. Antennae whitish with blackish annulations; ciliations in
male 1. Antennae pale grey; apices of segments and tuft ochreous-whitish. Legs
whitish; anterior pair blackish; anterior and middle tarsi whitish wth blackish rings.
Forewings with costa gently arched, apex rounded, termen obliquely rounded; whitish-
grey with blackish dots and a few scattered scales; first discal at one-fourth, plical
beyond it, second discal at middle; a dot on three-fourths dorsum; second discal
connected by a line of dots with one-third costa, and by another line with three-fourths
dorsum; a fine partly interrupted line from two-thirds costa to tornus, indented beneath
costa and in middle; a terminal series of dots; cilia whitish-grey. Hindwings and cilia
grey.

Queensland: Injune in February (W. B. Barnard); one specimen.

ss

BY A. JEFFERIS TURNER. ILI?)

2119. MacHIMIA RESTRICTA Meyr. (1 and a fourth). Exot. Micro., ii, p. 310. (Cape
York, Townsville, Yeppoon, Caloundra, Warwick, Injune.)

2120. MACHIMIA SOBRIELLA Meyr. (1). Proc. Linn. Soc. N.S.W.; 1882, p. 495. (Cape
York to Hobart, Adelaide, Pt. Lincoln.)

2121. MAcHIMIA PASTEA Turn., Proc. Roy. Soc. Tasm., 1926, p. 147. (Tasmania. )

2122. MACHIMIA BALANOTA Meyr., Proc. Linn. Soc. N.S-W., 1888, p. 1643. (Warragul.)

2123. MACHIMIA HOLOCHRA, nN. Sp. (dAwypos, wholly pale.)

6. 18-22 mm. Head and thorax ochreous-grey-whitish. Palpi with second joint
exceeding base of antennae, terminal joint one-half; whitish. Antennae whitish with
fuscous annulations; ciliations in male 3. Abdomen grey; apices of segments and tuft
whitish. Legs grey; tarsi with white rings; posterior pair white. Forewings with costa
gently arched, apex pointed, termen slightly rounded, slightly oblique; ochreous-whitish;
dots and slight irroration fuscous; stigmata minute or partly obsolete, first discal at one-
third, plical beyond it, second discal at three-fifths; a very fine outwardly oblique line
from two-thirds costa, angled in disc, and thence subterminal to tornus; cilia whitish.
Hindwings and cilia whitish.

West Australia: Denmark in March; Waroona in January (W. B. Barnard); six
specimens. Type in Queensland Museum.

2124. MACHIMIA MICROPTERA Turn. (1 and a half). Proc. Linn. Soc. N.S.W., 1916,
p. 372. (Mt. Tamborine, Macpherson Rge.)

2125. MAcHIMIA ANAEMICA Turn. (2). Ibid., 1916, p. 372. (Brisbane.)

2126. MAcHIMIA NEOCHLORA Meyr. (1). Ibid., 1882, p. 495. (Darwin, Townsville,
Duaringa, Toowoomba. )

2127. MACHIMIA MELLICHROA Low., ibid., 1897, p. 267. (Gisborne.)

2128. MACHIMIA MESOGAEA Turn., ibid., 1916, p. 371. (Adavale.)

2129. MACHIMIA ALBULA, n. sp. (albulus, whitish.)

6. 16-17 mm. Head white. Palpi with second joint exceeding base of antennae,
terminal joint two-thirds; grey. Antennae whitish with fuscous rings; ciliations in
male 3. Thorax whitish-grey. Abdomen grey; apices of segments and tuft whitish.
Legs whitish; anterior tarsi with fuscous rings. Forewings narrow, dilated posteriorly,
costa gently arched, apex pointed, termen slightly rounded, oblique; whitish-grey with
fuscous markings; a dot on base of costa, a sub-basal curved line of three dots; first
discal at one-third, plical beyond it, second discal double, at two-thirds; sometimes a
slender dentate line from two-fifths costa to mid-dorsum; a small spot on two-thirds
costa connected by an outwardly curved line with dorsum before tornus; two subapical
costal dots; a dotted terminal line; cilia white. Hindwings and cilia whitish.

Queensland: Injune in February; Carnarvon Rge. in December (W. B. Barnard);
three specimens. Type in Queensland Museum.

2130. MACHIMIA RHAPHIDUCHA, Nl. Sp. (¢adidouvxos, needle-streaked. )

6.. 28 mm. Head whitish. Palpi with second joint three times length of face, white
sprinkled with fuscous, basal third dark fuscous externally. Antennae grey-whitish
sprinkled with dark fuscous; ciliations in male 2 and a half. Thorax whitish-grey.
Abdomen grey; sides and tuft whitish. Legs whitish; anterior pair fuscous anteriorly.
Forewings elongate, costa gently arched, apex obtusely pointed, termen slightly rounded,
oblique; whitish with slender blackish streaks; a short streak from base of costa,
another in cell, and several very slender streaks between veins running to costa and
termen; cilia white. Hindwings grey-whitish; cilia white.

Queensland: Tweed Hds. (Burleigh) in September (W. B. Barnard); two specimens.
Type in Queensland Museum.

2131. MACHIMIA CUPHOSEMA, 1. Sp. (Kovdoonpuos, lightly marked.)

do. 20 mm. Head and thorax ochrecus-whitish. Palpi with terminal joint two-
thirds; whitish. Antennae whitish with fuscous annulations; ciliations in male 1.
Abdomen grey-whitish; apices of segments fuscous; tuft ochreous-whitish. Legs whitish;
anterior pair fuscous. Forewings with costa slightly arched, apex pointed, termen
obliquely rounded; ochreous-whitish with slender fuscous markings; an oblique line
from two-fifths costa half across wing; an oblique line from two-thirds costa, angled

118 REVISION OF AUSTRALIAN LEPIDOPTERA. OECOPHORIDAE. XIII,

inwards beneath middle to two-thirds dorsum; an irregularly dentate subterminal line;
an interrupted terminal line; cilia pale grey. Hindwings and cilia whitish.

Queensland: Macpherson Rge. (3,500 ft.) in December; one specimen.

2132. MACHIMIA DYSTHEATA, N. Sp. (dvobeaTos, insignificant.)

g. 18 mm. Head, thorax and abdomen grey-whitish. Palpi with terminal joint one-
half; whitish. Antennae whitish; ciliations in male 1. Legs whitish; anterior pair grey.
Forewings suboblong, costa gently arched, apex rounded, termen obliquely rounded;
grey-whitish; stigmata minute, fuscous, first discal at middle, plical beyond it, second
discal at three-fourths; some minute obscure costal and terminal dots; cilia whitish.
Hindwings and cilia whitish.

North Queensland: Cape York in October (W. B. Barnard); one specimen.

2133. MACHIMIA LERA, nN. Sp. (Anpos, trifling.)

¢g. 14 mm. Head pale grey. Palpi with terminal joint three-fifths; whitish.
Antennae whitish with fuscous annulations; ciliations in male 2.. Thorax white.
Abdomen pale grey. Forewings narrow, dilated posteriorly, costa gently arched, apex
rounded, termen slightly sinuate, slightly oblique; white; markings fuscous; a basal
costal dot and another at two-fifths; a small costal triangle before apex; a discal dot
at two-thirds; terminal edge fuscous; cilia ochreous-whitish. Hindwings and cilia
whitish.

North Queensland: Innisfail in November; one specimen.

2134. MACHIMIA METAGYPSA, N. Sp. (werayuwos, white posteriorly.)

9. 22 mm. Head and thorax pale grey. Palpi with second joint reaching base of
antennae, terminal joint three-fifths; pale grey. Antennae grey-whitish with dark
fuscous annulations. Abdomen white. Legs white; anterior pair grey with white tarsal
rings. Forewings rather narrow, costa slightly arched, apex obtuse, termen obliquely
rounded; pale grey; markings pale fuscous, indistinct; first discal at one-third, plical
beyond it, second discal before two-thirds; some fuscous sprinkling between second discal
and tornus; traces of a subterminal line; cilia whitish. Hindwings and cilia white.

West Australia: Albany in November; one specimen.

2135. MACHIMIA TRISEMA Low., Trans. Roy. Soc. S. Aust., 1907, p. 114. (Mackay.)

2136. MACHIMIA RHOECOZONA, N. Sp. (gorkofwvos, with crooked girdle.)

6d. 16 mm. Head and thorax pale yellow. Palpi with second joint reaching base
of antennae, terminal joint three-fifths; pale yellowish. Antennae grey; ciliations in
male 2 and a half. Abdomen grey. Legs fuscous; posterior pair ochreous-whitish. Fore-
wings somewhat dilated, costa moderately arched, apex pointed, termen sinuate, scarcely
oblique; pale yellow; markings fuscous; a narrow transverse fascia from costa before
middle to mid-dorsum, strongly bent in middle to form a posterior angle; a narrow
terminal fascia; cilia fuscous, apices whitish-ochreous except on apex and tornus. Hind-
wings and cilia pale grey.

New South Wales: Sydney in December: Wentworth Falls, near Katoomba, in
November (G. M. Goldfinch); two specimens. Type in Australian Museum.

2137. MACHIMIA XANTHISMA Turn. (2). Trans. Roy. Soc. S. Awst., 1917, p. 114.
(Bend Lomond, Ebor.)

171. Gen. EocHrois Meyr.

Proc. Linn. Soc. N.S.W., 1882, p. 448.

Palpi ascending, recurved; second joint reaching base of antennae; terminal joint
shorter than second, stout, acute. Antennae without pecten. Forewings with 7 and 8
stalked, 7 to termen. Hindwings with 5 from middle of cell. Type, H. pulverulenta.

2138. HocHROIS PULVERULENTA Meyr., Proc. Linn. Soc. N.S.W., 1882, p. 454.
(Toowoomba, Stanthorpe, Ebor, Katoomba, Cooma, Victoria.)

2139. EocHROIS ACUTELLA WIK., xxix, p. 766. (Hmerald, Pt. Macquarie, Sydney.)

2140. EocHROIS EBENOSTICHA Turn., Trans. Roy. Soc. S. Aust., 1917, p. 110. (Albany,
Perth.)

Ee

a ee Se eS eee es ee ee

BY A. JEFFERIS TURNER. 119

172. Gen. Lreprpozancia Turn.

.Proc. Linn. Soc. N.S.W., 1916, p. 375.

Palpi very long, ascending, recurved; second joint three times length of face,
thickened and slightly rough anteriorly; terminal joint shorter than second, slender,
acute. Antennae without basal pecten. Forewings with 7 and 8 stalked, 7 to apex.
Hindwings with 5 from middle of cell. Male unknown. Monotypical.

2141. LeEpPIDOZANCLA ZATREPHES Turn., Proc. Linn. Soc. N.S.W., 1916, p. 376. (Bris-
bane, Injune.)

173. Gen. HopromoreHa Turn.

Proc. Linn. Soc. N.S.W., 1916, p. 3738.

Palpi long, ascending, recurved; second joint reaching or exceeding base of antennae,
moderate, smooth; terminal joint shorter than second, more or less slender, acute.
Antennae without basal pecten. Forewings with 7 and 8 stalked, 7 to apex. Hindwings
with 5 from middle of cell or from nearer 6 than 4. Type, H. abalienella. Seven species.

2142. HopLoMoRPHA ABALIENELLA WIk., xxix, p. 762. = colonias Meyr., Proc. Linn.
Soc. N.S.W., 1887, p. 942. (Atherton, Brisbane, Tyringham, Scone, Bairnsdale.)

2143. HopLoMorRPHA PORPHYRASPIS Turn., Trans. Roy. Soc. S. Aust., 1896, p. 8. (Bris-
bane, Mt. Tamborine, Macpherson Rge., Toowoomba. )

2144. HopromorPHA TERATOPA Meyr., Haxot. Micro., ii, p. 310. (Killarney, Lismore.)

2145. HorpLoMORPHA CAMELAEA Meyr., Proc. Linn. Soc. N.S.W., 1887, p. 943. (Stan-
thorpe, Toowoomba, Sydney, Katoomba, Beechworth, Gisborne, Beaconsfield.)

2146. HopLomMorPHA EPICOSMA Turn., ibid., 1916, p. 374. (Mt. Tamborine, Uki.)

2147. HopLomorPHA CAMINODES Turn., ibid., 1916, p. 375. (Brisbane, Mt. Tamborine,
Tweed Hds., Killarney, Murwillumbah.)

2148. HopLoMORPHA STYPHLODES, Nn. Sp. (oaTvddwdns, stern.)

3, 2. 21-26 mm. Head and thorax fuscous. Palpi with second joint exceeding base
of antennae, terminal joint two-thirds; pale fuscous. Antennae pale grey annulated with
dark fuscous; ciliations in male 1 and a fourth. Abdomen fuscous; apices of segments
and tuft whitish-ochreous. Legs whitish; anterior pair fuscous. Forewings with costa
slightly arched, apex rounded, termen obliquely rounded; pale fuscous with dark fuscous
dots; costal edge whitish; discals approximated, first discal at one-third, plical beneath
it, second discal at middle; a curved subterminal line of minute dots, sometimes indis-
tinct; cilia grey. Hindwings grey; cilia grey-whitish with darker sub-basal line.

Queensland: Macpherson Rge. (3,000 ft.) in December and January; eight specimens.

Alphabetical List of the Species of Machimia.

absumptella (2007), achroa (2115), acmaea (2084), albula (2129), alma (2008),
amata (2035), ambrosias (2078), anaemica (2125), arrhodea (2103), atoecha (2040),
atripunctatella (2045), balanota (2122), baliosticha (2109), brachytricha (2046),
brevicilia (2038), callianassa (2000), carnea (2043), cholodella (2015), coccoscela (2059),
coenosa (2096), complanula (2017), conspersa (1998), costimacula (2028), craterombra
(2051), crocoxantha (2067), cryptorrhoda (2061), cuphosema (2131), cylicotypa (2056),
cyphopleura (2071), defessa (2112), delosticta (2014), demotica (2104), diphracta (2004),
dystheata (2132), ebenosticta (2118), elaeodes (2090) endoleuca (2107), eoxantha (2039),
erebocrossa (2111), ewbrocha (2093), eudoxa (2068), eugramma: (2064), euselma (2072),
gypsopyga (2062), habrocosma (2076), habroptera (2021), habroschema (2029), haplo-
ceros (2034), hebes (2100), hemiscia (2092), hemiteles (2089), holochra (2123), holoclera
(2047), homopolia (2117), hypertricha (1999), idiosema (2053), incarnatella (2080),
inclusella (2069), interjecta (2114), iospila (2074), iozona (2048), laetiferana (2002),
leiochroa (2005), lera (2133), leucerythra (2033), limbata (2036), loxomita (2023),
melanoplecta (2003), mellichroa (2127), mesodesma (2108), mesogaea (2128), metagypsa
(2134), metailota (2075), metaxantha (2050), metriopis (2041), micromita (2009),
microptera (2124), miltopsara (2011), miltosticha (2088), mimica (2094), mitescens
(2010), moderatella (2044), mollis (2087), myodes (2113), neochlora (2126), nephospila
(2101), notoporphyra (2055), ocellifera (2052), ochra (2032), oecophorella (2066),
oncospila (2025), pandora (2081), parthenopa (2073), pastea (2121), pelosticta (2091),

120 REVISION OF AUSTRALIAN LEPIDOPTERA. OECOPHORIDAE. XIII.

pericosma (2070), phaeoporphyra (2102), phanozona (2058), phoenopis (2006), phylarcha
(2083), picturata (2057), platyporphyra (2054), plausibilis (2085), poliarcha (2106),
polydesma (2077), praepedita (2013), pudica (2030), pyrrhopasta (2098), repandula
(2042), restricta (2119), rhaphiducha (2130), rhizobola (2082), rhodochila (2110),
rhodopis (2079), rhodopleura (2097), rhodoxantha (2027), rhoecozona (2136), rufa (2022),
rufescens (2020), rufogrisea (2060), sarcophanes (2105), sarcoptera (2065), sarcoxantha
(2001), sericata (2016), serva (2012), severa (2095), similis (2037), simplex (2086),
sincerella (2026), sobriella (2120), spatiosa (2031), stenomorpha (2099), stenorrhoda
(2018), stygnodes (2116), submissa (2019), thaumasticha (1997), triphaenatella (2063),
trisema (2135), umbratica (2024), unguentaria (2049), xanthisma (2137).

121

STUDIES ON AUSTRALIAN MARINE ALGAE. II.
NOTES EXTENDING THE KNOWN GEOGRAPHICAL RANGE OF CERTAIN SPECIES.
By VALERIE May, M.Sc. (C.S.1.R., Marine Biological Laboratory, Cronulla, N.S.W.*)

[Read 25th July, 1945.]

During any detailed systematic study of a section of a flora, one accumulates pieces
of information concerning other members of that flora as a whole. During the
programme of work on agar-producing algae on which the writer is at present engaged,
certain facts have emerged regarding the distribution of certain other algae in
Australia. These are discussed below. Unless otherwise stated, the specimens recorded
in this paper are either in. my own Herkarium or in the National Herbarium, Sydney.

CHLOROPHYCEAE.
VALONIA CONFERVOIDES Harv.

Prior to my recording, in 1940, that a specimen of this plant had been found at
Angowrie, northern New South Wales, it was known in Australia only from the north
Queensland coast (specimen from Dunk Island in Brisbane Herbarium, Queensland).
The following additional collections are now recorded:

Locality. Date. Collector. Notes.
Noosa Heads, Qd. xi.43. Valerie May. Drift.
Collaroy, near Sydney, N.S.W. 24.xi.44. Valerie May. Drift.
Collaroy, near Sydney, N.S.W. 27.xii.44. Valerie May. Drift.

Long Reef, near Sydney, N.S.W. 3.xii.44. Valerie May. Attached to rock; not
exposed at low tide.
APJOHNIA LAETEVIRENS Harv.
New Record for New South Wales.
Previously this plant had been collected from South Australia, Victoria, and “rarely

in Tasmania” (Lucas, 1936). The following collection is a new record for New
South Wales:
Locality. Date. Collector. Notes.
Collaroy, near Sydney, N.S.W. 7.vii.44. Valerie May. Drift after storm.
MELANOPHYCEAE.

HypDROCLATHRUS CLATHRATUS (Bory) Howe.
New Record for New South Wales.

Previously this plant had been collected in both Victoria and Queensland, as well
as in South Australia and Western Australia, so that it is surprising that no previous
‘record exists of its occurrence in New South Wales. Collections from southern
Queensland are included for the sake of completeness.

Earlier records in Australia referred to this species under the name of dH.
cancellatus Bory, which, according to Setchell and Gardner (1925), should be included

in H. clathratus.

* Contribution No. 40 from the Laboratory.

122 STUDIES ON AUSTRALIAN MARINE ALGAE. II,

Locality. Date. Collector. Notes.
Urangan, near Maryborough, Qd. 18.viii.43. Valerie May. Drift.
Near mouth of Mary River, Qd. 19.viii.48. Valerie May. Trawled.
Hervey Bay, near Maryborough, 25.vili.43. Valerie May. Trawled.
Qd.

Moreton Big Banks, Moreton Bay, 2.x11.43. Valerie May. Attached.
Qd.

Port Hacking, N.S.W. v.44. R. Bouchier.

Bundeena and Lillipilli, Port 8.ii1.45. Valerie May. Trawled.

Hacking, N.S.W.

ENDARACHNE BINGHAMIAE J. Ag.
New Record for Australia.

Setchell and Gardner (1925), in figuring this plant, have emphasized its macro-
scopic resemblance to Ilea Fascia (Muell.) Fries. This resemblance is presumably
responsible for Hndarachne not being recognized earlier in Australia, for, near Sydney,
it appears to be quite prevalent. One sample of it was collected, but left unnamed,
by the late Mr. A. H. S. Lucas. Our plant matches well the figures given by Setchell

and Gardner (1925). The localities from which it has so far been collected are as
follows:

Locality. Date. Collector. Notes.
Bondi, near Sydney, N.S.W. ® Vel On AN He Seeduulcase
Mona Vale, near Sydney, N.S.W. 13.v.44. Valerie May.
Mona Vale, near Sydney, N.S.W. 11.i11.45. Valerie May.
Long Reef, near Sydney, N.S.W. 31.vii.44. Valerie May. Growing on rock
Newport, near Sydney, N.S.W. 21.x.44. Valerie May. \ platforms on rocks
Newport, near Sydney, N.S.W. 18.11.45. Valerie May. | exposed at low tide.
Kurnell, near Sydney, N.S.W. 16.xi1.44. Valerie May.
Stanwell Park, near Sydney, 26.i111.45. Valerie May. J
N.S.W.
RHODOPHYCEAE.

NEMALION MULTIFIDUM (W. and M.) J. Ag.
New Record for Australia.

The identification of this plant is based on comparison of material with Harvey .
(1846-51, Pl. 36), and with Newton (1931, pp. 256-7). The microscopic anatomy agrees
well with these references, but the dichotomous branching is rather less frequent
than appears usual for the species. In this character the species approaches
N. elminthoides Batt. (= N. lwbricum Duby).

This is the only record of the genus Nemalion in Australia. An earlier record
(N. insigne Harv.) is now regarded as belonging to another genus (Helminthocladia
do INE Ne

All collections have been made from ocean headlands, from rock faces exposed
between tides, but receiving surf spray for much of the time. Both cystocarps and
antheridia were present in material of the 11.iii1.45 collection listed below.

Locality. Date. Collector. Notes.
Mona Vale, near Sydney, N.S.W. 11.v.44. Valerie May.
Mona Vale, near Sydney, N.S.W. 11.11.45. Valerie May. Fertile monoecious.
Newport, near Sydney, N.S.W. 21.x.44. Valerie May.
Stanwell Park, near Sydney, 26.111.45. Valerie May.
N.S.W.

EUCHEUMA GELATINAE (Hsp.) J. Ag.
New Record for south Western Australia. '
EH. speciosum (Sond.) J. Ag. is the species of this genus best known in Australia,

being collected from south Western Australia and also, in smaller quantities, from
Tasmania.

BY VALERIE MAY. 123

BE. gelatinae (Esp.) J. Ag. was recorded by Mme. Webei-van Bosse (1928) as from
“Cotes de la Nouvelle Hollande’”’. Recently I have seen samples of what appears to be
this species, collected from the following locality:

Locality. Date. Collector. Notes.
Abrolhos Islands, off W. Aust. v.44. D. L. Serventy. Drift.

These algae were collected in connection with the Western Australian agar
production programme, and I understand from Dr. Serventy that the Western
Australian manufacturers have been unable to detect a difference in the agar yield
obtained when using H. gelatinae as compared with that obtained when using the more
usual H. speciosum. 3

EHUCHEUMA MURICATUM (Gmel.) Weber-van Bosse.
New Record for Western Australia.

This species—recorded by the synonym E. spinosum (L.) J. Ag.—was known from
Thursday Island and Dunk Island, off the coast of Queensland (the latter specimen is
in the Brisbane Herbarium, Queensland) and has now been collected from the
following locality:

Locality. Date. Collector. Notes.
Pelsart Island, Abrolhos Group, xi. 44. D. L. Serventy. Drift on west
off W. Aust. (lagoon) side

of island.

It seems likely that further collections of tltis plant may be expected from more
northerly areas of Western Australia.

ANTITHAMNION PLUMULA (Ellis) Thur.
New Record for New South Wales.

This plant had been collected previously in Australia at Port Phillip Heads by
J. Br. Wilson and at Georgetown, Tasmania, by W. H. Harvey (both collections are now
in the National Herbarium, Sydney), but Harvey (1863) records it as “rare”’.

It is now reported by local fishermen as being extremely prevalent along the
whole New South Wales coast from Bateman’s Bay to Jervis Bay, at a depth ranging
from 20 to 50 fathoms, in areas which the fishermen knew previously as clean, sandy
bottoms and on which they have trawled for flathead during the last two years. This
plant has, in fact, been suggested as a possible menace to the flathead trawling
industry. It may be noteworthy that recently also samples of it were obtained from
D’Entrecasteaux Channel, Tasmania.

Locality. Date. Collector. Notes.
Off Ulladulla, N.S.W. liebe ken Sheard: Trawled, 20 fathoms.
D’Entrecasteaux Channel, Tasm. 8.ix.44. J. A. Tubb. Trawled, 5 fathoms.

LITHOTHAMNION LICHENOIDES (Ell. and Sol.) Heydrich forma PATENA
(H. and H.) Foslie.
New Record for New South Wales.
This rather distinctive form was previously known from Victoria and South
Australia. It is now reported as follows:

Locality. Date. Collector. Notes.
Long Reef, near Sydney, N.S.W. 3.xii.44. Valerie May. On attached Corallina
sp. on rock platform.

SUMMARY.

New collections are reported of certain marine algae, extending the known range
of geographical distribution in Australia of the species concerned. Four species are
recorded for the first time in New South Wales, one species is recorded for the first
time in Western Australia and one in south Western Australia.

Endarachne Binghamiae J. Ag. and Nemalion multifidum (W. and M.) J. Ag. are
recorded for the first time in Australia.

124 STUDIES ON AUSTRALIAN MARINE ALGAE. II.

REFERENCES.

HARVEY, W. H., 1846-51.—Phycologia Britannica. Vol. 3. London.
, 1863.—Synoptic Catalogue to Phycologia Australica. Vol. 5. London.

Lucas, A. H. S., 1936.—The Seaweeds of South Australia, Part 1. Introduction and the
Green and Brown Seaweeds. Adelaide.

May, VALERIE, 1940.—Notes and Exhibits. Proc. LINN. Soc. N.S.W., 65 (1-2) : xxx-xxxi.

NEWTON, Lity, 1931.—A Handbook of the British Seaweeds. London.

SETCHELL, W. A., and GARDNER, N. L., 1925.—The Marine Algae of the Pacific Coast of North
America, Part 3. Melanophyceae. Univ. Calif. Publ. in Bot., 8 (3): 383-898, Pls. 34-107.

WHBHR-VAN Bossnp, A., 1928.—Liste des Algues du Siboga. iv. Rhodophyceae, 3 me. part.
Siboga Expéditie. Monographie 59 d, 393-533, Pls. xi-xvi.

125

A BRYOZOAN FAUNA FROM THE LAKE’S CREEK QUARRY, ROCKHAMPTON,
QUEENSLAND.

By Joan Crockrorp, M.Se., Linnean Macleay Fellow of the Society in Palaeontology.
(Twelve Text-figures. )

{Read 27th June, 1945.]

INTRODUCTION.

A collection of specimens rich in Bryozoa from in and around the Lake’s Creek
Quarry near Rockhampton in Queensland was recently lent to me by Mr. O. A. Jones
of the University of Queensland. A brief summary of the opinions published regarding
the age of the Lake’s Creek beds, and of the Trachypora wilkinsoni* horizon which occurs
behind the Quarry, has been given by Bryan and Jones (1944, 43, 71). The commonest
species in the bryozoan fauna are described in this paper, and amongst the fourteen
species and varieties here described or recorded, there are six previously described
species; these six species are all forms which are known only from the Lower Permian.

The forms present in the collection which I have used are almost exclusively
Fenestrellinidae; a few poorly preserved Batostomellidae also occur, but they are not
well enough preserved for description. The specimens are preserved almost entirely as
casts, which are often crushed and distorted; many of these casts do, however, show
their specific characters very perfectly indeed.

Of the six previously described species which are here recorded from Lake’s_ Creek
Quarry, Fenestrellina aspratilis (Bassler), 1929, occurs in both the Bitaoeni and Basleo
-Beds in Timor, and is probably the same as F.. girtyi Elias, 1937, from the Dark Lime-
stone, Guadalupian Group, Guadalupe Mts., Texas; Fenestrellina granulifera Crockford,
1941, occurs in New South Wales in the Fenestella Shales of the Branxton Beds, Upper
Marine Series, and in Tasmania in the Permian at Huon Rd., Mt. Wellington;
Fenestrellina canthariformis Crockford, 1941, also occurs in the Fenestella Shales, and
occurs in the Springsure district of Queensland at Consuelo Creek, 2 miles above Cattle
Creek, Horizon A in the Dilly Stage of Reid (1930, 95), which is also loc. 9 in
Whitehouse’s list of fossils at the end of Reid’s paper; Polypora woodsi Etheridge, 1892,
occurs at several localities in the Fenestella Shales, and at Ulladulla, on the South Coast
of New South Wales, in the Permian at Marlborough, Tasmania, in both the Callytharra
and Nooncanbah Series in Western Australia, and in the Bitaoeni Beds of Timor (as
Polypora tripliseriata Bassler, 1929); Polypora woodsi also occurs at the same locality
as F’. canthariformis near Springsure, and Etheridge has recorded it from ‘‘below Sonoma
Rd. crossing, Coral Ck., Bowen R.”’, Queensland; Polypora virga Laseron, 1918, is
common in the Fenestella Shales of the Upper Marine Series, but does occur as well in
the Allandale Stage of the Lower Marine Series, and it also occurs at Marlborough,
Tasmania; Minilya duplaris Crockford, 1944, occurs in Western Australia from the
Callytharra Series through to the Wandagee and Nooncanbah, and is one of the
commonest species in the Western Australian Permian; it also occurs in the Northern
Territory in the Port Keats Bore, and at the same locality as F. canthariformis and
P. woodsi near Springsure; this species is probably the same as the species figured as
Fenestella perelegans Meek, 1871, by Waagen and Pichl (1885) from the Middle Productus
Limestone of the Salt Ra.; (this Indian species is not the same as the specimens
originally figured by Meek from North America). So far as the age of the strata at

*Thamnopora wilkinsoni (Etheridge), Hill, 1943 (J. Roy. Soc. W. Aust., 27 (for 1940-41):
Glo)

0

126 BRYOZOAN FAUNA FROM ROCKHAMPTON,

Lake’s Creek Quarry is concerned, therefore, the evidence from these Bryozoa is not
sufficient to give close correlation with any one stage of the Permian sequence in
Western Australia, Timor, or New South Wales, although they do indicate that the age
of the Lake’s Creek beds is Lower Permian.

Reid (1930, 41-2) stated that the Lake’s Creek Quarry and the Gympie Beds were
tied to the Neerkol Series of Upper Carboniferous age “to some extent, palaeontologically,
notably with Protoretepora, Rhombopora laza, Poiypora smithii, and, provisionally,
Aviculopecten squamuliferus, Productus subquadratus (Lake’s Creek type), and possibly
Stenopora”’. So far as the four bryozoans mentioned in support of this correlation are
concerned, there could be no evidence whatsoever in the presence of the forms mentioned
to support a correlation between Lake’s Creek Quarry and the Neerkol. Protoretepora,
in the sense in which the name has been used in the past, refers to the presence of any
coarse fenestrate bryozoan, generally to some species of Polypora: in Hastern Australia

~e_
°
=~

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Qo

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Text-fig. 1.—Fenestrellina canthariformis Crockford. Celluliferous surface of a specimen
(F.7970, Univ. Queensland Colln.) from the Trachypora horizon behind Lake’s Creek Quarry,
x¢ IM;

Text-fig. 2—Minilya duplaris Crockford. Celluliferous surface of a specimen (F.7973,
Univ. Queensland Colln.) from the Trachypora horizon behind Lake’s Creek Quarry, x 10.

Text-fig. 3.—Fenestrellina sparsinodata, n. sp. Celluliferous surface of the holotype, x 10.

Text-fig. 4.—Fenestrellina granulifera Crockford. Celluliferous surface of a specimen
(F.7962, Univ. Queensland Colln.) from Lake’s Creek Quarry, x 10.

Text-fig. 5.—Fenestrellina aspratilis Bassler. Celluliferous surface of a specimen (F.7961B,
Univ. Queensland Colln.) from Lake’s Creek Quarry, x 10.

Text-fig. 6.—Fenestrellina simulatrix, n. sp. Celluliferous surface of the holotype. x 10.

BY JOAN CROCKFORD. ILzAY

Polypora is. fairly common in almost all the Upper Palaeozoic deposits in which
fenestellids occur, and the genus itself ranges from the Ordovician to the Permian, so
that, unless the same species were proved to occur in the Neerkol and at Lake’s Creek
(and this has not been done), the presence of this form is of no account in correlation.
Rhombopora laxa (Etheridge), 1872, was figured with the original description
(Etheridge, 1872, 332, Pl. xxv, figs. 2, 2a), and has also been figured by R. Etheridge, jnr.
(1892, 224, Pl. ix, figs. 8, 9), but neither of these descriptions and figures are sufficiently
clear for recognition of this form, since none of the characters important in the deter-
mination of ramose Bryozoa is described or illustrated. Polypora smithii Etheridge,
1892, also could not be recognized from the original description and figures, which cover
more than one species; and Stenopora, which is common in the Permian, occurs, though
: rarely, in the Carboniferous also, so that the occurrence of the genus does not indicate
any special horizon. None of the bryozoan species quoted by Reid, therefore, could be
used for correlation between the Neerkol and Lake’s Creek Beds.

Whitehouse (1928, 1929) has also discussed the age of the Lake’s Creek Quarry
beds, and has suggested that they should be correlated with the horizon of the Greta
Coal Measures in New South Wales and the Yatton Limestone and Collinsville Coal
Measures of the lower part of the Middle Bowen Series in the Bowen River Coalfield
in Queensland. I have, unfortunately, no specimens from the Bowen River Coalfield
with which I could compare these specimens. Dr. Whitehouse placed all the beds below
the top of the Greta Coal Measures in the Carboniferous; at present the boundary
between the Permian and the Carboniferous is placed lower in the sequence, being
placed by different workers either at the base of the Lower Marine Series or at the
Hurydesma cordatum horizon in the lower part of this Series.

DESCRIPTION OF SPECIES.
Order CrRYPTOSTOMATA Vine.
Family FENESTRELLINIDAE Bassler.
Genus FENESTRELLINA d’Orbigny, 1849.

FENESTRELLINA SIMULATRIX, Nn. sp. Text-fig. 6.

Holotype: F7950A, Univ. Queensland Colln.

Locality: Lake’s Creek Quarry, Rockhampton, Queensland (holotype); Trachypora
horizon behind Lake’s Creek Quarry (F7951A, B, Univ. Queensland Colln.).

Fine Fenestrellina, with 3 zooecia to a fenestrule; carina slight; nodes sharp, small,
not very closely spaced.

The zoarium is fenestrate; there are from 11 to 13 fenestrules, and about 15 to 29
branches, in 10 mm. The branches are thin, 0:27 to 0:35 mm. in width, with a slight
but usually distinct median carina, which bears a single row of small, sharp nodes,
elongated parallel to the length of the branch along their bases; these nodes are spaced
0:27 to 0-5 mm. apart, and there are about 25 in 10 mm. The zooecial apertures are
placed on the rather flattened sides of the branches, and they do not project into the
fenestrules; the apertures are circular, 0:14 mm. in diameter, and are surrounded by
only very slight peristomes; there are about three apertures to a fenestrule, and 31 in
10 mm., the distance between the centres of successive apertures being 0-25 to 0-43 mm.
The fenestrules are rectangular, 0-6 to 0:95 mm. long, and between 0-24 and 0-4 mm.
wide; the width of the dissepiments is from 0-06 to 0:19 mm., and the length of one
fenestrule and one dissepiment is from 0:7 to 1:08 mm.

This species resembles rather closely Fenestrellina dispersa Crockford, 1943, a very
common form in the Permian of New South Wales, Tasmania and Central Queensland;
constant differences in the measurements of the spacing of the nodes and of the zooecia
in these specimens from Lake’s Creek, however, indicate that this form is a distinct
species. Fenestrellina pulchradorsalis (Bassler), 1929, from the Somohole and Bitaoeni
Beds of Timor, is also a very similar species; the spacing of its zooecia is similar, but,
although Bassler mentions the presence of nodes in his description of this form, he gives
no indication of their spacing, and this cannot be determined from the illustrations, so
that complete comparison with this Timor species is not possible.

128 BRYOZOAN FAUNA FROM ROCKHAMPTON,

FENESTRELLINA SPARSINODATA, Nn. Sp. Text-fig. 3.

Holotype: F.7952, Univ. Queensland Colln.

Locality: Trachypora horizon behind Lake’s Creek Quarry, Rockhampton.

Fine Fenestrellina, 3 zooecia to a fenestrule; nodes small and widely spaced on the
carina.

The zoarium is fenestrate; in 10 mm. there are from about 12 to 20 branches, and
10 to 12 fenestrules. The branches are straight, 0-25 to 0-41 mm. wide, and there are
two rows of small, oval zooecial apertures, 0:14 x 0:1 mm. in diameter, separated by a
thin but sharp median carina, on which there is a single row of small, sharp, widely
spaced nodes; the nodes are placed 0:66 to 0:92 mm. apart, and there are about 12 nodes
in 10 mm. The apertures are surrounded by slight peristomes; the distance between the
centres of successive apertures is 0:25 to 0-4 mm., and there are 32 apertures in 10 mm.,
with usually 3, rarely 4, to a fenestrule. The fenestrules are oval, 0:6 to 1:03 mm. long
and 0:13 to 0:36 mm. wide; the dissepiments are 0-11 to 0:33 mm. wide, and the length of
one fenestrule and one dissepiment is from 0:82 to 1:22 mm. The reverse surface of both
branches and dissepiments is smooth and evenly rounded, and they are both of about
the same thickness.

Fenestrellina disjecta Crockford, 1944, from the Wandagee and Nooncanbah Series
of Western Australia, is similar in its general appearance to this species, but its apertures
are more closely, and its nodes more widely, spaced.

FENESTRELLINA ROCKHAMPTONENSIS, nh. sp. Text-fig. 10.

Holotype: F.7953A, Univ. Queensland Colln.

Locality: Lake’s Creek Quarry, Rockhampton.

Coarse Fenestrellina, 4 zooecia to a fenestrule; carina broad and indistinct; nodes
large, distant, and irregularly spaced.

The zoarium is fenestrate; there are 11 to 14 branches horizontally, and 6 to 9
fenestrules vertically, in 10 mm. The branches are broad, 0°33 to 0-54 mm. wide; there
are two rows of zooecial apertures, separated by a broad, indistinct carina, on which
there is a single row of very large, irregularly spaced nodes; the distance between
successive nodes is from 0-47 to 1:85 mm., and on an average there are about 10 nodes
in 10 mm. These nodes are up to 0°45 x 0:29 mm. in diameter at the base. The surface
of the carina and between the apertures is ornamented by rows of tiny granules and by
discontinuous grooves. The zooecial apertures are rounded, 0-2 mm. in diameter, and
they are placed on the slightly sloping sides of the branches. No peristomes are
developed; the distance between the centres of successive apertures is 0:25 to 0-46 mm.,
and the average number of zooecia in 10 mm. is 28. The fenestrules are oval, 0-79 to
1-5 mm. long and 0-4 to 0:79 mm. wide; the width of the dissepiments is 0:08 to 0:38 mm.,
the total length of one fenestrule and one dissepiment being from 1:08 to 1-76 mm. On
the reverse surface both branches and dissepiments appear to be smooth and evenly
rounded, the dissepiments being less thick than the branches.

The closer spacing of the zooecia, and the irregular and more distant spacing of the
nodes, distinguish this species from Fenestrellina granulifera.

FENESTRELLINA SPARSA, Nn. Sp. Text-fig. 7.

Holotype: F.7954, Univ. Queensland Colln.

Locality: Trachypora horizon behind Lake’s Creek Quarry, Rockhampton.

Coarse Fenestrellina, branches sinuous, 4 to 6 zooecia to a fenestrule; nodes distant
and irregularly spaced, but large.

There are 6 to 12 branches, and from slightly less than 5 to 6 fenestrules, in 10 mm.
The branches are typically not straight, but rather flexuous; they are 0-44 to 0-67 mm.
in width. Between the two rows of apertures there is only a very slight median carina,
on which there is a single row of large, widely spaced nodes, whose bases are elongated
parallel to the length of the branches. The distance between the nodes is from 1:14 to
1:82 mm.; there are about 7 nodes in 10 mm. The apertures are rounded, 0-16 mm. in
diameter, and are surrounded by very thin, slight peristomes; there are 4 to 6 apertures
to a fenestrule, the distance between the centres of successive apertures being 0-3 to

BY JOAN CROCKFORD. 129

0-51 mm.; there are 25 apertures in 10 mm. Increase to three rows of zooecia occurs
only immediately before branching. The fenestrules are 1:36 to 2:57 mm. long, and
from 0-4 to 0:95 mm. wide; the width of the dissepiments is from 0-17 to 0-55 mm., and
the length of one fenestrule and one dissepiment from 1:54 to 2:93 mm. On the reverse
surface both branches and dissepiments are smooth and evenly rounded, and they are
of about the same thickness. The zooecia themselves are rather triangular in outline.

Text-fig. 7.—Fenestrellina sparsa, n. sp. Celluliferous surface of the holotype, x 10.
Text-fig. §.—Fenestrellina spinifera, n. sp. Celluliferous surface of the holotype, x 10.

FENESTRELLINA SPINIFERA, Nn. Sp. Text-fig. 8.

Holotype: F.7955A, Univ. Queensland Colln.

Locality: Trachypora horizon behind Lake’s Creek Quarry, Rockhampton.

Coarse Fenestrellina, 3 to 6 zooecia to a fenestrule; slight carina with small,
comparatively closely spaced nodes.

There are 9 to 16 branches, and 5 to 6-5 fenestrules, in 10 mm. The branches are
straight to slightly sinuous, 0-33 to 0-65 mm. wide, and there is a slight, rounded median
carina, which bears a single row of small, rather closely spaced nodes; these nodes are
0-43 to 0:59 mm. apart, and there are 20 in 10 mm. There are two rows of zooecial
apertures; 3 to 6 apertures occur to a fenestrule, and 24 in 10 mm., the distance between
the centres of successive apertures being 0-32 to 0-55 mm.; the apertures are rounded,
0-13 mm. in diameter, and they are surrounded by thin, slight peristomes. The
fenestrules are from 1-0 to 2:12 mm. long, and 0:14 to 0-92 mm. wide; the width of the
dissepiments is 0:12 to 0-48 mm., and the length of one fenestrule and one dissepiment
from 1:44 to 2:35 mm. The reverse surface of both the branches and the dissepiments
are evenly rounded, the branches being thicker than the dissepiments; the reverse of
the branches was apparently ornamented by coarse granules, so far as is shown in
the specimens available.

FENESTRELLINA EXPANSA, nh. Sp. Text-fig. 11.
._ Holotype: F.7956A, Univ. Queensland Colln.
Locality: Lake’s Creek Quarry, Rockhampton (holotype); Yrachypora horizon
behind Lake’s Creek Quarry (F.7957, Univ. Queensland Colln.).
Very coarse Fenestrellina, 4 to 10 zooecia to a fenestrule; carina high, rounded, and
unthout nodes.

130 BRYOZOAN FAUNA FROM ROCKHAMPTON,

Text-fig. 9.—Polypora minuta, n. sp. Celluliferous surface of the holotype, x 10.
Text-fig. 10.—Fenestrellina rockhamptonensis, n. sp. Celluliferous surface of the holotype,

Text-fig. 11.—Fenestrellina expansa, n. sp. Celluliferous surface of the holotype, x 10.
Text-fig. 12.—Fenestrellina expansa var. nodulifera, n. var. Celluliferous surface of the
holotype, x 10. y

BY JOAN CROCKEORD. 131

The zoarium is fenestrate, and is very coarse-meshed, there being 6 to 7:5 branches,
and only 2 to 3:5 fenestrules, in 10 mm. The branches are flexuous, and they bifurcate
very frequently, usually within three fenestrules; there are two rows of zooecial aper-
tures, increasing to three only immediately before bifurcation; there is a broad, rounded,
and rather high carina along the centre of the branches, but no nodes appear to be
developed on it. The apertures are placed on the slightly flattened sides of the branches;
they are large and round, about 0:24 mm. in diameter, and are surrounded by slight
peristomes. The celluliferous surface between the apertures was ornamented by fine
discontinuous ridges and grooves. There are from 4 to 10 apertures to a fenestrule, and
about 21 in 10 mm., the distance between the centres of successive apertures being
0-38 to 0-58 mm. The fenestrules are irregularly rectangular; they are from 1:1 to
5:3 mm. long and from 0:4 to 1-45 mm. wide; the width of the dissepiments is from
0-5 to 1:0 mm. On the reverse surface both branches and dissepiments were evenly
rounded and apparently without ornamentation; the branches were much thicker than
the dissepiments.

This species resembles Fenestrellina regina (Bassler), 1929, from the Amarassi (7?)
Beds of Timor; the irregularity in the length and shape of the fenestrules in these
Queensland specimens is, however, different from the habit of the zoarium shown in
Bassler’s figures of F’. regina, and the spacing of the zooecia in F. regina (about 27 to
28 in 10 mm.) is much closer than in F. expansa. Another species which shows
resemblances to this form is F. chapmani Crockford, 1944, from the Callytharra Series
(Permian) of Western Australia; F. chapmani, however, has only a slight carina, with
very large and prominent nodes, absent in F. exrpansa.

FENESTRELLINA EXPANSA Val. NODULIFERA, Nn. var. Text-fig. 12.

Holotype: F.7958A, Univ. Queensland Colln.

Locality: Trachypora horizon behind Lake’s Creek Quarry, Rockhampton.

Very coarse Fenestrellina, 6 to 8 zooecia to a fenestrule; carina slight, with large
nodes developed at very distant but fairly regular intervals.

There are about nine branches, and three fenestrules, in 10 mm. The branches are
straight, 0-55 to 0-71 mm. long, and bear two rows of zooecial apertures separated by a
slight, median carina, on which very large nodes are developed; these nodes are placed
at distant but usually fairly regular intervals, generally being between 3-1 and 3-5 mm.
apart. The apertures are oval, about 0:3 x 0:17 mm. in diameter; there are 6 to 8
apertures to a fenestrule, and 21 in 10 mm., the distance between the centres of
successive apertures being 0-38 to 0-57 mm. The fenestrules are normally from 2-1 to
3:7 mm. long, although abnormal fenestrules of small size do occur, and they are
0-15 to 0-87 mm. wide; the width of the dissepiments is 0-19 to 0-84 mm., the length
of one fenestrule and one dissepiment being 2:98 to 4-2 mm.

This form is so similar to F. expansa, except in the occurrence of large nodes,
that it seems best to consider it a variety of the more abundant species. It is
distinguished from F. chapmani by the very different spacing of its nodes.

FENESTRELLINA ASPRATILIS (Bassler), 1929. Text-fig. 5.

Fenestella aspratilis Bassler, 1929, 76, Pl. cexli (17), figs. 14-17; Fenestella spinulosa
Condra ?, Girty, 1908, 137, Pl. xix, fig. 4: Fenestrellina girtyi Elias, 1937, 314, fig. 1.

In the specimens from Lake’s Creek Quarry (F.7950B, F.7959, F.7960A, F.7961A, B,
Univ. Queensland Colln.), there are 16 to 20 branches, and 14 to 17 fenestrules, in 10 mm.
The branches are straight, 0-24 to 0:32 mm. wide; there is a slight median carina, on
which there is a single row of sharp, high nodes, elongated along their bases; these
nodes are spaced from 0-25 to 0:38 mm. apart, and there are about 31 in 10 mm. There
are two rows of small, circular zooecial apertures, 0:1 mm. in diameter, surrounded by
slight peristomes; the distance between the centres of successive apertures is from
0-25 to 0-38 mm., and there are 32 apertures in 10 mm. The apertures are frequently
placed so that one is opposite the end of each dissepiment and one at the centre of each
fenestrule, and here the fenestrules are hour-glass shaped, but the arrangement of the
apertures is not always regular. The fenestrules are 0-47 to 0-71 mm. long and 0-19 to

132 BRYOZOAN FAUNA FROM ROCKHAMPTON,

0-43 mm. wide, and the width of the dissepiments is from 0-08 to 0:19 mm.; the length
of one fenestrule and one dissepiment is from 0-55 to 0-87 mm. On the reverse surface
the branches are rather thicker than the dissepiments; both are evenly rounded, and
in places the backs of the branches appear to have been coarsely granular. The full
range of variation in measurements is usually shown in the one specimen.

, Elias (1937, 314) discussed at some length the characters of Fenestella spinulosa
Condra ?, Girty, which he named Fenestrellina girtyi Elias, and he pointed out its
affinities to the Fenestrellina mimica group of fenestellids; it appears from the measure-
ments given by him and by Girty, and from additional measurements taken on Girty’s
diagrams, that this species from the ‘‘Dark Limestone” of the Guadalupian Group in the
Guadalupe Mts. of Texas, is the same as F.. aspratilis (Bassler), from the Bitaoeni and
Basleo Beds of Timor; the measurements of specimens from these two areas and from
Lake’s Creek are compared in Table 1. Girty’s specimens evidently did not show the
nodes ornamenting the reverse surface described by Bassler, and this ornamentation is
only doubtfully shown in the Lake’s Creek specimens; this, however, is not sufficient
basis for the separation of these specimens into different species, as, when the reverse
surface is weathered, these coarse granules readily disappear, and they do not show the
scars shown when the nodes along the carina have been weathered away; in other
species in which coarse granules are developed on the reverse surface, they are
frequently not at all constantly developed, even over the surface of the one specimen.

FENESTRELLINA GRANULIFERA Crockford, 1941. Text-fig. 4.
Fenestrellina granulifera Crockford, 1941, 509, Pl. xxi, fig. 4; 1943, 266.

This species is one of the commonest forms in material from Lake’s Creek Quarry
(specimens F.7950C-—H, F.7960A, F.7962—-5, F.7966A—D, F.7967—8, Univ. Queensland Colln.),
and occurs also in material from the Trachypora horizon behind the same Quarry
(F.7969, Univ. Queensland Colln.). The characters shown by these specimens corres-
pond very closely with those of the holotype; the range of variation is rather greater
in the Queensland specimens, as would be expected from the larger number of specimens
available for study. In these specimens there are from 6 to 9 fenestrules, and 12 to 16
branches, in 10 mm.; the branches, which are 0-28 to 0°52 mm. wide, have a slight
median carina, which bears a single row of large, blunt nodes, placed 0:4 to 0:73 mm.
apart, with 18 in 10 mm. The zooecial apertures are rather large, about 0-2 mm. in
diameter, and are not very closely spaced, there being about 25 in 10 mm., and usually
4 to a fenestrule; the distance between the centres of successive apertures is from
0-3 to 0-55 mm. The fenestrules are 0-8 to 1:76 mm. long, and from 0-13 to 0:92 mm.
wide; the width of the dissepiments is from 0-08 to 0-4 mm., and the length of one
fenestrule and one dissepiment usually between 1:0 and 1-9 mm.

FENESTRELLINA CANTHARIFORMIS Crockford, 1941. Text-fig. 1.
Fenestreilina canthariformis Crockford, 1941, 510, Text-fig. 2a.

Three specimens (F.7970—2, Univ. Queensland Colln.) amongst the material from the
Trachypora horizon behind Lake’s Creek Quarry agree in their characters with the
holotype of this species; one of these specimens, showing the characteristic very closely
spaced nodes and apertures, is figured in Text-fig. 1.

Genus Mininya Crockford, 1944.
MINILYA DUPLARIS Crockford, 1944. Text-fig. 2.

Minilya duplaris Crockford, 1944, 173, Pl. 1, figs. 5, 7; Text-fig. 1, C, D.

One specimen of this species was found in material from the Trachypora horizon
behind Lake’s Creek Quarry (F.7973, Univ. Queensland Colln.), and a second in material
from Lake’s Creek Quarry (No. 55 in Reid’s Collection of fossils in the Univ. Queensland
Colln.). Although the preservation of these specimens is indifferent, they are charac-
teristic specimens of this species, very common in the Permian of Western Australia.
In addition to occurring at this locality, M. duplaris also occurs—associated with
Fenestrellina horologia (Bretnall), another common Western Australian species, and a

BY JOAN CROCKFORD. 118334

number of other fenestrate Bryozoa—at Consuelo Creek, 2 miles above Cattle Creek,
Springsure district, Queensland.

Genus Potyrora McCoy, 1845.
POLYPORA MINUTA, n. Sp. Text-fig. 9.

Holotype: F.7974A, Univ. Queensland Colln.

Locality: Trachypora horizon behind Lake’s Creek Quarry (holotype); Lake’s Creek
Quarry (F.7975, Univ. Queensland Colln.).

Fine Polypora, zooecia in 3 rows, with 3 zooecia in each row to a fenestrule; surface
ornamented by discontinuous ridges and grooves between the apertures, and by a few
small nodes.

There are 11 to 12 branches, and 10 to 12 fenestrules, in 10 mm. he branches are
narrow, 0:43 to 0-52 mm. wide; normally there are three rows of zooecial apertures, with
two just after and four just before bifurcation. The apertures are slightly oval, about
0-17 x 0-14 mm. in diameter; they are surrounded by very thin, high peristomes, and are
frequently closed by a centrally perforated calcareous plate. There are 3 apertures to
a fenestrule, and 28 to 29 in 10 mm., the distance between the centres of successive
apertures being 0-28 to 0-43 mm. The fenestrules are oval, 0-63 to 1:0 mm. long and
0:3 to 0-55 mm. wide; the width of the dissepiments is 0-1 to 0-2 mm., and the length of
one fenestrule and one dissepiment 0-82 to 1:13 mm. On the reverse surface the branches
are flattened, and the dissepiments, which are much thinner, are rounded. The casts
showed the longitudinal striae shown on the worn reverse surface of most fenestellids,
but were not well enough preserved to show if any other ornamentation occurred.

PoLypora virGa Laseron, 1918.

Polypora virga Laseron, 1918, 192, Pl. vii, fig. 4, Pl. viii, fig. 2; Polypora virga
Laseron, Crockford, 1941, 410, Pl. xix, fig. 3.

A single specimen (F.7976, Univ. Queensland Colln.), from Lake’s Creek Quarry,
is referred to this species, which is a common form in the Upper Marine Series of New
South Wales, and which occurs also in the Lower Marine Series in New South Wales,
and in the Permian at Marlborough, Tasmania. In its measurements and general
appearance this specimen from Queensland compares closely with the neotype.

POLYPORA woopst (Etheridge), 1892.

Protoretepora ampla (Lonsdale), de Koninck, 1878, 42, t. 8, figs. 5, a-c; [non]
Fenestella ampla Lonsdale, 1844, 163; Protoretepora sp. indet. Etheridge, 1880, 16;
Protoretepora ampla var. woodsi Etheridge, 1892, 222, Pl. 8, fig. 12; Polypora tumula
Laseron, 1918, 191, Pl. vii, fig. 3, Pl. ix; Polypora tripliseriata Bassler, 1929, 79, Pl. cexlii
(18), figs. 14-16.

Two typical specimens (F.7953B, C, Univ. Queensland Colln.) of this common species
occur in material from Lake’s Creek Quarry.

SUMMARY.

Fourteen species and varieties of Fenestella and Polypora occurring in Permian
strata in and near the Lake’s Creek Quarry in the Rockhampton district of Queensland
are described. Of these forms, six are species previously described from Lower Permian
rocks in Hastern or Western Australia or in Timor, five are described as new species
and one as a new variety. Bryozoa have previously been recorded from this Quarry,
and their occurrence has been quoted in support of correlations between the Lake’s Creek
Quarry beds and the Neerkol Series of Upper Carboniferous age; it is here shown that
such a correlation based on the Bryozoa is untenable, the age of the Lake’s Creek beds
being Lower Permian.

BIBLIOGRAPHY.
BASSuLER, R. S., 1929.—The Permian Bryozoa of Timor. Paldontologie von Timor, xvi Lief.,
XXvViii.
BRYAN, W. H., and Jonps, O. A., 1944.—A Revised Glossary of Queensland Stratigraphy. Univ.
Qd., Pap. Dept. Geol.. n.s., 2 (11).

P

134 BRYOZOAN FAUNA FROM ROCKHAMPTON,

CROCKFORD, JOAN, 1941a.—Permian Bryozoa from Eastern Australia, Part i: A Revision of Some
Previously-named Species of Fenestrellinidae (Fenestellidae). J. Roy. Soc. N.S.W., 74: 397.

——_—_— , 1941b.—Permian Bryozoa from Hastern Australia, Part ii: New Species from the
Upper Marine Series of New South Wales. Ibid., 74: 502.

, 1943.—Permian Bryozoa of Eastern Australia, Part iii: Batostomellidae and Fenestrel-

linidae from Queensland, New South Wales, and Tasmania. Ibid., 76: 258.

, 1944a.—Bryozoa from the Wandagee and Nooncanbah Series (Permian) of Western

Australia, Part i. J. Roy. Soc. W. Auwst., 28: 165.

, 1944b.—A Revision of some Previously Described Species of Bryozoa from the Upper
Palaeozoic of Western Australia. Ibid., 28: 187.

————., 1944c.— Bryozoa from the Permian of Western Australia. Part i. Cyclostomata and
Cryptostomata from the North-West Basin and Kimberley District. Proc. LINN. Soc.
N.S.W., 69 (3-4) : 139. ;

Evias, M. K., 1937.—Stratigraphic Significance of Some Late Paleozoic Fenestrate Bryozoans.
J. Paleont., 11 (4): 306.

ETHERIDGE, R., snr., 1872.—Palaeozoic and Mesozoic Fossils from Queensland. Quart. J. Geol.
Soce% 28 shi.

ETHERIDGE, R., jnr., 1880.—On a Collection of Fossils from the Bowen R. Coalfield and the
Limestone of the Fanning R., North Queensland. Proc. Phys. Soc. Edinb., 5: 1.

—————,, 1892.—-In Jack, R. L., and Etheridge, R., jnr., Geology and Palaeontology of Queens-
land and New Guinea. Qd. Geol., Surv., Publ. No. 92.

Girty, G. H., 1908.—Guadalupian Fauna. U.S. Geol. Surv., Prof. Pap. 58.

KoNINCK, L. G. de, 1877.—Recherches sur les fossiles paléozoiques de la Nouvelle Galle du Sud.
Mém. Soc. Sci. Liége, (2) 8: 1. Reprinted in English in Mem. Geol. Surv. N.S.W.,
Palaeont., 6.

LASERON, C. F., 1918.—Notes on Some Permo-Carboniferous Fenestellidae, with Descriptions of
.New Species. J. Roy. Soc. N.S.W., 52: 181.

LONSDALE, W., 1844.—In Darwin, C., Geological Observations on Volcanic Islands.

MerEK, F. B., 1872.—In Hayden’s Final Report on the Geological Survey of Nebraska, etc.,
Part 2. Paleontology.

ORBIGNY, A. d’, 1849.—Sur Quelques Genres Nouveaux de Mollusques Bryozoaires. Rev. Mag.
Zool.) hy 499. a

Reip, J. H., 1928.—Central Queensland Geological Section. Qd. Govt. Min. J., 29: 384.

, 1936.—The Queensland Upper Palaeozoic Succession. Qd. Geol. Surv. Publ. No. 278.
WAAGEN, W., and PICHL, J., 1887.—Salt Range Fossils. Palaeontologica Indica, (13) I.
WHITEHOUSE, F. W., 1928.—Central Queensland Geology. Qd. Govt. Min. J.,.29: 441.

, 1929.—In Bryan, W. H., Report of Carboniferous and Permo-Carboniferous Correlation

Committee. Rep. Aust. Ass. Adv. Sci., Hobart, 1928, 19: 74.

, 1930.—Report on a Collection of Fossils made by Mr. J. H. Reid, in the Springsure

District. Qd. Govt. Min. J., 31: 156.

135

MISCELLANEOUS NOTES ON AUSTRALIAN DIPTERA. XI.
EVOLUTION OF CHARACTERS IN THE ORDER: VENATION OF THE NEMESTRINIDAE.
By G. H. Harpy, Queensland University, Brisbane.

(Two Text-figures. )

[Read 25th July, 1945.]

Foreword.—The direction which the evolutionary trends in Diptera is presumed to
take, is given by Williston (1908), his main idea being developed below, as on present
evidence it remains satisfactory, though published thirty-seven years ago. Since Williston
wrote, however, doubt has arisen as to whether the terms applied to the structures of
the Diptera are in accord with homologies of other orders, and amendments are being
suggested by various morphologists to rectify the position. These amendments are
advancing the formation of a phylogenetical classification and, in addition, it is now
possible to render into simple sequence some complex and apparently interrelated minor
developments that are used in taxonomic work.

Regarding the present study of venation in the Nemestrinidae,* it must be noted that
considerable confusion exists in the interpretation of main veins and cross-veins, no
method being known to determine their status other than by comparative morphology.
Alexander (1928) has shown that the so-called cross-vein between R, and R, is the vein
R, itself coalescing with R, He gives a supernumerary vein between R, and R; which
was originally regarded as being part of R, at its fureation with R,.

This development suggests that the second radial-median cross-vein in the Cyrtidae
may be the vein M, coalescing with R,; and the supposed M, coalescing with M, is due to
a cross-vein, and is not part of a main vein. In the ultimate, it is expected that the veins
holding the same position in the wings of various families are not all homologues, but it
is customary to retain the Tillyard notation until valid reasons are given to make a
change.

I am indebted to Mr. Lucius Allen, B.A., for abbreviating some of the paragraphs
of my manuscript, and to Professor EH. F. Simmonds for mathematical aid when deter-
mining the sequence of shapes evolved in the varied abdominal forms, a summary of
which is given below. Also Mr. Edgar Riek, B.Sc., recently rediscovered the wing-
folding mimetic fly at Sunnybank (September, 1944) and studied it with me on a
comparative basis with its model, thus enabling the account below to be more detailed
than otherwise would have been the case.

The Longitudinally Folding Wings.—The flies Cerioides subarmata C. & B. and
C. alaplicata Hardy (Syrphidae) have wings that fold longitudinally in the manner

* The notation of veins given by Tillyard does not conform to homologies of veins within
the order. So far Alexander has shown that in the Nemestrinidae there is one link with a branch
of the Tipuloidea, so presumably this is a connection leading to the Tabanoidea. In the Cyrtidae
there are two radial-median cross-veins which is the number also traced in the Syrphoidea and
may prove another link between two superfamilies if the morphological affinity between these
two families is correctly assessed by Crampton. Clues like these are rare and hard to identify,
and each author placed his own valuation on their significance. In Tillyard’s view there can
hardly be two radial-median cross-veins, and in manuscript he notated the second in the
Cyrtidae as being M, which, if correct, suggests that M, is eliminated from the veins of most
Syrphidae. This and other consequences may make Tillyard’s notation for the Muscoidea utterly
collapse. There can be no scheme of venation yet acceptable to cover the Diptera, but the
Tillyard scheme already is modified for the Tipuloidea along phylogenetical developments, and,
in its constituent parts, the radial field is notated differently one from the other, agreeing with
the Tillyard notation only in the Tanyderidae.

136 MISCELLANEOUS NOTES ON AUSTRALIAN DIPTERA. XI,

somewhat resembling the folding wings in Diploptera, thus enhancing the fly’s
resemblance to those wasps (Hardy, 1933, p. 420). The flexure in both cases is evidently
automatic as it occurs only when the wing is brought back to the resting position and,
at least on the wasp, no musculature of the forewing can be involved; in other respects
there are marked differences in the manner of this flexure.

In the wasp, starting from the apex of the line, the flexure usually crosses the tip
of the median vein, but sometimes lies behind it, and reaches the posterior wing border
near the base, passing through the area between the median and cubital veins, and
crosses the cubital vein where this is slightly flattened and broadened. The folding
takes place along this line in a manner that brings the posterior part of the wing to
lie below the anterior part, venter to venter.

In the fly the line of flexure runs from the wing margin between the median and
radial veins, crosses the media apically where it bends rearwards and without affecting
that vein, then follows the strong crease adjacent to the media through the vein M,,,
which is strongly depressed at this point of furcation and perhaps completely broken
there; the flexure coincides with this crease to reach a considerable distance along it to
a point from where it passes over the fold just anteriorly placed to the vein bordering
the alula along which the flexure reaches the wing margin behind the wing base. Those
wing sclerites lying at the base, one each side of the median crease, anteriorly and
posteriorly, appear to move towards each other when the wing is folded in a manner
that brings the posterior part of the wing to lie above, nearly touching, the anterior
part, meeting dorsum to dorsum. It is not certain if this apparent motion of the wing
sclerites be due to the motion of the folding only, as it might be due in part to some
undetected muscular action perhaps causing the flexure.

That break at the furcation of the median vein may yet be detected more widely
than is at present known, for it occurs in the genus Apiocera (see Norris, 1936, p. 54),
and is known as the thyridium on other extant orders and in some fossil wings. This
thyridium may have some connection with wing folding wherever it occurs and thus
indicate that once wing folding was more widespread. Further evidence may be
indicated by wing-folded pupae of the families Blepharoceridae and Simulidae, leaving
fold-creases in the adult which no longer folds the wings.

Those flies that have an upward flexure of the alula when the wings are brought
to rest show that presumably the jugal fold is all that remains in these cases of what
may have been a more complete folding system, and other flies are showing indications
of wing folding in other ways.

The Transversely Bending Wings.—Nearly all other flies known to me, including
other Cerioides, retain the wings expanded in the one plane apart from the alula
already mentioned. The few exceptions have wings that bend transversely when at
rest and do not complete the folding. This transverse flexure is found in the Psychodidae,
in the genus Ophiodesma (Stratiomyiidae), in certain flies of the Acalyptrata and in the
genus Chlororhina (Calliphoridae), thus showing a wide distribution over the order
with the maximum bending in the Psychodidae where the wings have a downward
flexture with an acute angle near the base, and in the others forming an obtuse angle
at or beyond half-wing length. In the Blepharoceridae there is a transverse crease
marking a line of folding that is comparable to this line of flexure in the Brachycera
and which may have had the same origin.

The Vena Spuria.—Possibly that convex vena spuria found in most Syrphidae is
another survival of a primitive character, marking a remnant of some former condition
in so far as creases in other families strongly suggest that these once had a vena spuria,
too. Lundbeck (1916, p. 33) states that “. . . in Bombylius we find a wing-fold much
recalling the vena spuria’”’, and although there are many similar traces elsewhere, this
character is missing in Australian species of Bombylius.

The claim that the vena spuria is a spurious vein rests upon the fact that it cannot
be brought into alignment with any notation of veins, but all notations are unsatisfactory,
exhibiting many abnormalities which are not explained.

The Colours, Hairs and Processes—The eye-colour and the body-colour, already
discussed in these ProcrEpINGs (Hardy, 19389 and 1940 respectively), may sometimes

BY G. H. HARDY. 137

depend upon a fundamental alteration like that which brings variegated eye-colouration
and the development of a powdery covering which produces a new range of colours on
the body, but the nature of these developments is not understood. The change that
takes place in the terminal structures (Hardy, 1944) and the development of bristles,
tubercles, spines, etc., are features that appear to become increasingly complex and then
simplify, finally disappearing, leaving hardly a trace of the phenomenon in the advanced
stages.

Spines that arise from the cuticle as local protuberances must not be confused
with those granulations that may cover wide areas and are formed by deepening hair-
pits. A highly roughened surface exhibited in the latter case seems to be associated with
a reduction of hair length, and in addition, the more hirsute types of flies, wherever
they occur, lie low in the phylogeny of their respective branches. This does not mean
that the most primitive flies were densely hairy, but rather that an early increase in
hair density may have been linked with the disappearance of something else.

Two Types of Spine.—Thoracic spines are found in various but isolated genera and
also in all genera and species of the tribe Chrysopogonini (illustrated by Hardy, 1934,
fig. 15). These spines have a consistency in size and shape that suggests a considerable
stability not equalled by any scutellar spines formed by cuticular projections. It may
be true that excrescences of many kinds arise on insects of senile strain, as reported by
James (19386), but the cause of cuticular spines, wherever they occur, seems to be the
reduction of surface area covered by the sclerite without a corresponding reduction of
the volume of the sclerite, mainly that of the scutellum, whereas the development of
the pair of spines referred to on the thorax, seems to be linked with the elimination of
vestiture, since each of these two hairs is transformed first into a strong bristle, then
into a spine, and the rest of the mesonotum is left bare.

Antennal Tubercle.—A trend towards developing an antennal tubercle occurs widely
over the order and is well known in the genus Cerioides (illustrated by Nicholson, 1927,
Text-figs. 1, C and D), where the tubercle is seen to vary from short to excessively long.
This development on the fly takes place subsequent to emergence from the puparium
and apparently it originated within the subfamily Cerioidinae, not as an inheritance from
some more primitive type. By holding the point of a pin against the head at the base
of the antennae of a newly emerged fly, it is easy to restrain development of the
tubercle, and this action produces no abnormality on the head, nor yet any alteration
to the antennae, thus showing that the bulk that would have formed the tubercle has
been absorbed within the head mass. Such experiments were carried out on a number
of C. ornata Saund. reared from waste comb of a beehive; the tubercle also was stunted
in its growth by similar experiments at later periods. Occasionally flies with a shortened
tubercle are caught, the full extension of the tubercle evidently having been prevented
by some extraneous circumstance.

The Articulating Scutellum.—lit is difficult to interpret the case of the articulating
scutellum (described by Hardy, 1944, p. 61) in Neoexaireta spinigera Wied. (Stratio-
myiidae) one of the so-called mimetic flies with pictured wings and clubbed abdomen,
both evidently rather primitive characters. Here the scutellum is spined and is hinged
to swing up and down in unison with the motion of the wings, whilst (at least on the
relaxed fly) the halteres move in the opposite direction. If this be a true scutellum,
then a unique character has arisen without any apparent utility. If, however, it be
the true metanotum, as has been claimed, then it becomes obvious that the muscles of
articulation survive from those of the metanotum. A claim may yet be made that it is
the true first tergite of the abdomen.

The Upstanding Scutellum.—tThis articulation of the scutellum may account for the
various angles at which the scutellum is set within the Stratiomyiidae, notably in the
tribe Pachygasterini, where some species are found with the upstanding scutellum as
opposed to the normal horizontally fixed form.

As opposed to this true type of upstanding scutellum is another found in some
Therevidae (illustrated by Mann, 1929, fig. 2, D), which is caused by an enlarged tubercle
arising from the dorsal surface of the scutellum which itself is horizontally held as seen
by the marginal bristles lying on the true apical margin below the tubercle. This

138 MISCELLANEOUS NOTES ON AUSTRALIAN DIPTERA. XI,

upward thrust of the dorsal surface could be caused by shrinkage in the length of the
scutellum without sufficient reduction in bulk.

Abdominal Shape.—The shape of the abdomen is fairly consistent within a genus,
few genera showing any marked variation amongst the species. Such variation represents
slightly different stages in the evolutionary process and is not sufficient evidence to
indicate a complete sequence of changes. It is convenient to assume that the primitive
shape was cylindrical with most segments as long as, or a little longer than, they were
broad. The advance can then be conceived as taking two trends in conformity with
increasing egg-capacity. The abdomen becomes either (a) attenuated with each segment
over three times longer than broad, or (b) broadened so that each segment becomes
over three times broader than long, both these proportions being within those observed
on flies.

When these two trends appear in different segments of the abdomen of one individual,
the two possibilities are realized, viz., the tapering abdomen that widens basally (c),
and the clubbed abdomen that widens apically (d). In the former case, that is in (c),
there develop the shortened basal segments (e), and the highly reduced apical segments
(f) (the so-called ‘“postabdomen”), whilst from the combined (e) and (f) there evolves
the sphero-conical abdomen (g), the form seen in most Muscoidea.

Reduced to the simple components (a) and (0), it is conceivable that many shapes
can be brought about by adjusting length, breadth and depth of the abdomen without
altering the total surface area and capacity of the abdomen, but if one alone were used,
then (a@) would result in a reduction in volume or capacity of the abdomen relative to
surface area, and (b), becoming spheroid under practical conditions, would increase the
relative capacity as can be seen mathematically through the well-known theorem in the
calculus of variations which states that for any given surface area, the volume is
greatest when the surface is a sphere. There is a marked trend to produce the sphero-
conical abdomen throughout the order, and the character even occurs in flies carrying
what may be regarded as primitive characters.

Amongst the long series of shapes derived from the cylindrical form is that in
which the abdomen is broadened in the middle segments. In this type, because of the
necessity to maintain a reasonable capacity, the abdomen becomes depressed and assumes
the depressed oval form, and through further shortening finally reaches the disc shape
as found in many Stratiomyiidae. Further, it is easy to trace the development from the
cylindrical to the conical and to the sphero-conical shapes, and also the trend towards
the clubbed form which is evidently quite an early evolved type as it is based upon a
rather primitive cylindrical abdomen.

Mimetic Pattern.—It is thus seen that abdominal formation, like mimetic colours
discussed by Hardy (1940), is an inevitable process that needs no “enhancement by
natural selection”, as Nicholson (1927) and others have supposed, “in order to satisfy
the phenomenon of mimicry’. If there be any natural selective agency playing its part
in the matter of mimetic Diptera, it would be in maintaining these mimetic characters
which form part of the normal process of development, coming low in the series of
developing characters, and in preventing them from proceeding to more advanced
characters. Environmental factors are maintaining, but did not produce the mimicry.

The Old Theory of Development Revised.—Throughout my investigations on
characters of Diptera, I can find no sign other than that leading to the conclusion that
most characters have a potential distribution over the whole order, the same structures
being induced and eliminated widely, whilst colours are being developed in succession,
each colour in its sequence, following one after the other. James (1936) credibly states
that “In general, there is reason to believe that reduction in parts is a sign of evolu-
tionary development ...”, whilst Williston (1908) forestalled the view in his discussion,
giving nine cases of such reduction, a process which he believed to be irreversible, all
or nearly all being polyphyletic, resulting in numerous cases of parallel resemblances.
Finally he suggested that the Nycteribia and Mallophaga are perhaps the most highly
specialized of all insects, having travelled furthest from the starting point. Some of
Williston’s remarks are not justified in the light of recent information; nevertheless, the
general trend of the idea remains undisputed. Each of those characters he quotes must

BY G. H. HARDY. 139

have developed at the time of building up and isolation of the order and now has reached
its period of elimination, leading one to suppose that reduction in parts is the only
criterion of progress.

The Theory of Compensatory Adaptation.—Some characters and combinations of
characters would seem to have a lethal effect as they develop a stage that does not permit
adaptation to changing environment, and that ultimately must eliminate the line carrying
them. Apart from these, the development of new characters may serve the purpose of
compensation for characters that are lost in the mutual loss and gain taking place
between them. Practically every new structure developed is accompanied by a diminution
of some nearby structure, or expressed simply, a part increases with the diminution of
some other part and vice versa. It becomes increasingly apparent that following on the
loss in adjacent parts, the structures which compensate for the loss may, in a similar
manner, dwindle and disappear at some subsequent stage, the new loss being usually
compensated by further developments. In all cases, however, the total loss exceeds the
gain, making a net loss phylogenetically and reaching simplicity of structure in the
ultimate result.

The whole conception may be rendered clearer if it be assumed that the distin-
guishing characters between species are reflecting only the different stages reached by
each character in the inevitable sequence through which the varied characters are passing.
During the progress of time some of these characters are retained indefinitely whilst
others pass away again relatively soon. The summation of characters forming the
individual species may appear to be a complex, each character diverging “fan-shape” in
the various species from the parent stock, but this is due to the various tracks taken by
them under the influence of the evolutionary process, and actually the drift is towards
a final simplicity to which all tracks lead. Just as there is a line of progress to be
traced arising from the common ancestral form that existed in the remote past, so also
there is a general simplification towards which the ultimately surviving types may drift,
no matter how complex the intervening tracks taken appear to be.

The Application of the Theory of Compensation to Taronomy.—The manner in which
taxonomists are arranging the order Diptera is promoting this idea of compensatory
characters, but in many cases the authors reverse the sequence of development, regarding,
as they do, a simple unspecialized character as primitive, and those forming side issues
and destined to disappear relatively soon, as being advanced or highly specialized
characters, there being no data upon which to check such conclusions. Many of their
views are demonstrably wrong on evidence already accumulating. In the ultimate,
numerous characters are both advanced and primitive, so that, unless qualified, the
use of these terms may mislead. If used as a working hypothesis, the present theory
may lead to better interpretations of characters and more definite conclusions in phylo-
genetical studies.

The Application to Studies in Terminalia.—The trend already seen in the develop-
ment of terminalia, is to transform parts into structures hard to recognize in the new
form, whilst new processes and depressions may arise and become incorporated with the
transformations. Fantastic shapes may develop, these being devoid of any apparent
utility, but some may become adapted to useful purposes though not essential ones. The
main features affecting development in the Muscoidea have been dealt with independently
by Crampton (1936) and by Hardy (1942), this superfamily clearly showing the compli-
cated nature of the subject; but the two authors are not in agreement about details.
Hardy (1934) has shown the development for the Asilidae, this being wholly by
reduction in the female.

The Transposition of Claspers—There is nothing in literature that may suggest how
it comes about that in the Tabanoidea the claspers are hinged to swing laterally and in
the Asiloidea vertically. Quite obviously the Tabanoidea retain the primitive form and
the Asiloidea have developed a secondary pair of claspers, having lost the primary pair
everywhere except, perhaps, in the Bombyliidae. A possible intermediate stage is found
in the Nemestrinidae where, as in the Bombyliidae, the claspers have moved upwards
to lie on the dorsal surface. Whereas in the Nemestrinidae the claspers swing laterally
in the normal way, those in the Bombyliidae swing vertically as seen in other Asiloidea.

140 MISCELLANEOUS NOTES ON AUSTRALIAN DIPTERA. XI,

This may have been brought about in the Bombyliidae, by the supports of the claspers
moving ventrally to form one fused unit on the ventral side of the terminalia, no longer
being laterally placed, and thus displacing the direction of muscular tension. In the
Nemestrinidae, the presence of claspers on the dorsal surface of the supports which
remain laterally placed, does not alter the muscular tension and they move laterally
as ,elsewhere in the Tabanoidea. Besides the claspers, however, there occur what
Mackerras regards with some doubt on its homologies, as being the basi-dististyles. The
pair of basi-dististyles curve upwards and arise from a low position on the clasper
supports, taking the form very much like most claspers in the Asiloidea and may
possibly be homologous with them. These basi-dististyles have a very limited are
in which to swing laterally, and the apices, being near the claspers, restrict the are of
swing of the claspers. There is nothing of this movement suggested by Mackerras but
it was observed on a specimen of Trichophthalma bivittata Westw. (E. Riek, Sunnybank,
August, 1944) given to me for study whilst in the relaxed condition.

There has been a recent tendency by authors to bring the Nemestrinidae and the
Bombyliidae together on the basis of their adult structure, but differences in larval
structure, the female terminalia and other characters separate them into different
superfamilies. The supposed affinity evidently denotes a common ancestry, but mainly
with parallel development playing the leading part.

General Remarks—James (1936), dealing with some evolutionary trends in the
Stratiomyiidae, gives an essential outlook that varies slightly from the present one,
expressing the view that scutellar spines come and go, but he does not carry his
observations to the extent here outlined, and further, there are some modern taxonomic
papers showing agreement in the manner in which the authors adopt phylogenetical
lines of classification and show a very evident progress by reduction of parts, and
comparatively few new structures are evolved in any one branch. The most striking
differences in structure are arising through modifications in size and shape of existing
parts, and some of those that might appear to have arisen de novo are certainly primitive
and largely obsolete. Amongst these rare survivals are three classed as such above,
namely, the folding wings of a mimetic fly, the vena spuria of the Syrphidae and the
articulating scutellum of a Stratiomyiid fly. Hitherto a large part of the broader lines
of classification in the Brachycera has rested upon wing venation, about which very
little is adequately understood (as indicated below) and the homologizing of the veins
within the order is not yet complete. Tillyard considered no difficulty was in the way
of constructing an archetype for the order, but when this was done it was found that
the data gathered, subsequently failed to fall into alignment. It may be better to form
an archetype for each family on an independent basis and work back to that of the order
through the archetype of superfamilies.

VENATION OF THE NEMESTRINIDAE.

The Radial Field—Alexander (1929) has shown that in some extant Nematocera
the radial sector is dichotomously twice branched, which is in agreement with the
generally held theory of the Dipterous venation and is made to cover the whole order
irrespective of exceptions that were becoming apparent. The radial field in some
Nematocera has five distinct branches, but in the ascending seale these become reduced
to four, which is the maximum number found in Brachycera. The first branch of the
radial sector, R. is definitely shown to coalesce with R, and traces of this are left in the
apparent cross-vein joining R, and R;, erroneously called the interradial cross-vein by
Tillyard. This condition is traced in the Brachycera by R. coalescing with R, in the genus
Nycterimyia, and there is no reason apparent to dispute the idea that it was in this
manner that the radial field in the Brachycera became reduced to four veins.

Another feature of importance found by Alexander in the Tipuloidea is the trans-
ferring of the dichotomously twice-branched radial sector to the pectinate type. On this
account the radial sector first furcation divides R, from the main stem and R, divides
away separately; finally R, leaves R., coalescing with R, towards its apex, the evidence
of which is seen in Nycterimyia.

BY G. H. HARDY. 141

The vein R, branching away independently from the main stem and its partial
disruption, forming the “appendix”, is sufficiently wide in distribution amongst the
Tabanoidea and Asiloidea to indicate that this happens in the Brachycera. In this
interpretation of the radial field Alexander assumes that a supernumerary vein occurs
between R, and R; and it is best to regard this as a cross-vein that became important,
permitting the elimination of the basal section of R,, and finally gave the deceptive
appearance of R, branching from R, that misled earlier authors.

This outlook leads to a better interpretation of the radial field in the family
Nemestrinidae and apparent anomalies are thereby explained.

The Median Field.—According to Mackerras (1925), the genus Hazeretoneura
resembles the Leptidae very closely, the wing being primitive in shape as also is the
disposition of M,. If this be so, then on Tillyard’s basis of notation a reasonable inter-
pretation of the median field would be along lines adopted by him for the Leptidae and
Tabanidae, but the manner in which Mackerras treats the venation not only shows
divergence from the Leptid type but also shows a cross-vein between M, and M,. A
cross-vein in this position is unusual but occurs in the Tanyderidae, and its presence
has yet to be shown in any family of the Brachycera. Usually the vein M, takes the
position of this cross-vein by coalescing with M, but it is not certain that this always
occurs, certain primitive genera being possible exceptions. It is usual to regard this
possible cross-vein, wherever it occurs, as being the course of the vein M,, a policy
followed here. The supposedly similar cross-vein in Nycterimorpha is certainly a main
vein, and when so treated it clarifies a very confusing type of venation.

Between the bend of vein M, and the oblique vein in Exeretoneura, occurs the inter-
median cross-vein (i-m) permitting the coalescing of M, and M, to reaeh the wing
margin at a point remote from the oblique vein. When i—m becomes eliminated, it brings
M,,, to lie near the apex of the oblique vein, whilst M., originally at the apex, takes a
position parallel with M, as seen in Trichophthalma and other genera.

A Summary of Wing Characters—The figures (Figs. 1 and 2) represent all the
characters discussed in the radial and median fields of Australian genera, the broken
lines representing the course of veins that have disappeared. In no case has R, shown
a trend to coalesce with either R, or R, which is seen to occur in exotic genera. M, may
coalesce apically with either M. or R,; in both Australian and exotic genera. Very few
supernumerary veins ever occur between the branches of the radial sector and the
median field reaching the hind border of the wing, but are common in some exotic
forms including Nemestrina aegyptica Wied., which has quite a network over the area.
A supernumerary vein is not known to occur between R, and R,.*

Figure 1 represents a hypothetical type of venation showing the manner of notation
of the veins and may even represent the archetype. It is possible, however, that another
cross-vein will be needed to represent the true M, which would then leave that part
of M, adjacent to M, to be regarded as a cross-vein, which it was considered to be by
Mackerras although he did not state his evidence. An appended figure shows the position
of this possibly true M,.

* According to Alexander (these PROCEEDINGS, lii, 1927, 48), in some Tipulidae a super-
numerary vein occurs between R, and R,, whilst a cross-vein that is similarly placed always
takes the oblique form or is in series with the vein branches and is never transversely placed
in the manner of that supernumerary vein. Theoretically the supernumerary veins are the
remnant of the “lace-wing” type of venation of the pre-Dipterous ancestry, and the cross-veins
are the persisting supernumerary veins placed advantageously for maintaining mechanical
strength during the course of vein reduction. Main veins are simple or branched units, but some
so-called branches may be complexes, with a cross-vein formed in series with two remnants of
branches, appearing as a single unit. Any two or more branches join to form a complex, the
maximum development being on Neorhaphiomidas (Apioceratidae) where two of the median
branches conjoin all the radial branches in a common apex. Further complications arise with
partial and total elimination of branches.

As far as I know supernumerary veins are liable to occur only in some Tipulidae, most
Nemestrinidae and occasionally in the Bombyliidae and even in the Asilidae and are interpreted
in accord with the notation adopted just as are the cross-veins.

142 MISCELLANEOUS NOTES ON AUSTRALIAN DIPTERA. XI,

HYPOTHETICAL
SCHEME

CYCLOPSIDEA

Fig. 1.—Venation of the Nemestrinidae.

Se, subcosta; R, radial field, the branches being marked by numerals; R,, radial sector at
its furcation with R,; M, median field with branches marked by numerals; Cu,, cubital vein ;
A, anal vein; i-v, interradial cross-vein; r-m, radial-median cross-vein; m-cw, median-cubital
eross-vein.

The oblique vein crossing the wing at an angle is composed of parts of various veins in
the radial and median fields and is also called the “diagonal vein’. This oblique vein is not
formed in the hypothetical scheme which venation is itself very like that in the Cyrtidae. The
figure shows the position of a second radial-median cross-vein in the Cyrtidae and which may
be the true vein M,, and if so the cross-vein then becomes apparent between M, and M, as
discussed in the text. Separate sketches show some details in the venation of Trichophthalma
and broken lines represent the course of veins that are missing in the various genera.

BY G. H. HARDY.

ATRIADOPS

SS

143

Fig. 2.—Venation of the Nemestrinidae. (For explanation see below ishiress al)

144 MISCELLANEOUS NOTES ON AUSTRALIAN DIPTERA. XI,

The accompanying table of characters gives a summary of twelve points that are
apparently important for phylogenetical consideration and are divided into those of
primary or primitive nature and those of advanced development. These two grades are
complementary but distinct identities.

Seven primary characters are listed and it will be noted that Haxeretoneura and
Nycterimorpha do not have any in common. Between these two genera come Nycterimyia
and Trichopsidea with one, Atriadops and Cyclopsidea with two, and Trichophthalma
with three out of the five carried by Hxeretoneura which, together with Trichophthalma,
bears none of the advanced characters recorded.

In Trichopsidea the vein M, coalesces with M, and there is a trend for M, and M, to
coalesce at their bases (i.e., become stalked). In Atriadops these two veins remain
apart, and the basal coalescing of M, is with Cu, in Nycterimyia and Cyclopsidea, a
condition that leads to the abnormal development in Nycterimorpha. In this last genus
the apical coalescing of M, and M, is far in advance of the oblique vein, and gives the
venation a peculiar character, which Mackerras had difficulty in elucidating.

The various veins are depicted in the figures on the present interpretation and the
distinguishing features there seen are incorporated in the following key which has a
phylogenetical basis. Nycterimyia, Atriadops and the abnormal Nycterimorpha are so
closely akin that the venational differences may prove to be only subgeneric in value.
The related Trichopsidea is parasitic on grasshoppers and it seems probable that the
other three are also parasitic on Acridioidea.

Key to Genera of Australian Nemestrinidae.

i WWathstoum radial) veins reachines the pmarnrsint independently erases tienen ie einieneinioiene 2
With only three radial veins reaching the margin, the vein R, being eliminated or
BOX 1211 Gl baler {0 RRS ue A UR ei fund iar ey Pe ane Ce ER ae a Oe ca a eeO CHEE so oareMENS cas Ona Cos Gin © aiOlO o10"S-6 3

to

With the furcation of R, from the radial sector present and only one median vein lies
parallel with R,. The veins M, and M, remain remote from the oblique vein ........
ig sa hay capleusarre utes ten ead wine Susrwastetye Ge aay Sue tal cA te Mee G scree maa eetnanione Se tee Gat en sateen te ea a Exeretoneura Macq.
With the furcation of R, from the radial sector obliterated. Two median veins lie parallel
with R, and the veins M, and M, are approximate to the apex of the oblique vein ....
sje call dovies wivetinl © elcealoTnyjes ecotis toe, safe e\ weevte tte Strays Gea ae eee Tat SIRE ETE ee cH OU Trichophthalma Westw.
With R, at its furcation from the radial sector, together with the cross-vein i-r forming

ae)

ay SecondarysCross-veing between Evam angen cusctsntare) sit teneney een er Neneh tne ey eee eee 4

Without such secondary cross-vein, Ry and “4-7 being; entirely obliterated i vam) tien 5

4. With M, and M, reaching the wing margin independently .......... Cyclopsidea Mackerras

With M, and M, coalescing before reaching the wing margin .......... Trichopsidea Westw.
Only one free median vein present, M, being absent. R, is present at its furcation from

thesnadial (sectors: Guniques ps 3 skal orcs cy Seen eo Re ee Nycterimyia Lichtw.

5. With veins M, and M, lying parallel with R, and reaching the wing margin independently
SUGE aha eile: sarsel eerehy Sau cch od say otal alee ces re pe eure aches too tes val: Sates ida RUIN LOO 8 vec De yc Sa A Be -... Atriadops Wandol.
With veins M, and R. coalescing apically for a considerable distance, leaving only M,
lying parallel with RM co seuinssef te eNolene suey enenscs ameter aR Aa EST ee Nycterimorpha Lichtw.

TRICHOPHTHALMA BIVITTATA Westw.

Westwood, Philos. Mag., Lond., vi, 1835, 448; Edwards, Dipt. Patagonia and 8. Chile,
v, 1930, 180. TJ. eques Schiner, Reise Novara Dipt., 1868, 110.

This species is the genotype of Trichophthalma, the identity of which species has
been confirmed by Edwards, who examined the type, recording the matter in a footnote.
Schiner’s name, under which the species has long been standing in collections, becomes
a synonym. Care is needed to avoid confusion between the names bivitta Walker and
bivittata Westwood, both species being plentiful in collections and belonging to different
series. Confusion made by earlier authors in this respect has been clarified by Hardy
(1924) and by Mackerras (1925).

Hab.—New South Wales and south-eastern Queensland.

CYCLOPSIDEA HARDYI Mackerras.

Only one specimen is known and was captured by Mr. J. H. Buzacott of the
Queensland Sugar Bureau, and not by myself as recorded erroneously by Mackerras.

BY G. H. HARDY. 145

Owing to an accident before returning to me, only the terminalia are now left, so the
published figure of the venation has been used to interpret the characters.

A Comparative Table of Characters.

Genus. Primary Characters. Advanced Characters.

oblique
coalesce

from

M,

and
apically in advance of

and M, coalesce.
characters in common

characters developed,
with Everetoneura.

occasionally.

remote

vein,

base.

characters retained.

M,
Number of advanced

M, and M, free at apex.
M, and Cu, coalesce at

the oblique vein
Number of primary
Number of primary

i-r present.

M, free at base.

M, and R, coalesce.
M, absent.

i-m present, making M,—,
M,

R, present at furcation.
r-m present at least

4 radial veins.

|

|

|

|

|
on

|
uo

a

|

|

f
aie
fe

Exeretoneura .. |

eS)

Trichophthalma + =— + — = + + | = — = — — 4 —
Cyclopsidea .. | — = | = — = + = i] 1 2

Atriadops oo || Peete — — — ee dk
Nycterimyia .. —- + = ae = = =

Nycterimorpha - = + = = = =

|
{
Trichopsidea a = ate = = = | ea ae “red = Se

TRICHOPSIDEA OESTRACEA Westw.

The following note is taken from the typescript minutes of a meeting of the
Entomological Society of Queensland, held on the 11th July, 1945: “Mr. Sloan submitted
some specimens of Nemestrinid flies and their hosts with the following explanatory
note: The bitter bark, Alstonia constricta, is a small tree which tends to be a pest in
some scrub areas where seedling growth is rapid in established pastures. In some
seasons considerable numbers of these trees die and two Cerambycids, Disterna
plumifera Pase. and Prosoplus woodlarkiana Montr., are sometimes held responsible for
the phenomenon. At Bracewell, in March last, dead trees were common but no live
insects were present in the wood. However, the twigs were studded with the exit holes
normally associated with longicorns of this kind.

On several trees, innumerable flies, each with its wings outstretched, were resting
on the twiggy growth. More than fifty could be counted without any difficulty on a
single tree eight or nine feet high. The flies were more or less torpid and could be
captured ... by simply enclosing them in a glass tube. Each fly had its ovipositor
inserted into one of the longicorn emergence holes and, on sectioning the twigs,
innumerable eggs were found inside each cavity. Each hole must have contained 500
or more eggs of the insect.”

A few of these female flies submitted to me for identification conformed to
T. oestracea Westw., though slightly darker, being perhaps denuded of the slight greyish
pulverulent overlay on the abdomen. It is evident that they share with some Asilidae
the habit of ovipositing into beetle holes though the species has been reared only from
grasshoppers. The specific determination needs confirmation on the males.

NYCTERIMYIA HORNI Lichtwardt.

Apart from the holotype, specimens have not been recorded in literature, and the
species is known to me through a single male specimen in the Department of Agriculture,
Brisbane, captured by Mr. W. B. Barnard, at Toowoomba, on the 10th February, 1927.
The specimen has its terminalia exerted and held conspicuously in the diverted position.

146 MISCELLANEOUS NOTES ON AUSTRALIAN DIPTERA. XI.

ADDENDA.

Mr. E. Riek has informed me that the type of Cerioides mastersi Ferg. shows the
characters associated with wing folding and the type of C. apicalis Ferg. has a very
definite thyridium. :

A short note “On flies that fold their wings” in The Entomologist’s Monthly
Magazine, 1xxxi: 1945, 93-4, draws attention to the implications of this and other
anomalies. The abnormal course of veins in Nemestrinidae may be due to the survival
of primitive characters. It may be added that in the light of Alexander’s discoveries,
the breach between the Nematocera and Brachycera appears to be wider than credited,
and the Brachycera are not the lineal descendants of Nematocera but instead share with
it a common ancestry. Even the venation of Tanyderidae may be inadequate to assess
the homologies with the Brachycera.

REFERENCES.

ALEXANDER, C. P., 1929.—Int. Congr. Ent., 2: 700-7.

CRAMPTON, G. C., 1936.—Bull. Brooklyn Ent. Soc., 33: 141-9.
Harpy, G. H., 1924.—Proc. Linn. Soc. N.S.W., 49: 447-60.

—_—— , 1933.—Ibid., 58: 420.

————,, 1934.—- Ann. Mag. Nat. Hist., (10) 13: 504.

————, 1939—Proc. LINN. Soc. N.S.W., 64: 34-5.

————.,, 1940.—Ibid., 65: 484-5, 490.

———_—,, 1942.—_Nature, 149 (3781): 441-2.

——_——,, 1944.—- Proc. Roy. Ent. Soc. Lond:, 19: 52-69.
————,, 1945.—-Proc. Roy. Soc. Qd., 56: 81-4.

JAMES, M. T., 1936.—Ann. Ent. Soc. Amer., 29: 634-6.
LUNDBECK, W., 1916.—Diptera Danica, 5:.10-104.

MACKERRAS, I. M., 1925.—Proc. LINN. Soc. N.S.W., 50: 489-561.
MANN, J. S., 1929.—Aust. Zool., 6: 17-49.

NICHOLSON, A. J., 1927.—Ibid., 5: 10-104.

Norris, K. B., 1936.—J. Roy. Soc. W. Awst., 22: 49-70.
WILLISTON, S. W., 1908.—Manual of North American Diptera, 3rd Ed., 405 pp.

147

PRINCIPAL MICROSPORE-TYPES IN THE PERMIAN COALS OF
NEW SOUTH WALES.

By J. A. DuLtHuUNTY, B.Sc.*
(Plate vii; three Text-figures. )

[Read 25th July, 1945.]

INTRODUCTION.

The preliminary study of microspore-types recorded in this paper was carried out
as a basis for investigating microspore assemblages on various horizons in the Kamilaroi
(Permian) coal measures of New South Wales. The descriptions are intended to
provide a survey of general types rather than a palaeobotanical contribution, as the
ultimate purpose of the work in its geological application, if characteristic assemblages
are found to persist on coal-bearing horizons, is to assist stratigraphical correlation
of coal seams. The types are defined in terms of a generalized classification, designed
to group closely related microspores, the characterization of which will require detailed
morphological studies in botanical research. It is anticipated that such work will
eventually show that many of the general types described here may be subdivided into
several specific types of individual character. No attempt has been made to identify
the microspores with regard to the plans which produced them, as this also will require
specialized botanical work.

In the case of some of the types it is difficult at this stage to differentiate between
spores and pollen grains, and it is proposed to refer, tentatively, to all types as micro-
spores for the purpose of present descriptions.

CLASSIFICATION BASED ON PHYSICAL FEATURES.

A tabular system of classification depending on physical features has been drawn
up to serve as a key to the microspore-types. This system (Table 1) consists of
reference squares corresponding to two sets of major features: different kinds of
ornamentation and number of wings set out along the top, and shape of body and
nature of tetrad scar indicated on the left-hand side. The numbered squares represent
principal possibilities in type variation. The microspores are allotted type-numbers,
according to the numbers of the squares to which they correspond. When several
microspores corresponding to one square differ only in size or degree of development
of certain features, they are distinguished by letters added to the type-numbers. For
example, ellipsoidal, monolete microspores with striate ornamentation correspond to
square 23, but differences in arrangement of striae (transverse and longitudinal) give
two types, 23A and 23B. Microspores corresponding to some of the possible types have
not been found, and such squares remain blank to provide for the inclusion of additional
types, if such should be recognized. This system of tabular classification does not
infer any genetical relation between different types. In the absence of exact knowledge
of relations between microspores and plant species, it enables the counting of micro-
spores and determination of assemblages in terms of general types. But, at the same
time, it is recognized that variations in proportions of different types from one horizon
to another must indicate proportional modification in plant assemblages, resulting from
environmental or evolutionary changes.

* This investigation was undertaken when the writer held a Linnean Macleay Fellowship
in Geology.

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BY J. A. DULHUNTY , 149

The dimensions of the microspores are given in microns, measurements being made
in the optical plane in positions and along directions indicated in each case. A limited
number of types is illustrated by photomicrographs in Plate vii, showing general
appearance, and all types are illustrated by line drawings in Figs. 1, 2 and 3 to show
detailed features. All illustrations are at standard magnification of 400 diameters,
giving size comparison.*

MerHop or ISOLATING MIcCROSPORES AND PREPARING PERMANENT MOUNTS.

The technique used in isolating microspores together with fragments of resistant
plant tissue is similar to that described by Raistrick and Marshall (1939). Each
coal-sample is crushed to pass a 20 mesh I.M.M. sieve, producing as little fine powder
as possible, and the portion retained on a 30 mesh sieve is used for treatment. About
4 gm. is placed in a 100 ml. conical flask with 6 gm. of potassium chlorate and 60 ml.
of concentrated nitric acid—the flask being immersed in water to prevent heating during
the early stages of reaction. Oxidation is allowed to continue for 2 to 24 hours,
depending on room temperature and hardness of coal. The residue is then washed five
times with water by decantation to remove the acid solution. Owing to the granular
nature of the residue, only two to three minutes are necessary for settling between
each wash. The oxidized coal-substance is then dissolved by adding 50 ml. .of 10%
(Sp. Gr. 1.11) sodium hydroxide’solution and stirring, after which water is added, and
the extremely fine residue containing microspores allowed to settle for one hour before
decantation. It is then washed five times, one hour being allowed for settling each
time. After final decantation 2 to 3 ml. of strong alcoholic safranin is added, and the
moist residue allowed to stand for half an hour. It is then stirred vigorously, allowed
to settle for about 15 seconds, and the thick supernatant liquid containing microspores
poured onto filter paper. The short time allowed for settling before filtering assists in
obtaining an improved concentration of microspores by eliminating much of the mineral
matter which sinks and remains behind in the flask.

The use of granular coal (between 20 and 30 mesh I.M.M. sieve) for oxidation as
described above, eliminates fine powder which contains a large proportion of fractured
microspores, fine mineral matter and powdery carbon from fusain bands. These are
undesirable in the final microspore-concentrate, as they decrease the clarity of mounts
for microscopic examination. Influence on proportions of microspore-types, due to
removal of fine coal-powder, was tested by separate oxidation of both granular and fine
materials from one coal-sample. Counts made on the final mounts showed almost
identical proportions of microspore-types; but, in the case of the finely powdered
portion, counting was difficult owing to large numbers of fractured microspores, and
the presence of finely-divided mineral and carbonaceous matter.

In the preparation of permanent mounts about 1 ml. of strong glycerine-jelly is
melted by warming, and small portions of the moist microspore-concentrate are mixed
in until a drop of jelly, mounted on a slide and examined microscopically, shows the
desired concentration has been reached. Three permanent mounts are then prepared
from each microspore-concentrate.

SYSTEMATIC DESCRIPTIONS OF MICROSPORE-TYPES.
Type 1A. (Fig. 1. Plate vii, 1A.)
Angular tetrahedral; sharply-defined apices; flat or slightly convex sides; triangular
outline. Apices of tetrahedron to opposite sides 20--36u. Trilete; well-developed

triradiate sutures extending to distal apices, and frequently opened towards proximal
apex. Exine psilate.

* As a result of a suggestion by Dr. A. B. Edwards (Department of Geology, University of
Melbourne), and consultation with other workers interested in microspore work on Australian
coals, it is proposed that the tabular system of type-numbering adopted in this paper (Table 1)
be used in subsequent work on both Permian and Mesozoic microspore-types, but that in addition
the letters P, T, J or C be placed before the type numbers to signify Permian, Triassic, Jurassic
or Cretaceous, respectively.. For example, the Permian type 3A (described here) would become
P3A, and a similar type, if found in Jurassic coal, would be J3A.

Q

150 MICROSPORE-TYPES IN THE PERMIAN OF N.S.W.,

Tipe LBs (bies a Plater vais Bs)

Angular tetrahedral; similar to Type 1A, but much larger; 55-904 from apices to
opposite sides in proximal view. Trilete; sutures frequently opened as far as the distal
apices. Exine psilate.

Tape Ams (ies Plates vail 2A)

Subangular tetrahedral; rounded apices; convex sides; ill-defined triangular outline.
Trilete; somewhat curved lines extending towards distal apices tend to assume
triradiate or crude spiral arrangement. Size 36-554 from apices to opposite sides in
proximal view. Exine psilate.

50
MICRONS

Bp Jie Aus DWI En UN TY; 51

Type 3A. (Fig. 1... Plate vii, 3A.)
Ellipsoidal. Monolete dehiscence along a line running full length of body gives

rise to longitudinal opening expanded towards extremities. Length 40-654; width in
lateral view 24-36u. Exine psilate. d

Type 3B. (Hig. 1.. Plate vii, 3B.)

Generally ellipsoidal; elongated curved body; curved, oval outline. Appears to be
monolete with closed suture on more convex side of curved body. Length 90-140u;

PEON
wy Ye

ANY
HF 00.9"

IN

S
Ay

(oe) 50 100
(a
MICRONS

152 MICROSPORE-TYPES IN THE PERMIAN OF N.5.W.,

width 40-754. Exine generally psilate, although several longitudinal wrinkles or folds
appear on the walls.

Type: 3C. (Hig. 1. Plate vii, 3C:)
Ellipsoidal. Monolete; longitudinal suture, at times opened with lips turned out-
wards, runs full length of body. Length 20-304; width 12-17y. Exine psilate.

Type 4A. (Fig. 1. Plate vii, 4A.)

Spheroidal; somewhat flattened proximal and distal sides; circular to slightly oval
outline. Trilete with frequently opened sutures extending from centre to margin in full
proximal view. Diameter 20-—40u. Exine psilate.

Type 4B. (Fig. 1. Plate vii, 4B.)

Flattened spheroidal shape generally similar to Type 4A, but much larger and more
variable in size; diameter 75-1354. Trilete with well-developed triradiate sutures
extending to, or beyond, margin in proximal view. Dehiscence common along sutures,
in some instances tending to trisect the spore case.

Type 4C. (Fig. 1. Plate vii, 4C.)

Spheroidal with much flattened proximal and distal sides. Circular outline in axial
view. Trilete; triradiate sutures extend short distances from their origin, but give
rise to small, well-defined triangular opening at centre. Diameter 40-80u. Exine psilate.

Type oAwy ihe i Plate: vais) An)

Spheroidal with slight distal and proximal flattening giving circular or slightly
oval outline. Monolete dehiscence gives rise to large oval opening on proximal side.
Normally small but varies considerably; diameter 16—-40u. Exine psilate.

Type dB. (Fig. 1. Plate vii, 5B.)

Spheroidal to suboblate with circular or slightly oval outline. Monolete with ill-
defined suture or elongated area appearing straight, or curved, depending on position,
but seldom exhibiting distinct opening. Size variable, 45-120u. Walls appear relatively
thin. Exine psilate.

TypevoC.- (hice ter elatey wai Cs)

Spheroidal; distinctly flattened and circular outline in axial view. Appears to be
monolete, although regular sutures or distinct openings have not been observed. Usually
very small with diameter of 17—20u, but varies from 14—-40u. Exine psilate and very
thick-walled giving a dark rim round the body.

Type 6A. (Fig. 1. Plate vii, 6A.)

Angular tetrahedral; well-defined apices; flat or slightly convex sides; triangular
outline. Trilete with long, frequently opened sutures extending to distal apices. Size
variable, 40-100u from apices of the tetrahedron to opposite sides. Exine granulate,
normally of fine granular texture, but somewhat variable.

Type 7A. (Fig. 1. Plate vii, 7A.)

Subangular tetrahedral; rounded apices; convex sides; ill-defined triangular outline.
Trilete; curved triradiate sutures seldom reaching distal apices. Size variable, 50-110
from apices of tetrahedron to opposite sides. Exine granulate, normally of fine granular
texture, but somewhat variable. i |

Type SA. (Fig. 1. Plate vii, 8A.)

Ellipsoidal. Monolete; dehiscence extending full length of body produces longitudinal
marginal opening usually expanded towards its extremities. Length 40-504; width in
lateral view 23-28u. Exine granulate of medium-fine granular texture.

Type 9A. (Fig. 1. Plate vii, 9A.)

Generally spheroidal; somewhat flattened; circular to slightly oval outline. A large
depressed triangular area on one side of the body indicates a tetrad scar of trilete
character. Diameter 24-38u. Exine granulate of coarse granular texture.

BY J. A. DULHUNTY 153

Type 9B. (Fig. 1. Plate vii, 9B.)

Spheroidal; slightly to distinctly flattened; circular outline. Trilete with well-
developed triradiate slits, which frequently tend to trisect the spore. Size variable;
diameter from 50-100«. Exine granulate varying from fine to medium-coarse granular

texture.

AQ

' USNS \

SS

\
aN

AN

wee

ie) 50 100
MICRONS

154 MICROSPORE-TYPES IN THE PERMIAN OF N.S.W.,

Type 10A. (Fig. 1. Plate vii, 10A.)

Spheroidal to suboblate; somewhat flattened; circular to suboval outline. Monolete
tetrad scar up to 54 in width at centre. Diameter 40-60u. Exine granulate of medium-
fine granular texture.

Type 10B. (Fig. 1. Plate vii, 10B.)

Spheroidal to suboblate; slightly flattened; circular to suboval outline. Monolete;
well-developed slit extending the full width of the spore, in lateral view is usually open
and expanded towards its extremities. Lips of the slit may protrude outwards. Diameter
50-80u. Exine granulate of medium-fine granular texture.

Type 138A. (Fig. 1. Plate vii, 13A.)

Ellipsoidal with elongated oval outline. Monolete with indistinct suture situated
marginally in lateral view. Length 50-704; width in lateral view 30—40u. Exine usually
exhibits several longitudinal folds or wrinkles, and a fine reticulum of anastomosing
ridges between the folds.

Type 14A. (Fig. 1. Plate vii, 14A.)

Spheroidal; much flattened on proximal and distal sides; circular outline in axial
view. Trilete with short triradiate slits normally opened to form a small triangular pit.
Diameter 40-90u. Exine reticulate with fine network of ridges. Ektexine thick, seen
as dark band round the body in optical section.

Type 15A. (Fig. 1. Plate vii, 15A.)

Spheroidal to suboblate; slightly flattened; circular to suboval outline. Tentatively
placed as monolete, although no definite slit or opening has been observed. Diameter
40-65u. Exine reticulate, traversed by a widely-spaced system of large, anastomosing
grooves about 4u in width, and 10y apart.

Type 16A. (Fig. 2. Plate vii, 16A.)

Tetrahedral; well-defined apices; flat to slightly convex sides; triangular outline.
Trilete with slits extending to distal apices, and frequently opened. Size 40-704 from
apices to opposite bases of the tetrahedron. Exine echinate with small spines 3—4y long,
and 1—2u wide at base, set 2-3 apart.

Type 17A. (Wig..2. Plate vii, 17A.)

Subangular tetrahedral; rounded apices; convex sides; rounded triangular outline.
Trilete with short, ill-defined sutures extending towards distal apices. Diameter about
30-60u. Exine echinate with large widely spaced spines of regular size pattern. Spines
up to 7u in length, and 3u in width at base, set about 5u apart.

Type 18A. (Fig. 2. Plate vii, 18A.)

Generally ellipsoidal. Definite evidence of tetrad scar or dehiscence lacking; but a
longitudinal collapsed area suggests monolete character. Length 90-110u; width 45—-55n.
Exine echinate, spines up to 7u in length and 3” in width at base, set about 3u apart.

Type 19A. (Fig. 2. Plate vii, 19A.)

Spherical or slightly oblate with circular or slightly oval outline. Trilete with
short ill-defined sutures, rarely open. Diameter 28—40u. Exine echinate with spines of
constant size pattern on each individual, but varying on different individuals from
3 to 6 in length, and 1:5 to 2-54 in width at base, typically spaced about 6y apart.

RY Dee OA] (Hick 2am atemystine 2 Ae)

Spheroidal, but usually collapsed on one side, giving an outline which is completely
circular or semi-circular with one side flattened depending on position. Nature of
dehiscence or tetrad scar uncertain; but collapsed area on one side suggests monolete
character. Diameter of sphere 60-80u. Exine echinate with spines up to 6u in length
and 3 in width at base, set about 3u apart.

BY J. A. DULHUNTY 155

Type 21A. (Fig. 2. Plate vii, 21A.)

Tetrahedral; well-defined apices; slightly convex sides; generally triangular outline.
Trilete with straight, well-defined triradiate sutures extending to distal apices. Size,
90-1104 from apices to opposite sides of tetrahedron. Situated at the centres of the
three distal interfaces are three dome-shaped protuberances, each about 25u wide at its
base, projecting about 124 beyond the margin of the body. In some cases a small cone-
shaped projection, about Su at base and 7u long, is present at the top of each dome. The
exine of the three proximal faces is marked by coarse striae radiating from the base of
each dome towards the triradiate sutures. The sutures are normally closed; but in
some individuals they have been seen open to a width of 20u, indicating definite
dehiscence.

Mype 2A. (Rig. 2: Plate vii, 23A.)

Ellipsoidal. Monolete suture extending full length of body along-lateral margin;
normally closed. A narrow frill-like fringe of smooth exine, 10-134 wide, is associated
with the suture, and projects beyond outline of body in lateral view. Length 50-60u;
width 40—50u in lateral view. Exine ornamented with transverse striae about 3u apart
extending full width of body.

ypey23B. (His, 2) Plate vii, 23/82)

Ellipsoidal. Monolete with straight lateral suture, usually closed, extending full
length of body. Length 55-654; width 40-454. Exine ornamented with longitudinal
striae about 3-54 wide extending full length of body.

Type 26A. -(Fig. 2. Plate vii, 26A.)

Angular tetrahedral; apices well-defined; sides flat or slightly convex; triangular
outline. Trilete with triradiate sutures extending to distal apices. Size 64-734 from
apices of tetrahedron to opposite sides. Exine verrucate with small, closely-packed,
-rounded elevations about 2-54 in diameter and ly apart.

Type 28A. (Hig. 2. Plate vii, 28A.)

Ellipsoidal; ellipse of low eccentricity; suboval outline. Monolete with well-defined
suture running full length of body. Length 50-70u; width 40-50u in lateral view. Exine
verrucate with small, widely-spaced, rounded elevations about 2u in diameter, and
4—-5u apart.

Type 29A. (Fig. 2. Plate vii, 29A.)

Approximately spheroidal; slightly flattened; rounded to slightly oval outline.
Diameter 35-50u. Trilete with well-marked triradiate sutures extending to margin in
full proximal optical section. Exine verrucate with rounded elevations about 2yu in
diameter and 3u apart.

MM DCAD Bree 2h) Plater vil, 29.B:)

Spheroidal; slightly flattened; circular outline. Trilete with well-developed
triradiate sutures, frequently opened, extending to margin in full proximal optical
section. Diameter 130-170u. Exine verrucate with large rounded elevations 4-5y in
diameter, and 2-3 apart.

Type 380A. (Fig. 2. Plate vii, 30A.)

Spheroidal to suboblate; slightly flattened; circular to suboval outline. Monolete
with short, ill-defined suture, at times accompanied by a small collapsed area. Diameter
35-504. EHxine verrucate with large, rounded elevations 6—7u% in diameter, and 2-3
apart.

Type 382A. (Fig. 2. Plate vii, 32A.)

Monowinged with subangular, tetrahedral body exhibiting rounded apices; convex
sides; rounded triangular outline. Trilete with well-defined triradiate sutures seen as
double lines along proximal interfaces. Body psilate, 50-80u from apices of tetrahedron
to opposite sides. Wing marked with radiating venation, and situated in one plane

156 MICROSPORE-TYPES IN THE PERMIAN OF N.S.W.,

round body and attached to distal interfaces of the tetrahedron, has an approximately
circular outline and crenulate margin; wing 15-20 wide.

Type 388A. (Fig. 2. Plate vii, 33A.)

Monowinged with ellipsoidal body. Monolete; single longitudinal suture about 5u
wide extending almost full length of body. Body psilate, 60-80% long, and 40-604 wide.
Single narrow wing with oval outline, and with faint reticulate marking varying in
width from 3u to 104 in one individual, situated in lateral plane round body.

Type 338B. (Fig. 2. Plate vii, 33B.)

Monowinged with ellipsoidal body. Monolete; single longitudinal suture, 5—-7u wide,
extending almost full length of body. Body psilate, 80-110u long, and 40-60% wide.
Single wing, 28—40u wide with oval outline, situated in lateral plane round body; faintly
reticulate, and somewhat granular appearance.

Type 34A. (Fig. 3. Plate vii, 34A.)

Monowinged with flattened spheroidal body. Trilete character indicated by short,
indistinct triradiate sutures seen only in well-preserved examples. Body psilate, 50-80u
in diameter. Single circular wing, 12-184 wide, continuous in one plane round body,
and marked with radiating reticulum. Several concentric lines appear at junction
between wing and body.

Type 34B. (Fig. 3. Plate vii, 34B.)

Monowinged with flattened spheroidal body. Trilete characters similar to Type 34A.
Body psilate, 90-1204 in diameter. Single circular wing, 20-40u wide, continuous in
one plane round body, and marked with radiating reticulum. Several concentric lines
occupy relatively wide zone at junction between wing and body.

Type 84C. (Fig. 3. Plate vii, 34C.)

Monowinged with spheroidal body. Trilete characters similar to Types 34A and
34B. Body psilate, 100-130u in diameter. Single circular wing, 25-35u wide, continuous
in one plane round body, and ornamented by fine granular marking. Several concentric
lines appear at junction between wing and body.

Type 385A. (Fig. 3. Plate vii, 35A.)

Monowinged with flattened spheroidal body. Monolete character indicated by
indistinct, narrow. germinal area extending across body. Body psilate, 50-80u in
diameter. Single wing with oval outline and reticulate venation, continuous in one
plane round body, and situated with short axis of elliptical outline in direction of
germinal area. Width of wing 9-124 in direction of germinal area, and 20-30 in
opposite direction. Narrow zone at junction between body and wing, widest in direction
normal to that of germinal area.

Type 388A. (Fig. 3. Plate vii, 38A.)

Biwinged with ellipsoidal body. Monolete, exhibiting narrow, longitudinal germinal
area extending full length of body. Body psilate, 40-80% long, and 30-60u wide. Two
wings, marked with radiating venation and normally equal to or slightly larger than the
body but of variable shape and size, are situated symmetrically on either side of the
germinal area towards which zones of attachment extend.

Type 40A. (Fig. 3. Plate vii, 40A.)

Biwinged with slightly flattened spheroidal body. Monolete, exhibiting narrow
germinal area extending across body between roots of wings. Body, 25—40u in diameter,
frequently exhibits coarse striae transverse to germinal area. Two relatively small,
somewhat elongated wings marked with radiating venation, and normally slightly wider
and considerably longer than diameter of body, but of variable shape and size, are
situated symmetrically on either side of germinal area towards which zones of attach-
ment extend.

BY J. A. DULILUNTY 157

Type 40B. (Fig. 3. Plate vii, 40B.)

Biwinged with slightly flattened spheroidal body. Monolete, exhibiting narrow
germinal area extending across body between roots of wings. Body psilate, 35-50u in
diameter. Two relatively large, rounded wings marked with reticulate venation,
frequently twice width of body but of variable shape and size, are situated symmetrically
on either side of the germinal area with large zones of attachment.

Type 40C. (Fig. 3. Plate vii, 40C.)

Biwinged with flattened spheroidal body. Monolete character similar to Type 40B.
Body psilate, 50-S0u in diameter. Two wings with reticulate venation, normally rounded
or slightly elongated and about same size as body but of variable size and shape, are
situated symmetrically on either side of germinal area with large zones of attachment.

Type 40D. (Fig. 3. Plate vii, 40D.)

Biwinged with flattened spheroidal body. Germinal area not evident, but probably
situated between roots of wings. Body 35-55u in diameter, marked with coarse striae
between wings. Two small lobe-like wings with wrinkled or folded surfaces, attached
symmetrically to one side of the body, protrude only a limited distance beyond its
margin, and are directed away from the proximal part of the spore.

Type 41A. (Fig. 3. Plate vii, 41A.)

This microspore has been tentatively classified in terms of Table 1, as its morphology
is not yet clearly understood. The illustration in Fig. 3 is intended to show the appear-
ance in optical section. The general features are typical, and the type has been
recognized in coals from a number of different seams. The spore appears to consist of
a flat disc-shaped portion (45—70u in diameter), to one side of which is attached a large,
elevated structure projecting considerably beyond the margin of the disc. The exine is
psilate, and no opening or dehiscence has been observed.

ACKNOWLEDGEMENTS.

The author wishes to acknowledge the assistance of F. V. Mercer, B.Sc., Department
of Botany, University of Sydney, in the interpretation of microspore structures, and
valuable discussion with Dr. A. B. Walkom, Director, Australian Museum. The
terminology adopted by Erdtman (1943) has been adhered to as closely as possible in
describing the spore-types.

REFERENCES.
RAIstTRick, A., and MARSHALL, C. E., 1939.—Nature and Origin of Coal and Coal Seams. English

Universities Press Ltd., London, p. 130.
BERDTMAN, G., 1943.—An Introduction to Pollen Analysis. Chronica Botanica Company, Waltham,

Mass., U.S.A.

EXPLANATION OF PLATE VII.
Photomicrographs of some of the principal microspore-types occurring in the New South
Wales Permian coals, x 400.

158

PETROLOGICAL STUDIES IN THE ORDOVICIAN OF NEW SOUTH WALES. III.
THE COMPOSITION AND ORIGIN OF THE UPPER ORDOVICIAN GRAPTOLITE-BEARING SLATES.
By GERMAINE A. JOPLIN, B.Sc., Ph.D., Linnean Macleay Fellow of the Society in Geology
(Four Text-figures. )

[Read 25th July, 1945.]

Contents.
Page.
I. Introduction ae foe” Megeteem, wee eyo Nets bce Meme Bs here Seah a ues en heen Ree 5 9 - aLiNss
Il. The graptolite- penis slates see ten SAN votre as 9. toe (ue ey agatiene Sat one We PAMELA RIES a LL tS
1. General description ewan acne ice GE wk RUN aN © yay: th a ene Pc Bioy es 5g. lS
2. Stratigraphical position te ae ee 42 ose a Ris Sie rece sg ALOU
III. Origin of the black slates of New South Wales ad FV ns ah as ae eat eae aR ee Mh eucd is |
1. Possibility of silicification PRs S eee Nicer a 5 SPER ees ee at Bich | ar euen? a errs «Loh
2. Origin of the siliceous material Byer an Lost Prise Ata cero OD
3. Amount of volcanic ash available for the oration ie alates ae ‘ BG a AO
4. The occurrence of lavas and ash in the Ordovician sequence Bieeetene aes so LOY
IV. Conditions necessary for the accumulation of black shales eee cee bc on eens emeliG 3
1. Earlier hypotheses pat APAATE. ES te cried ae an ois ae Pat cei Lee RSG a ae ta calk jes
2. The volcanic ash nepornecia PERC RES) omeL SECON aN Renal arene MLAS OMEN Ua da ites CO pier eRe nie, ero,
Vi. SUMmMary 52. Pe a Se BE ee ed ie a ae ats eye oo Re eee Saw Recta wees aThO,
Vi. Ne neicdeement. oe PSEA ee cont cis aaNet ae Fe Per ene esa ee hieee is Kecae eT
VII. References Be eas Cen tie Oke ae sete ie See RT ae a ae ae Be aera emt rE red (KO)

I. INTRODUCTION.

Some years ago, when making a detailed study of the Ordovician rocks at Cooma,
I had occasion to analyse a graptolite-bearing black slate of Upper Ordovician age.
A number of more highly altered pelitic sediments previously analysed showed a
very uniform composition, with a silica range of about 54-61%, and it was with
some surprise that I found that this less altered type contained over 80% of silica.
Two other rocks, on approximately the same stratigraphical horizon and from widely
separated parts of the same district, also proved, on analysis, to be highly siliceous, and
this led to an investigation of the graptolite-bearing slates from other localities in New
South Wales. The results of this study are presented in the present paper.

II. THE GRAPTOLITE-BEARING SLATES.
1. General Description.

In every area where these rocks occur (see map, Fig. 1) there is a remarkable
lithological uniformity and they are found either as black or dark bluish-grey slates, as
light grey or white fissile slates or as cherts. The graptolites are often very well
preserved and appear either as black carbonaceous films, as white micaceous films or as
reddish-brown films of iron oxide. The last type of preservation seems to be more
common in the light coloured, fissile varieties which have probably been oxidized and
leached; except for a lower carbon content they do not differ markedly in composition
from the black slates with which they are usually interbedded (see Table 2, Analyses
C and L).

The slates are usually in a low grade of regional metamorphism, and a well-marked
slaty cleavage, particularly in the light coloured leached types, is often well developed.
Slate quarries have been worked at Ariah Park, near Temora, and the Yalgogrin
occurrence was investigated with a view to exploitation as roofing slates. Sometimes
the slates are much veined with quartz and, when near a granite contact, chiastolite is

BY GERMAINE A. JOPLIN. 159

LOCALITY MAP

showing

DISTRIBUTION OF UPPER ORDOVICIAN s a pore

—
Narromine— af
\ al ;
Tomingley Wellington
*Apsley
Peak Hill
Condebolin-——Molong aad
Parkes O
range
1
Weia Cadia Blayney
* Ungarie *Mandurama fitoken BAY
Wyalong Trunkey
ee hae | SYDNEY
oe Jy)}/ j |
Ariah Park ate VY, |
oe aralgal / Me H
. Temora Claes WoLLonconc

Bray ton

Yass,|. \Tewrang. tation lf
fin Jerfaw YaGoulBurn ~ SH
Wagga ay BeBune nell me
CANBERRA ey Siew 1S Ray a
Queanbeyan ify YL) Sy
yi

SN era oan
Albay Rien C A
Berndale
3
\

—
= |e

Bombala
ae

at

147° [

often developed. In the chiastolite slates graptolites are easily recognizable and appear
to be preserved in a white micaceous material.

Although the slates have been examined under very high magnification, much of the
material is not resolved by the microscope. The rocks are often banded with finely
carbonaceous seams alternating with carbon-free or carbon-poor bands (Fig. 2B). The
non-carbonaceous seams appear to consist of tiny flakes of sericite, small zircons,
needles of rutile and occasional tiny grains of quartz in a faintly green almost isotropic
base which is not completely resolved. Minute Y-shaped bodies sometimes occur, and
these may possibly represent sponge-spicules, whilst tiny rounded patches of chalcedony
are doubtfully interpreted as radiolaria. Other, less regular, patches of radiating
chalcedony have also been noted, and it is possible that some of the base may consist
of this mineral. This chalcedony and the occasional minute quartz veins point to
secondary silicification, but it appears to be an internal rearrangement of a primary
constituent. The amount of quartz and sericite varies in the different slates and when
present as distinct grains the quartz often shows a curved fracture.

Biotite is developed in some of the slightly metamorphosed types and chiastolite
and incipient cordierite are not uncommon when the slate occurs near a granite contact
(Hig. 2C).

160 PETROLOGICAL STUDIES IN THE ORDOVICIAN OF N.S.W. III,

Reference to Table 2 will show that slates from seven widely separated localities in
New South Wales show marked chemical similarity. Though there is a range of some
10% of silica, it is always very high, and with the possible exception of Analysis A,
there is every reason to believe that no silica has been added from an outside source.
The analyses are arranged in order of decreasing silica without reference to their
locality, and it can be seen that there is a slight increase of alumina and potash
accompanying the decrease in silica and that the amount of ferric oxide is a little
variable, but for all these differences, the variations are only minor ones and it can be
seen that the slates all belong to a single rock-series.

The graptolite-bearing rocks are interbedded with buff coloured slates of slightly
coarser texture or with more sandy types which range from fine-grained quartzites
to medium-grained sandstones coloured white, buff, red, grey or black. In these, negative
erystals of pyrites are not uncommon.

Near Tallong, in the Shoalhaven Gorge at Badgery’s Crossing, perfectly preserved
graptolites occur as black carbonaceous films in a dark grey quartzite. Under the
microscope this rock is seen to consist of somewhat rounded and sub-angular quartz
grains in a matrix of sericite, chlorite and some isotropic material. Zircons and rutile
can be recognized. In rather similar rocks at Cooma (Jeplin, 1942, p. 160) occasional
small grains of oligoclase-andesine (Ab,An,-Ab,,An,,) occur with rounded or
sub-angular grains of quartz. The occurrence of fairly fresh felspar in these rocks
suggests wind action, ice action, volcanic action or close proximity to the source of
supply. The conchoidal fracture of the quartz grains (Fig. 2A) may be interpreted as
the result of sub-aerial weathering and temperature changes (Waterhouse and Browne,
1929) or as the result of shattering by volcanic activity (Pirsson, 1915).

2. Stratigraphical Position.

Reference to Table 1 will show that Upper Ordovician graptolites have been
recorded from a number of widely separated localities in New South Wales, but it will
also show that intensive collecting and detailed descriptive work has been confined only

an Sn ae B eee C 1mm

1M

A. Low-grade psammite in Coolringdon Beds on Berridale Road, near Slack’s Creek, SW. of
Cooma. The rock shows rounded and sub-angular grains in an argillaceous and ferruginous
matrix. Note curved fracture in quartz grains, and small patches of clastic mica. x12.

B. Siliceous slate with carbonaceous bands from Wambrook Creek at crossing of
Adaminaby Road, W. of Cooma. Note tiny clastic grains of quartz and iron ore. x 12.

C. Chiastolite-slate, Gygederick Hill near Berridale. Former slight schistosity indicated
by orientation of incipient cordierite crystals. The groundmass consists mainly of finely divided
carbonaceous material and quartz. x12.

Reprinted from these PROCEEDINGS. Ixvii, 1942, 161.

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of
6

‘TL WIAVI,

BY GERMAINE A. JOPLIN. 161

to a few areas. In consequence of this, no zoning is possible at present, and as several
graptolite zones are possibly represented among the collections from a single locality, it
was considered useless to publish lists of the various species; furthermore it was also
impossible to tell precisely from which graptolite-horizon each analysed specimen had
been taken, and it seems likely that the analysed rocks listed in Table 2 represent a
number of different horizons in the Upper Ordovician sequence. In Table 1 a complete
list of publications for each locality is given, and though some of these refer to general
information concerning the slates, most deal with the graptolites, and to these the
reader is referred for detailed information on the fauna. Most of these references have
been compiled by Keble and Benson (1939), but Table 1 has been included in the
present paper since it has been arranged in such a way that all available information
about a single locality may be seen at a glance and may be compared with available
information from other Upper Ordovician localities.

Schists believed to be of Ordovician age occur at Cooma, Albury, Jingellic, Junee
Reefs and possibly in the Trunkey and Yalgogrin districts. At Cooma these occur in
close proximity to the Upper Ordovician graptolite-bearing slates and there appears
to be a gradual progressive metamorphism from slates into schists. In the metamorphic
area of North-East Victoria, of which the Albury district is the northern extension, a
similar progressive change has been observed from graptolite-bearing Upper Ordovician
slates into high grade schists, and on this account, the schists have been regarded as
Upper Ordovician sediments (Howitt, 1889; Browne, 1914, 1929, 1943; Tattam, 1929;
Joplin, 1942, 1943). Reference to Tables 2 and 3 will show that the slates and schists
form two chemically distinct rock-series, and furthermore, in the only locality in New
South Wales, namely Cooma, where both series have been studied, they appear to occupy
distinct stratigraphical horizons and no interbedding of the one type with the other has
ever been observed. In this area the siliceous slates have been termed the Coolringdon
Beds and the aluminous schists the Binjura Beds (Joplin, 1942, 1943). Interbedded with
the schists of the Binjura Beds, however, sandy schists, containing some plagioclase
felspar, often occur and these have a composition rather similar to the siliceous slates
(Joplin, 1942, p. 161, Table 2, Anal. IV).

If the schists are of Ordovician age and are not interbedded with the siliceous
slates, then they must be either above or below them. According to Dr. W. R. Browne’s
interpretation of the structure (Browne, 1943) the Binjura Beds overlie the Upper
Ordovician slates.

My interpretation of the structure at Cooma (Joplin, 1943) places the Coolringdon
Beds above the Binjura Beds and below the Bransby Beds—a series consisting of
porphyries, rhyolites, tuffs and limestones (see Fig. 3).

Unfortunately the isoclinal folding and the possibility of overthrusting at Cooma
make the interpretation of the structure very difficult, and as the Coolringdon and
Binjura Beds have not yet been found together elsewhere in New South Wales, only
indirect evidence can be used in interpreting the structure. At Trunkey, Raggatt (1934)
finds black slates above schists, but as graptolites have not been found, their age is not
certain. It is hoped that graptolites may be found at Jingellic where the Binjura
schists are highly carbonaceous and in a comparatively low grade of metamorphism, but
in the meanwhile the relative age of the Coolringdon and Binjura Beds must remain in
doubt.

III. OriciIn oF THE BLACK SLATES oF NEW SOUTH WALES.
1. Possibility of Silicification.

The occurrence of quartz veins and the not infrequent proximity of granite (see
Table 1) immediately raises the question of silicification of the black slates. In fact
many of the siliceous slates have been described as “‘silicified slates’’ and there is no
doubt that local silicification has taken place in some areas. In selecting material for
analysis, however, care was taken to avoid any specimens containing quartz veins or
any apparent addition of silica from an outside source. As pointed out above, a
microscopic examination sometimes revealed that there had been a slight silicification
which could be attributed to an internal arrangement of the original silica of the rock.

III,

ORDOVICIAN OF N.S.W.

ICAL STUDIES IN THE

1
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164 PETROLOGICAL STUDIES IN THE ORDOVICIAN OF N.S.W. _ III,

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BINJURA BEDS

Basic Granulite ( Basalt).
ata ar st Cooma Gneiss.

1a Be
Columnar section showing approximate sequence of the Ordovician as exposed in the Cooma
Region. The thickness of beds has not been measured and the section is therefore not drawn to
scale. Note presence of tuffs above and below the Coolringdon Beds and rhyolites in the
Bransby Beds.
Reprinted from these PROCEEDINGS, Ixviii, 1943, 161.

As previously indicated, the two chemically distinct rock series listed in Tables 2
and 3 are never found interbedded, but there is a close field association at Cooma, in
certain areas of North-East Victoria, and possibly in the Trunkey district of New
South Wales (Raggatt, 1934; Browne, 1935). As Table 3 lists the only normal pelitic
rocks anywhere in the vicinity of the siliceous slates, it must be assumed that if the
latter are the result of silicification then the normal pelite was the parent rock.

In the first place it would seem to be rather a coincidence that all the normal
pelites were silicified to about the same degree and that no gradations in the progressive
silicification occur. It is true that the slates show a variation of about 10% of silica
and that there is a sympathetic variation in the other oxides, but apart from other
differences there is a large gap between the series that could reasonably be expected
to be filled if progressive silicification had occurred. There is sometimes a suggestion

BY GERMAINE A. JOPLIN. 165

that the presence of carbon promotes the addition of silica and if there is no further
selection than this, then all other oxides should be present in about the same amount as
those of the original rock. If any other single oxide were selectively replaced, then it
must have been necessarily an oxide present in large amount, for some 25% of silica
must have been added. Actually only alumina could have been present in sufficient
amount to meet this requirement, and an inspection of the analyses will show that
there is no justification for such an assumption. Assuming some constituent to have
remained constant, it would be possible to calculate what has been added to or
subtracted from the aluminous pelite in order to convert it into the siliceous pelite,
but this would lead to the dangerous assumption that one rock had actually been
derived from the other. In this case no such fact is known and actually with sufficient
manipulation of chemical analyses a rock of any type might be calculated into another.
Thus, while a two-way passage of oxides is not denied, it is telt that there is no
reasonable chemical evidence for such an assumption, and that such calculations might
prove a very fallacious argument. It is possible to assume, however, that there had
been no selection in the silicification, and if such were the case then the other oxides
of the slates should be present in the same ratios as those of the normal pelites. To
test out this assumption an A-F-C diagram was constructed with A = the molecular
proportion of alumina after satisfying alkalis, with C = the molecular proportion of lime
and F = the molecular proportion of magnesia plus total ferrous oxide after satisfying
titania for ilmenite. As the relative amounts of ferrous and ferric oxide vary with the
degree of metamorphism, the making of magnetite brought out untrue differences
between rocks that had approximately the same total iron, and it was considered that
a more valid comparison could be made between the analyses if ferric oxide was
reduced and added to ferrous oxide. As sulphur has been estimated only in a few of
the analyses, pyrites was not calculated. Further, it was considered inadvisable to -
satisfy phosphorus pentoxide with lime for apatite, as it seems not unlikely that the
phosphorus may be present in some of the low grade rocks as vVivianite or some earthy
form. Furthermore, phosphorus pentoxide has not been determined in some of the
older analyses of rhyolites and tuffs, with which the analyses of the slates are compared.
: Inspection of the triangular diagram (Fig. 4) will show that the normal pelites
occupy a very restricted field and that the field of the siliceous pelites, though not
quite so well defined, is distinct from that of the other type. As SiO. has not been taken
into account in this diagram, it has been possible to compare the ratios of Al,O,;, Fe.0,,
FeO, MgO, CaO, Na.O, K.O and TiO. in both rock series, and there is thus evidence
for the conclusion that the ratios of the various oxides in the two rock-series differ
and that there is no evidence for the assumption that the slates are silicified normal
pelites.

2. Origin of the Siliceous Material.

If the slates are not silicified slates but siliceous slates, then the origin of such
a sediment must be considered. Inspection of the analyses of silts and fine, deep-
water sediments (Clarke, 1916; Twenhofel, 1926) throws no light on the matter, but
Fig. 4 shows that a number of rhyolites and rhyolite tuffs compare with the slates in
the ratios of their respective oxides. Although only twenty-eight analyses of the
lavas and tutfs have been plotted, the analyses have been chosen from various parts
of the world, and although the area enclosed within the detted line is not strictly
the field of the rhyolites, it possibly represents that of most normal types. Reference to
Fig. 4 shows that the field of the siliceous slates falls within the less well defined one
of the rhyolites and rhyolitic tuffs. Moreover, No. 12, a rhyolite from the Ordovician
at Cooma, falls within the slate field.

It is therefore suggested that the slates consist largely of redistributed rhyolite tuff,

In describing the slates it was mentioned that they were interbedded with more
sandy types containing sub-angular quartz grains and occasional grains of andesine
(Fig. 14), and such features would be in accord with the suggestion of a pyroclastic
origin. Actually a little tuffaceous material of a coarser nature has been recognized in

166 PETROLOGICAL STUDIES IN THE ORDOVICIAN OF N.S.W. III,

the Binjura Beds and in the Bransby Beds of the Cooma district (Joplin, 1942, 1943),
but this appears small by comparison with the great amounts of volcanic dust that
must be postulated for the formation of the siliceous slates.

Fig. 4.

A-F-C diagram showing plots of analyses of siliceous slates (A, B, etc, Table 2), of
normal pelites (a, b, ete., Table 3) and of rhyolitic tuffs (triangles) and rhyolites (circles) as
listed below.

1. Voleanic Tuff. Tooloom, N.S.W.—In W.T., No. 37: 801.

2. Brisbane Tuff. Queensland.—C. Briggs, 1929, Proc. Roy. Soc. Qd., 40: 156.
3. Rhyolite Tuff. Westmoreland, England.—In W.T., No. 26: 801.

4. Liparite Tuff. Hritrea.—In W.T.; No. 33: 801.

5. Liparite Tuff. Sardinia.—In W.T., No. 30: 801.

6. Rhyolite Ash. Montana, U.S.A.—In W.T., No. 4: 797.

7. Rhyolite Tuff. Montana, U.S.A.—In Wi. Nos 53 799)

8. Rhyolite Tuff. (Lake bed deposit), Colorado, U.S.A.—In W.T., No. 19: 799.
9. Rhyolite Tuff. Oregon, U.S.A.—In W.T., No. 14: 799.

10. Rhyolite Tuff. California, U.S.A.—In W.T., No. 18: 799.

11. Rhyolite Tuff. California, U.S.A.—In W.T., No. 19:: 799.

12. Rhyolite. Cooma district, N.S.W.—G. Joplin, 1943, Proc. Linn. Soc. N.S.W., 68: 165.
13. Rhyolite Glass. Gloucester, N.S.W.—In W.T., No. 42: 97.

14. Rhyolite. Esk, Queensland.—C. Briggs, 1929, Proc. Roy. Soe. Qd., 40: 156.

15. Rhyolite.. Hauraki, N.Z.—In W.T., No 3: 107:

16. Rhyolite. Hauraki, N.Z.—In W.T., No. 45: 97.

17. Rhyolite. Hauraki, N.Z.—In W.T., No. 95: 91.

18. Rhyolite Perlite. Waiau Valley, Hauraki, N.Z.—In W.T., No. 101: 72:

19. Rhyolite- Tardree, Ireland.—F. H. Hatch, 1914, The Petrology cf the Igneous Rocks: 262.
20. Rhyolite (altered). Hungary.—In W.T., No. 29: 801.

21. Rhyolite. Hungary.—In W.T., No. 28: 801.

22. Rhyolite. Hungary.—In W.T., No. 58: 87.

3. Rhyolite. Japan.—In W.T., No. 39: 97.

24, )Rhyolite., California,) Us;SeAc— in Wil... NOt 19 R83:

25. Rhyolite:. California, (W:SVA:—in Wen: Now iis) sie

26. Rhyolite (altered). Idaho, U.S.A.—In W.T., No. 9: 799.

27. Pumice. Katmai, Alaska.—C. Fenner, 1926, J. Geol., 34: 695.

28. Pumice. Katmai, Alaska.—C. Fenner, 1926, ibid., 34: 695.

BY GERMAINE A. JOPLIN. 167

Above, it was mentioned that the graptolites sometimes occur in cherts, and it is
pertinent to note that this rock type is not infrequently of a tuffaceous origin. A good
example of chert formed in this way may be seen at Manildra, New South Wales
(Joplin and Culey, 1937) where there is a gradation from fossiliferous Silurian
breccias, through coarse and fine tuffs into cherts. Though these are of Silurian age
they bear a strong lithological resemblance to the Wellington cherts which contain
Upper Ordovician graptolites (Sherrard and Keble, 1928). It has also been suggested
that the fine banded cherts of the Upper Coal Measures in New South Wales are finely
divided tuffaceous material (David, 1907).

3. Amount of Volcanic Ash available for the Formation of Slates.

If it be assumed that the Upper Ordovician slates are deposits of volcanic ash or
dust, then it must be concluded that tremendous quantities of such pyroclastic material
were available. Possibly some of the shales may represent admixtures with
normal clastic sediments, but the bulk of the material must be presumed to be ot
volcanic origin. Siliceous waters associated with the volcanic activity may have played
a part in silicifying the ash deposits and any normal admixed clastic material.

The great bentonite deposits of the Upper Ordovician of North America (Nelson,
1922) are believed to represent ash deposits the volume of which has been estimated
as about 66 cubic miles. Nelson suggests that the source may have been a volcanic island
off the peninsula separating the Lowville Sea from the Palaeozoic Atlantic Ocean.

In reporting on the great Katmai eruption of 1912, Martin (1913) has given a
graphic description of what he considers to be “the most tremendous volcanic explosion
known in history’. Ash fell at a distance of 900 miles from the volcanic centre, and
the suspension of dust in the air caused complete darkness over an area of several
thousand square miles. It is estimated that about five cubic miles of material was
ejected from Katmai. According to Fenner (1925), the pyroclastic material consists
largely of fragments of glass which he believes to have been cemented as a result of
cohesion due to heat. Marshall (19385) postulates a similar origin for the ignimbrites
which cover large areas in the North Island of New Zealand, and regarding their
origin he states ‘they are thought to have been deposited from immense clouds or
showers of intensely heated but generally minute tragments of volcanic magma’. In
comparing the volume of material ejected from Katmai with other great eruptions
that have taken place in historical time, Martin states that a similar volume of five
cubic miles was emitted from Krakatau, whilst the explosive eruption of Tomboro, an
island near Java, produced an ash shower variously estimated as representing 28:6 to
50 cubic miles of material. Capps (1916) has shown that a great eruption in the
Upper Yukon ejected 10 cubic miles of ash which was spread over a minimum area of
140,000 square miles, the thickness ranging from 300 feet near the volcanic centre to
about an inch at the edge of the deposit. He considers that wind had helped to carry
the ash a distance of some 450 miles.

The magnitude of these eruptions shows that the suggested pyroclastic origin of
the New South Wales Upper Ordovician slates, though speculative, is not impossible.

4. The Occurrence of Lavas and Ash in the Ordovician Sequence Hlsewhere.

As shown above, the great bentonite deposits of Tennessee, Kentucky and Alabama,
and even further west (Nelson, 1922; Twenhofel, 1926; Schuchert and Dunbar, 1933),
are ash beds, possibly rhyolitic ash now converted into leverrierite or some other clay-
mineral having a composition very different from the original material. The horizon
ot this material is in the Middle Ordovician near the base of the Upper Ordovician on
and below that of the Trenton limestone.

In Britain, rhyolites and rhyolitic ash make up a large part of the Welsh succession
(Lake and Reynolds, 1912; Gregory and Barrett, 1931; Elles, 1940) and in the English
Lake district (Bailey and Weir, 1939; Stamp, 1923; Hutchings, 1892) ash slates, tuifs
and andesites occur in the Bala Series which make up the Upper Ordovician in that
part of Britain.

168 PETROLOGICAL STUDIES IN THE ORDOVICIAN OF N.S.W. III,

When such outbursts of volcanic activity characterized the Ordovician in other
parts of the world, it would be strange if Australia should have been exempt from
such activity.

IV. ConpirioNs NECESSARY FOR THE ACCUMULATION oF BLACK SHALES.

Writing on the association of graptolites with the black shales, Ruedemann (1934)
points out that “no other fossil fauna is so highly restricted to a single definite rock
facies’, and as this rock facies is the subject of the present paper, it seems pertinent
to examine the various hypotheses put forward to account for the accumulation of
these black sediments with their characteristic fauna.

1. Earlier Hypotheses.

It is generally agreed that the black carbonaceous material of the shales has been
derived trom the decay of masses of seaweed ta which the graptolites were either
attached or associated, and the Palaeozoic plankton has been compared to that found
in the modern sargasso-seas, but there is no consensus of opinion as to whether the
plankton is capable of sinking to the bottom of the ccean or whether it dies and
decays on the surface or is carried either alive or dead into the quiet embayments
and shallower seas as the result of storms.

On the assumption that the shales and fine muds are deep-water deposits and
that the formation of carbonaceous deposits requires reducing as opposed to oxidizing
conditions, most of the hypotheses relating to black shale accumulation demand that
deep, tranquil water is essential.

According to Marr (1925) the graptolite-shales of Europe were deposited in quiet
embayments where the bottom was fouled by the presence of hydrogen sulphide and
no life could exist. In a diagrammatic section, Marr shows that the benthonic forms
develop in the littoral zone of the embayments where there is sufficient oxygen to
maintain life, and the only fossils accumulated in the poisonous muds are the dead
planktonic or pseudo-planktonic forms which sink from the surtiace. .

This accounts for the pure graptolite fauna in which the benthonic forms are not
associated, but Ruedemann suggests “occasional and incomplete incursions, or overflows,
from a geosyncline” to account for the mixed faunas, such as that of the Utica shales
of North America, which pass into limestones.

Clarke (1903) suggested that the black shales accumulated in deep almost stagnant.
seas, such as the present Black Sea, where there was only slight: vertical circulation
between two layers of different salinity, and Schuchert (1910) believes that they form
in stagnant culs-de-sac or infill holes in the sea floor.

Neither of these theories accounts for the occasional mixed faunas or for the
conglomerates and sandstones which are sometimes found interbedded with the black
shales, and both Ulrich (1911) and Ruedemann (1897, 1911, 1934) have pointed out
that the world-wide distribution of the graptolites and their dependence upon ocean
currents for their. distribution negative any theory requiring stagnant conditions,
and furthermore, they discredit hypotheses requiring that the shales be deposited in
narrow embayments or culs-de-sac, for they point out that the marine currents could
not carry the plankton to the head of such a bay. It would, however, be possible for
wind to carry the plankton into the bay, but the world-wide distribution of this fauna
suggests that their preservation in such bays would be too limiting a condition.
Ruedemann (1911) concludes that the world-wide distribution of some species shows
that the Palaeozoic oceans were connected to permit intercommunication but sufficiently
separated to permit the development of provincial characters. Ulrich and Ruedemann
both consider that the occurrence of black, graptolite-bearing shale in the Levis,
Athens and Ouachita troughs, which are narrow strips hundreds of miles in length,
indicates that the shales were laid down in geosynclines open to the ocean at both
ends. As mentioned above, Ruedemann accounts for the mixed faunas and the
occasional admixture of littoral sediments by postulating an occasional break through
of the barrier and the formation of epicontinental seas. He states, however, that

BY GERMAINE A. JOPLIN. 169

“marine faunas are not found generally distributed through the mass of the black
shales. They occur in occasional thin seams in which, however, their remains are
likely to be very numerous, and the best of these—indeed it may be the only zone
of such fossils in hundreds of feet of shales—is usually in the basal foot or two’.
He thinks that “not depth but tranquillity” is the main factor and that deposition
probably takes place between the agitated water and the currentless sea. He suggests
that the waters may have been stagnant or the circulation imperfect over long periods
and that the marine life may have become extinct by fouling of the water.

Later, Ruedemann (1944) states, with reference to the shales of the Hudson Valley
Belt, ‘the graptolite shales are black, very fine-grained sediments which were deposited
at the lower slopes of continental shelves or at the bottom of the abysses, some at
12,000 + feet’. He supports this hypothesis as to their deep origin by pointing out
that radiolarian cherts may be found associated with black shales (Ruedemann and
Wilson, 1936) and that the radiolarian genera are very deep-water forms.

Grabau and O’Connell (1917) claim that the black shales are deposited in the
lagoons and bays of deltas where the plankton has been washed by exceptionally high
tides and storms. They review the two classical areas for graptolite-bearing shales,
namely, the Swedish deposits and those of Moffatdale in Scotland, and show that both
indicate a progressive overlap accompanying a positive movement of the strand-line.
They believe that the sediment is mainly of terrestrial rather than of marine origin
and that the holo- or epi-plankton has been buried in the deltas or lagoons. This theory
has not gained general acceptance, for though it accounts for shales of shallow water
origin, it is difficult to believe that storms and exceptional tides could account for
this widely distributed and common type of sediment.

Twenhofel (1915) suggests that the deposits may be connected with low tempera-

tures where the decay of the plankton is retarded and where the conditions are too
cold for the growth of the benthonic forms. Twenhofel wrote in 1915 “much has been
written relating to the origin of black shales, but judging from the divergence of
published opinion, no hypothesis has gained a general acceptance’. The same state-
ment can be made in 1945.
To gain general acceptance an hypothesis must account for the fine texture of the
shales, for the not infrequent intercalations with coarse-grained sediments, for the
occasional overlaps indicating shallow water deposition, for the elongated narrow strips
hundreds of miles in length indicating geosynclinal deposition, for the world-wide
distribution of some graptolites and the provincial development of others, for the
general absence of benthonic forms and the occasional mixed faunas, the benthos
usually being restricted to narrow seams often near the base of thick black shale
deposits, and finally for the carbon content and the excellent preservation of the
graptolites which indicates rapid burial and lack of oxidation.

2. The Volcanic Ash Hypothesis.

Studies in present-day mobile areas indicate that continental masses are often
fringed by island arcs on which active voleanoes are situated. At the.present time
such festoon islands occur on the western shores of the Pacific Ocean and cut off
comparatively tranquil seas from that ocean. On the eastern shores of the Pacific the
voleanoes are situated on the main continental mass and the festoon islands are absent.
Nelson pictured palaeogeographical conditions such as these in Ordovician times when
he stated that the bentonite was derived from a volcano situated on an island off a
peninsula separating the Lowville Sea from the Palaeozoic Atlantic. The great
eruptions of Tomboro and Krakatau, cited above, took place in such a mobile area and
both voleanoes are situated on islands. On the other hand, Katmai is situated on the
mainland of Alaska, and Fenner (1925) has been able to trace great rifts in the
Valley of Ten Thousand Smokes where much of the volcanic ash has accumulated.

As stated by Ulrich (1911) and Ruedemann (1911, 1934), the elongated masses of
black shale extending over hundreds of miles must have been laid down in a
geosyncline open at each end of the ocean to allow the ingress and egress of the

170 PETROLOGICAL STUDIES IN THE ORDOVICIAN OF N.S.W. III,
oceanic plankton, and it is to be expected that volcanoes would be situated along the
shores of such a great trough.

If large volumes of volcanic ash were suddenly poured on the masses of plankton
floating either in an island-fringed sea or in a geosyncline, they would founder and
sink. Their sudden burial provides the perfect conditions for the preservation of the
graptolites and for the slow decomposition of the associated seaweeds cut off from all
possibility of oxidation. Furthermore, if this load of ash and plankton chanced to fall
on any benthos life in the littoral zone of the sea or geosyncline, then the benthos
would immediately be killed and preserved as a seam at the base of the black
shale deposit. In the course of time the benthonic forms would probably invade the
shores again only to be killed off again by a further eruption from the same volcano
or from others situated further along the shore or fringing arc.

Hypotheses suggesting the shallow water origin of the graptolite-shales have to
depend upon severe storms and unusually high tides to sweep the plankton into the
shallow bays or deltas, but such deposits could be synchronous with the laying down
of the deeper water ash beds, for there is little doubt that the great volcanic outbursts
would be accompanied by tidal waves which would sweep the continental shores,
driving the plankton into the landlocked lagoons, where it would be immediately
buried by volcanic ash. Thus overlaps on coarser deposits and intercalations of
coarser sediments could be accounted for and formed either among delta deposits or
along the littoral zones. Volcanic ash could therefore form accumulations of the same
rock facies under apparently different environmental conditions.

In the case of the New South Wales rocks, it is suggested that the bulk of the
material is rhyolitic ash or dust, since rhyolites are known to occur in the Upper
Ordovician sequence at Cooma, and the composition of the siliceous slates appears
to conform to that of certain rhyolites and rhyolitic tuffs; but it must be emphasized
that rhyolitic material need not necessarily be the material that entombed the
graptolites and that this hypothesis might be applied to shales of very different
composition. Hence Twenhofel (1926) has shown that the bentonite is very different
in composition from the rhyolitic ash from which it was derived, and a difference in
composition could be caused either by subsequent alteration of the ash or by an
original difference in the type of volcanic material ejected. Actually volcanoes emitting
rhyolite are more often of the explosive type, and the acid type of material is more
often associated with terrestrial volcanoes.

V. SUMMARY.

It has been shown that the graptolite-bearing slates of the Upper Ordovician in
New South Wales are highly siliceous and that their siliceous nature is probably original.
It is suggested that they may have been formed as the result of large accumulations of
voleanic ash which encased the plankton and prevented oxidation of the carbon content.

This hypothesis for the origin of the graptolite-bearing black shales is considered
in the light of other hypotheses and of the necessity to account for all the observed
facts concerning black shale accumulation.

VI. ACKNOWLEDGEMENTS.

I am indebted to Mrs. Sherrard, Dr. Ida Brown, Dr. W. R. Browne and Mr. D. G.
Moye for kindly supplying me with material for this investigation. I should also like
to thank Professor L. A. Cotton for helpful discussion, Dr. W. R. Browne for discussion
and for his critical reading of the manuscript, and Professor E. S. Hills, of Melbourne
University, for kindly sending me a list of analysed Ordovician slates from Victoria.

VII. REFERENCES.
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————, 1898.—The Occurence of Graptolites in the Peak Hill District. Ibid., iv: 183.

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, 1909.—Notes on a Collection of Graptolites from Tallong, N.S.W. Ibid., 8: 339.

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, 1887.—Notes on the Area of Intrusive Rocks at Dargo. Ibid., 23: 127-164.
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HUTCHINGS, W. M., 1892.—Ash-slates and Other Rocks of the Lake District. Geol. Mag. Lond.,
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JOPLIN, G. A., 1942.—Petrological Studies in the Ordovician of New South Wales. i. The Cooma
Complex. Proc. LINN. Soc. N.S.W., 67: 156-196. 3

, 1943.—Id., ii. The Northern Extension of the Cooma Complex. Ibid., 68: 116.

——— and CuLey, A. G., 1937.—The Geological Structure and Stratigraphy of the Molong-
Manildra District. J. Roy. Soc. N.S.W., 71: 271.

KEBLE, R. A., and BENSON, W. N., 1939.—Graptolites of Australia: Bibliography and History of
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and SHERRARD, K., 1938.—Geol. Surv. N.S.W., A. R. Dept. Mines: 138.

LAKE, P., and REYNOLDS, S. H., 1912.—The Geology of Mynydd-y-Gader. Quart. J. Geol. Soc.,
68: 345-362.

LLASERON, C. F., 1909.—Exhibit. Proc. Linn. Soc. N.S.W., 34: 118.

Marr, J. E., 1925.—Conditions of Deposition of the Stockdale Shales of the Lake District.
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MARSHALL, F., 1935.—Acid Rocks of the Taupo-Rotorua Volcanic District. Trans. and Proc.
Hoy. Soc. NeZ., 64 (3) 2 328. :

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of the Most Tremendous Volcanic Eruptions known in History. Nat. Geogr. Mag., 24:
131-181.

Nayior, G. F. K., 1935a.—Note on the Geology of the Goulburn District with Special Reference
to the Palaeozoic Stratigraphy. J. Roy. Soc. N.S.W., 69: 75-85.

—, 1935b.—The Palaeozoic Sediments near Bungonia. Ibid., 69: 123.
, 19387.—Preliminary Note on the Occurrence of Palaeozoic Strata near Taralga,
INESSWes) MDG lls 45.

UA PETROLOGICAL STUDIES IN THE ORDOVICIAN OF N.S.W. III.

Naytor, G. F. K., 1938.—Graptolites of the Goulburn District, N.S.W. Part i. Some Forms and
Localities. Ibid., 72: 129-135.

NELSON, W. A., 1922.—Volcanic Ash Bed in the Ordovician of Tennessee, Kentucky and Alabama,
Bull. Geol. Soc. Amer., 33: 605.

Pirsson, L. V., 1915.—The Microscopical Characters of Volcanic Tuffs—a Study for Students.
Amer. J. Sci., 40: 191.

Raceatr, H. G., 1934.—Geology of the Trunkey-Tuena District. Address to Geological Section.
J. Roy. Soc. N.S.W., 68: xiviii.

RUEDEMANN, R., 1897.—Evidence of Current Action in the Ordovician of New York. Amer.
Geol., 19: 367-391. ;

——__——.,, 1911.—Conference on the Faunal Criteria in Palaeozoic Palaeogeography. Strati-
graphic Significance of the Wide Distribution of Graptolites. Bull. Geol. Soc. Amer.,
BAAR. AGS ra, i

——__—-—., 1934.— Palaeozoic Plankton of North America. Geol. Soc. Amer., Mem. 2.

, 1944.—The Hudson Valley Belt of Graptolite Shales and Negative Anomalies of

Gravity. Amer. J. Set., 242: 391-396.

and WILSON, T. Y., 1936.—Eastern New York Ordovician Cherts. Bull. Geol. Soe.
Amer., 47: 1535-1566.

ScHUCHERT, C., 1910.—Biologic Principles of Palaeogeography. Pop. Sci. Monthly, 598-9.

and DuNBaR, C. O., 1933.—A Text-book of Geology. Part ii. Historical Geology. New

NiOnk apy Lbs:

SuHerraArD, K., 1936.—Exhibit: Proc. Linn. Soc. N.S.W., 61: 1.

—, 1939.—Tnhe General Geology of the District east of Yass, N.S.W. Ibid., 64: 577-600.

—_—_—_—, 1942.—Upper Ordovician Graptolite Horizons in the Yass-Jerrawa District, N.S.W.
J. Roy. Soc. N.S.W., 76: 252.

—_———. and Kmpie, R. A., 1937.—The Occurrence of Graptolites near Yass, N:S.W. Proc
LINN. Soc. N.S.W., 62: 303-314.

SmirH, H.°G., 1899.—Exhibit. J. Roy. Soc. N.S.W., 33: xli.

Stamp, L. D., 1923.—An Introduction to Stratigraphy. London, p. 69.

Tarram, C. M., 1929.—The Metamorphic Rocks of North-east Victoria. Geol. Surv. Vict.,
Bull. 52.

TWENHOFEL, W. H., 1915.—Notes on Black Shales in the Making. Amer. J. Sci., 40: 272-280.

. 1926.—Treatise on Sedimentation. Baltimore, p. 295.

UnricH, E. O., 1911.—Revision of the Palaeozoic Systems. Bull. Geol. Soc. Amer., 22: 358.

WASHINGTON, H., 1911.—Chemical Analyses. U.S. Geol. Swrv., Prof. Pap. 99.

WATERHOUSE, L. L., and BROWNE, W. R., 1929.—Note on an Occurrence of Quartzite containing
Common Opal and Chalcedony at Tallong, N.S.W. J. Roy. Soc. N.S.W.. 63: 143, 146.

WooLnoucH, W. G., 1909.—The General Geology of Marulan and. Tallong. Proc. Linn. Soc.
INSSAWie oi 4n 82.

173

ON THE FAMILY SMARIDIDAE (ACARINA).
By R. V. SoutHcort, M.B., B.S.
(Four Text-figures. )

[Read 25th July, 1945.]

In 1941 Womersley and Southcott revised and redescribed the Australian
Smarididae, and made some alterations to the classification of the family; leaving it
with four genera—Smaris, Fessonia, Hirstiosoma and Sphaerotarsus. They were able
to record all of these genera except Fessonia from Australia, and thus were able to
redefine the genera (except Fessonia).

Although it was considered at the time that the previous definition of Fessonia,
and description of Fessonia papillosa (by Vitzthum, 1929) as having three sensillary
areas to the crista was not likely to be correct, it was not then considered wise to
alter the accepted definition of the genus. The doubt has been confirmed by the
discovery of Fessonia australiensis, n. sp., from the Northern Territory of Australia;
and an examination of this reveals the reason for the mistake: the crista is continued
beyond the anterior sensillary area for some distance, onto the nasus, and there
expands to form an area resembling an area sensilligera (which is frequently found in
this position), but lacking any sensillary setae. This new species is described in the
present paper.

In 1927 André erected the genus Phanolophus for P. nasica, a new species from
Algeria.

Womersley and Southcott (1941) described the larva of Smaris prominens (Banks,
1916) as the first proven larval Smaridid, and stated that it differed from all described
larval Erythraeidae in having the lateral tarsal claws identical. An examination of
the figures given for Hauptmannia Oudemans, 1910 by Oudemans (1912) and an
examination of some Australian material, shows that this larval genus, previously
considered an Erythraeid, has the lateral tarsal claws identical (though differing from
Smaris in that these are simple, not pulvilliform). It is considered that Hauptmannia
is a larval Smaridid; and on grounds of its distribution, it-is most likely to be the
larva of Hirstiosoma.

The differences between the genera of the family are profound, and the opportunity
is taken here to divide the family into sub-families:

Family SMARIDIDAE Kramer, 1878.

Key to the Sub-families with Genera.

A. Eyes anterior to both sensillary areas. Sensillary areas both well behind nasus.
B. Dorsal and ventral shields present. Without crista. Eyes two on each side ..........
ee RR Tan ei Oh a reap Shey eco uis (Re tee side lavenatisne: tere sub-fam. Smaridinae, n. sub-fam.
With only one genus: Smaris Latreille, 1796.
BB. No dorsal or ventral shields. With a well-marked crista which is produced forwards
onto the nasus. Eyes two on each side, just anterior to anterior sensillary area
Ee SPUN Te ESSER arse Flin: ci sticci ete aloe eaey bee oeteatol Sone ghee teneie, Blades sub-fam. Fessoniinae, n. sub-fam.
With only one genus: Fessonia von Heyden, 1826.
AA, Eyes between the levels of the anterior and posterior sensillary areas. Anterior sensillary
area on or just behind nasus.
C. Eyes one on each side. Anterior sensillary area right on nasus............2.e200-
RO SUR TT Ach ny eens teu penn! Miiapimeeudusyslcehey aU tueteleve sub-fam. Hirstiosominae, n. sub-fam.
With two genera: Hirstiosoma Womersley, 1934, and Sphaerotarsus Womersley,
1936.
CC. Eyes two on each side, closer to the posterior sensillary area. Anterior sensillary
area a little behind tip of nasus. Crista prolonged behind posterior sensillary
VT ODMR COME RTS alle oi Pan. aa apeke avin Navana talreneneaanzdeiees sub-fam. Phanolophinae, n. sub-fam.
With only one genus: Phanalophus André, 1927.

174 ON THE FAMILY SMARIDIDAE,

F
Qo Dv
q o
me
ew
FEessoniAé EROTARSUS PHANOLOPHUS
HIRSTICSeMmA SPH fasren aban 1427)

SMARIS

Fig. 1—Characteys of genera of adults of Smarididae.

Os

850 ¢

S55
CO

SSS—
iS)

wf Te Gi
Qi as

ZMn (Ae
Uren (dé

i

&
3)

, (a sy aS,
“B Way
f aS j ten
- 33)

SOUTHCOTT e

Fig. 2.—Smaris prominens (Banks, 1916). A-H, Adult. A, Dorsum of male showing plates
and clear areas (muscular insertions) ; B, Internal and external genitalia of male superimposed
upon figure of posterior dorsal shield; C, External genitalia of male; D, External genitalia of

female; E, Female internal genitalia; F, Nymph, external genitalia.

BY R. V. SOUTHCOTT. 175

Genus. Smaris Latreille.
Précis car. gén. Ins., 1796, 180.

SMARIS PROMINENS (Banks, 1916). Figs. 1 A, B; 2 A-F; 3 A-F.
Fessonia prominens Banks, 1916, 225. Smaris prominens Wom. and Sthet., 1941, 63.

The opportunity is taken here of figuring the external genitalia of male and female
adults, and nymph, also some other features. In the lymph the genitalia are occluded
by a thin oval plate, which has a median raphe originating in the midline anteriorly,
extending back almost to the posterior end of the plate. Sexes cannot be differentiated
in nymphs, either on characters of genitalia or dorsal muscular plates (as they can be
in the adults).

[00K

SOUT HCOTT

Fig. 3.—Smaris prominens (Banks, 1916). A-B, Adult. A, Mouth-parts from below (dorsal
view of palp on ieft); B, Tip of palp; C, Larval skin attached to post-larval pupa, in which
the pigment-masses are shown; D-F, Nymph; D, E, Mouth-parts from above and below; F, Tip
of palp.

SMARIS MAMILLATUS (Say, 1821).

Hrythraeus mamillatus Say, 1821, 70. Smaris mamillatus Oudemans, 1937, 952.
Erythraeus mamillatus Jacot, 1938, 125.

Oudemans (1937, 952) repeats Say’s description and refers the species to Smaris
(s.l.). Jacot (1938, 125) suggests Say’s species was a Labidostomma. The “indented
punctures” of Say are considered by Oudemans as muscular insertions and these are a
feature of Smaris (s.str.), and a “marginal impressed line’ is also present in Smaris
(s.str.) (see Womersley and Southcott, 1941, 64, fig. 1A, B, of Smaris prominens
(Banks) ). Say’s “few distant hairs’ are possibly the posterior sensillary setae.
Accordingly Say’s species is referred to Smaris (s.str.).

176 ON THE FAMILY SMARIDIDAE,

Genus FrEssonra von Heyden, 1826.
Vers. syst. Einth., in Isis, xix (6), 609.

FESSONIA AUSTRALIENSIS, n. 8p. Figs. 1 C; 4 A-L.

@ Adult: Red, body oval, pointed anteriorly, slightly waisted around middle. Length
0-85 mm., width 0-46 mm. Crista linear, with two sensillary areas, the anterior being
well behind the nasus. Crista continues beyond anterior sensillary area onto nasus.
Hach sensillary area with two ciliated sensillary hairs, anterior 51 long, posterior 79y;
anterior sensillary area also with two of the normal dorsal setae 20u long. Distance
between centres of sensillae 1464; from centre of anterior sensillae to tip of nasus 215y.
Eyes two on each side, just anterior to anterior sensillae, anterior being the larger and
the more medial. Palpi as figured. Dorsal setae numerous, brown, dorsally convex,
there with 6-8 lines of serrations, which mostly run down to the pedicel of the seta,
ventral surface of dorsal setae flattened, and ciliated down the centre; setae 18-33u
long, some longer near the nasus to 37u. Ventral setae with long strong ciliations,
setae 18-24u long. Legs: I, 990u long; II, 700u; III, 6904; IV, 990u (all including coxae).
Tarsus I, 1381p long by 41yu high; metatarsus I, 2104 long; tarsus IV, 109 long by 35yu high;
metatarsus IV, 1934 long. Cuticle of legs including coxae porose. Various types of
sensory setae are present on the legs, as in Smaris.

SOUTHCOT T

Fig. 4.—Fessonia australiensis, n. sp. A-L, Adult. <A, Dorsal view, entire, female; B,
Anterior part of dorsum; C, Eyes and crista of teratological specimen; D, Tarsus I and meta-
tarsus I; H-J, Adult setae, with scale to left; E-G, Dorsal surfaces of dorsal setae; H-I,
Ventral surfaces of dorsal setae; J, Ventral seta; K-L, Palp, K from above, L below. (All
figures except C from type 2.)

BY R. V. SOUTHOOTT. irre

Habitat—Northern Territory: Mataranka, found along the banks of the Roper River,
- under bark of Melaleuca, 28-29.111.1943, four adults (R.V.S.). (Type @ and three para-
types in author’s collection).

One specimen with an abnormal anterior sensillary area to the crista is figured in
IMs, Zo (Op

The anterior prolongation of the crista onto the nasus does not form a sensillary
area, though superficially it has that appearance, and has misled previous workers.
There are no sensillae on the nasus, only some of the normal dorsal setae being present.

A detailed comparison of this species with the European IF. papillosa (Hermann,
1804) (the genotype) is not possible, as no good description of that species has been
published. It must be emphasized that these descriptions must include measurements,
and it is also imperative that a detailed drawing of the dorsal seta be included for
specific determinations of these mites (this also applies in other families) to be possible.

Genus HirstiosomA Womersley.
Rec. S. Aust. Mus., v (2), 1934, 242.

HIRSTIOSOMA JACOTI, New name.

Smaris sericea Jacot, 1938, 123. [non] Zrombidium sericeum Say, 1821, 70.
Hirstiosoma sericea Wom. and Sthet., 1941, 63.

Oudemans (1937, 1406) repeats Say’s original description of sericeum, and considers
it possibly a species of Hutrombidium (?). Say’s statement ‘‘densely covered with a
short silken hair’ cannot be applied to any Smaridid.

Jacot (1938, 122) suggests that Say had mixed up his descriptions of Trombidium
scabrum (1821) and Trombidium sericeum, and refers the former to his new genus
Trombiculoides, and the latter to Smaris (s.l.). Jacot also states (p. 121) that he had
caught “four specimens of a Microtrombidium” in Say’s collecting localities, but does
not describe them. The suggestion naturally arises that these are possibly Say’s
sericeum. It is best to follow Oudemans’ decision on sericeum, and therefore Jacot’s
Smaris sericea must be renamed.

Genus SPHAEROTARSUS Womersley.
J. Linn. Soc., Lond. (Zool.), xl, 1936, 119, 269.

SPHAEROTARSUS WOMERSLEYI, new name.

Caeculisoma ripicola (as nymph) Womersley, 1934, 239. [non] Caeculisoma ripicola
(adult) Womersley, 1934, 238. Sphaerotarsus allmani Womersley, 1936, (part), 40, 119,
269. Sphaerotarsus ripicolus Wom. and Sthet., 1941, 73.

The description of the nymph of this species followed the description of the adult
Caeculisoma ripicola Womersley, 1934, with which it was wrongly correlated (this adult
belongs to the Erythraeidae and is now the genotype of Callidosoma Womersley, 1936).
Accordingly the name ripicolus, being based on a mis-identification, cannot stand for
the nymph (Article 31) and is changed as above.

List of Smarididae.

The family as reclassified above will comprise the following species:

Smaridinae, n. sub-fam.: Smaris squamatum (Herm., 1804) Europe (genotype);
S. mamillatus (Say, 1821) North America; S. nicoletii (Gervais, 1849) South America;
S. depilata (Berlese, 1888) South America; S. prominens (Banks, 1916) Australia.
Fessoniinae, n. sub-fam.: Fessonia papillosa (Herm., 1804) Europe (genotype);
F. australiensis, n. sp., Australia. Hirstiosominae, n. sub-fam.: Hirstiosoma scalaris
Wom., 1934, Australia (genotype); H. novae-hollandiae Wom., 1936, New Zealand; H.
tasmaniensis Wom. and Sthet., 1941, Tasmania; H. ampulligera (Berlese, 1887), Europe;
H. jacoti, new name, North America; Sphaerotarsus allmani Wom., 1936, Australia
(genotype); S. womersleyi, new name, Australia; S. leptopilus Wom. and Sthet., 1941,
Australia; 8. claviger Wom. and Sthet., 1941, Australia. Phanolophinae, n. sub-fam.:
Phanolophus nasica André, 1927 (genotype), North Africa.

178 ON THE FAMILY SMARIDIDAE.

The following species have been referred to the genus Smaris (s.l.) by various
authors; but from their descriptions cannot at present be classified generically:
Smaridia villosa Dugés, 1834 (see Oudemans, 1937, 962); Rhyncholophus phloginus
Koch, 1837 (Oudemans, 1937, 957, 958); Smaridia vestita Gervais, 1849 (Oudemans, 961) ;
Smaridia ampulligera var. longipes Berlese, 1888, 173; Smaris longilinealis Ewing, 1909,
61.

REFERENCES.

ANDRE, M., 1927.—Bull. Soc. Hist. nat. Afr. N.. 18: 169.
Banks, N., 1916.—Trans. Roy. Soc. S. Aust., 40: 225.
BERLESEH, A., 1888.—Bull. Soc. ent. Ital., 20: 171-222.
Ewine, H. E., 1909.—Trans. Acad. Sci. St. Lowis, 18: 53-77.
*______1910.—Univ. Ill. Bull. 7, 14, 120 pp. (Part cf University Studies, Vol. 3.)
Jacot, A. P., 1938.—Psyche, 45 (2-3): 121-132.
OUDEMANS, A. C., 1912.—Zool. Jahrb., Suppl. 14, Heft. 1.
, 1936-1937.—Krit. Hist. Overzg. d. Ac., iii A, 1936; iii C, 1937, 952-962; iii G, 1937.
*Say, T., 1821.—J. Acad. Nat. Sci. Philad., 2. (1): 59-82.
VITzTHUM, H., 1929.—Tierwelt Mitteleuropas, 3 (7): 69.
WoMERSLEY, H., 1934.—Rec. S. Aust. Mus., 5 (2): 238-9.
, 1936.—J. Linn. Soc. Lond. (Zool.), 40: 119, 269.
, and SoutHcotTt, R. V., 1941.—Trans. Roy. Soc. S. Aust. 65 (1): 61-78.

* References not seen by author.

Proc. Linn. Soc. N.S.W., 1945. PLATE VI.

(Kitzinger).

TeEsvensis

Geographic Variation in Hemiergis dec

PLATE VII.

Proc. Linn. Soc. N.S.W., 1945.

Microspore-Types in the Permian of N.S.W.

xt.

cs

Sa

179

THE HAIR TRACTS IN MARSUPIALS.
PART II. DESCRIPTION OF SPECIES, CONTINUED.*

By W. BOARDMAN.
(Department of Biology, The University of Queensland, Brisbane.)

(Plate viii; forty-three Text-figures.)

[Read 26th September, 1945.]

In addition to the further species dealt with in the present contribution,
supplementary notes, mostly based on fresh material, are recorded of species previously
described.

ACKNOWLEDGEMENTS.

All but a few of the specimens examined belong to the collection of the National
Museum, Melbourne. The late Mr. D. J. Mahony, when Director of the Museum, very
generously made them available for study. I am indebted to Mr. C. W. Brazenor,
Mammalogist at the same institution, for much help on questions of nomenclature.
My thanks are tendered also to Dr. A. B. Walkom, Director of the Australian Museum,
Sydney, and to Dr. J. Pearson, Director of the Tasmanian Museum, Hobart, for
permission to describe several species from the collections under their charge.

The drawings are the work of Mrs. Dorothy Reid.

Registration numbers prefixed with the letters “R” or ‘“‘C” refer to specimens from
the National Museum Collection. The repository of the remaining material is specifically
indicated in the text.

Suborder POLYPROTODONTIA.
Family DASYURIDAE.
Subfamily PHASCOGALINAE.
DASYCERCUS CRISTICAUDA Krefft. Figs. 1-8.

Material.—R.12517-9—three females, and R.12512, R.12521-2, R.12524,
R.12528—five males (length of head and body 56-77 mm.); Charlotte Waters,
Central Australia.

Wood Jones (19230) says of this species: ‘‘The arrangement of the hair upon the
whole of the head, body, and limbs is of basal simplicity. The face, head, and the
whole of the trunk and tail are clothed with hair having a uniform caudad trend.
Upon the limbs the hair is uniformly in a distal and post-axial direction. There are
no reversals, partings, convergences, or vortices anywhere.” This cannot be said of
the series before me which shows some departures from the primitive arrangement.

Head.—On the head dorsally, and laterally above the level of the genal papilla, the
hair-flow is mostly caudalwards; between the eye and the ear the current bends
upwards so that there the flow is upwards and caudally. Beneath this level on the
side of the face, from immediately behind the mystacial zone, the direction is caudal-
wards and ventrally along an are which is of shorter radius behind the genal papilla
and thus, in a restricted field, the flow is towards the mid-ventral line and at right
angles to it; about midway between the genal papilla and the mid-ventral line the flow

* The observations embodied in this paper were made during the tenure of an appointment
at the Institute of Anatomy, Canberra, and are published with the permission of the Common-
wealth Director-General of Health.

At

180 THE HAIR TRACTS IN MARSUPIALS. II,

again resumes a caudal and ventralwards trend so that, in all, the current follows a
somewhat S-shaped course. Between the mandibles, except in a narrow mid-ventral
strip, the hair proceeds caudomedially to coalesce without any definable boundary with
the current from the side of the face.

ed

Figs. 1-3.—Dasycercus cristicauda. 1. Ventro-lateral view of head and neck (R.12519).
2. Lateral view of head and neck (R.12518). 3. Constituent currents of the prescrotal triangle
(R.12512). The bilaterally paired convergent intervals are exaggerated for purpose of
illustration.

Neck.—The stream flowing caudalwards over the crown of the head is continued
without interruption along the dorsum of the neck and trunk. On the side of the neck
the trend is towards the mid-ventral line with varying degrees of obliquity—
caudoventrally where it runs on to the lateral aspect of the upper arm over the shoulder
and the root of the neck, at right angles to the long axis at about the middle of the
neck, cranioventrally in front of this; round the base of the auricle a recurved
course is followed. The recurved stream behind the base of the auricle encounters
below the stream from the side of the face and where these two opposing currents meet
a divergent interval is formed which lies immediately beneath and somewhat in
front of the ventral limit of the base of the ear. A mid-ventral convergent line is
present on the neck; it extends from about opposite the angle of the mandible along
the neck and on to the cranial fourth of the thorax.

Trunk.—The primitive arrangement holds throughout except for the presence in
the male of a prescrotal reversed area (v. infra). On the ventral trunk the hair flows
caudally and medially but the medial component is too weak to produce a mid-ventral
line of convergence. The pouch area does not interrupt the direction of hair-flow.

The prescrotal reversed area of the male is similar to that recorded for Antechinus
maculatus (Boardman, 19430). Its relationship to the scrotal stalk and the currents
entering into its formation are detailed in Fig. 3. This reversal is observable satis-
factorily only on specimens such as R.12512 in which the fur is well grown. A
contribution to the tuft-like convergent centre at the apex of the triangle is made by
a sharp recurving of the hairs forming its sides.

Limbs.—The primitive arrangement prevails except on the forearm where, proximal
of the ulnar-carpal vibrissae, the hair on the lateral and medial aspects of the limb
bends back to produce a reversed stream which flows along its postaxial margin to
end in a convergent centre just distal of the elbow. In the axilla a divergent interval
is formed similar to that figured for Cercartetus nanus unicolor (Fig. 14).

Remarks.—The close resemblance of this species to Antechinus maculatus should
be emphasized. The only noteworthy differences are the presence of a forearm reversal
in Dasycercus cristicauda, and a relative weakness in the recurved stream behind the
base of the ear in Antechinus maculatus which has prevented the formation of a clearly
defined divergent interval.

— eo eS

BY W. BOARDMAN. 181

DASYUROIDES BYRNEI Spencer.
Material. C.465-7—a male and two females (length of head and body
57-75 mm.); Central Australia.
No significant differences from Dasycercus cristicauda could be defined.

SMINTHOPSIS CRASSICAUDATA CENTRALIS Thomas.
Material.—R.12461—two specimens, a male and a female, probably litter
mates (crown-rump length about 24 mm.); Charlotte Waters, Central Australia.
Like the subspecies macrura (Boardman, 194380), centralis shows no differences
which are detectable (on the developmental stage available) from Antechinus maculatus.
The hair of the inguinal region is weakly developed, but the prescrotal reversed triangle
can readily be charted.

Subfamily DASYURINAE.
DASYURINUS GEOFFROII GEOFFROIL Gould.
Material—cC.476—-8—three litter mates comprising a male and two females
(crown-rump length about 59 mm.); Central Australia.

The principal difference between the subspecies geoffroii and fortis (v. infra) occurs
in the females. In the female of geoffroii the mid-ventral inguinal convergent interval
lies immediately in front of the cranial limit of the pouch (ef. Fig. 6). The males of
the two subspecies are similar in every respect.

DASYURINUS GEOFFROII FORTIS Thomas. Figs. 4-6.
Material—R.1931—2—a male and a female (length of head and body 117
mm.); Western Australia; October, 1875.
Head.—In this animal the head corresponds point for point with the condition
described above for Dasycercus cristicauda, except that the divergent interval below
the base of the ear is not so precisely defined.

AN
Li \\

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.

=

SS
W
EZ

N
EZ MEN AW AWAY 5

y

SN
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SS

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ic

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Figs. 4-6.—Dasyurinus geoffroii fortis. 4. The male from the ventral aspect. 5. Ventro-
lateral view of head, neck and upper thorax. 6. Abdomen of the female viewed ventrally.

182 THE HAIR TRACTS IN MARSUPIALS. II,

Neck.—The flow of hair on the dorsum, sides, and cranial two-thirds of the ventral
aspect of the neck is very much the same as described for Dasycercus cristicauda. On
the ventral surface in its caudal third, however, there is some difference due to
disturbance on the upper chest; this area will be referred to below.

Trunk.—A divergent centre is situated within the axilla and from it hair flows
laterally on to the medial aspect of the forearm, medially by a curved course (concave
caudally) on to the ventral chest, and in front towards and across the cranial axillary
fold and so by recurvature on to the lateral aspect of the upper arm. A weak feathering,
which takes its origin in the divergent centre, runs forward, inclined towards the
medial line, to about the middle of the neck; medially from it hair flows along a curved
course towards the mid-ventral line, laterally by a similar curved course over the root
of the neck on to the shoulder and front of the arm where it merges with the general
caudalwards current of the side of the body. The details of the relationships of these
currents are illustrated in Figs. 4 and 5.

On the dorsal and lateral aspects of the trunk the hair-flow is primitive. In both
the male and the female the current over the flanks curves round sharply towards the
mid-ventral line. In the male the bending round becomes a recurvature which results
in the formation of a convergent centre about midway between the attachments of the
limbs; a convergent centre is not formed in the female, but instead there is a distinct
hair-ridge which stops short caudally at a convergent interval in front of the pouch
area. The tracts of the inguinal region are primitive in both sexes; a prescrotal
triangle is present in the male.

Limbs.—A forearm reversal occurs as in Dasycercus cristicauda but with the
convergent centre more distally placed.

DASYUROPS MACULATUS Kerr.
Material.—C.441-3—two males and a female, probably litter mates (length
of head and body 102 mm.); Western Australia; purch. W. Groener, 30th April,
1880.
The male is similar in every way to Dasyurinus geoffroii fortis. The female shows
no differences from the female of the subspecies Dasyurinus geoffroii geoffroit.

SARCOPHILUS HARRISII Boitard. Figs. 7 and 8.

Material.—R.5459-5462—a group of four females (length of head and body
in each case 135 mm.); Wynyard, Tasmania; 20th July, 1914.
The general features of the disposition of the hair tracts in this species have already
been described (Boardman, 1943b). The present series of four females is ideal for
figuring and enables the previous description to be amplified by the provision of a
diagram of the hair arrangement of the ventral abdomen and in the gular region and
axilla. It will be noticed that the mid-ventral convergent centre on the trunk is
whorled in a clockwise direction. A re-examination of the male (A.6438, Australian
Museum Collection) shows that the scrotal stalk has the same relations to the inguinal

convergent interval as has the pouch in the female.

Family PERAMELIDAE.
ISOODON OBESULUS OBESULUS Shaw and Nodder. Fig. 9.
Material—C.451—a female (length of head and body 111 mm.); Central
Australia.

The features shown by the subspecies obesulus differ from fusciventer* only in
detail in the zone where the forward flow from the gular whorl system meets the
backward flow from the chin. This will be clear from a comparison of Figs. 9 and 10.
The gular whorl is single, on the right side, and counter-clockwise; its influence
cranially ceases immediately behind the interramal papilla.

* Specimens of Isoodon obesulus described (Boardman, 1943b) without subspecific designa-
tion are, by virtue of their locality, almost certainly the subspecies fusciventer. The arrange-
ment of hair on the throat between the mandibles is the same as that recorded and figured for
Perameles gunnii (this communication, Fig. 19).

BY W. BOARDMAN. 183

<=

—
Ww

3

ree
ae

VEE

QE

Ss
Ss
ee

©
SS

Figs. 7 and 8.—Sarcophilus harrisii. 7. The female abdomen from the ventral aspect.
8. Ventro-lateral view of the head, neck and thorax.
Fig. 9.—Isoodon obesulus obesulus. The gular field.

PERAMELES GUNNII Gray. Figs. 10-13.

Material—C.589—a female (length of head and body 133 mm.); locality
unknown; pres. Zoology and Acclimatization Society of Victoria, 1st June,
1872. C.590—a male (length of head and body 123 mm.); Victoria; coll.
D. Le Souef. R.12634—a femaie (length of head and body 128 mm.); Victoria;
15th November, 1920. R.13042—a female (length of head and body 147 mm.);
Kew, Victoria; 14th September, 1886.

Head and Neck.—Of the three females examined two have a pair of gular whorls
as in Perameles nasuta (Boardman, 1948a), one a single clockwise whorl on the left
side like the male of Jsoodon obesulus fusciventer (see footnote, p. 182); the arrangement
of hair currents between the mandibles is the same for both the females and the male.
The male presents so many abnormal features that it is considered separately below.

Trunk.—The general arrangement of the trunk currents is as described and figured
for Isoodon obesulus fusciventer (Boardman, 1943), Fig. 9) but a complication is
introduced in that the inguinal reversal has its origin in a pair of symmetrically
disposed whorls, the left member being clockwise, the right counter-clockwise. In the
female the centres of the whorled system lie about level with the cranial limit of the
pouch and only a short distance from the mid-ventral line. In the male the centres are
relatively more widely separated and so much further cranially that they lie medial
of the knee.

Limbs.—None of the specimens shows any signs of forearm reversal. On the
contrary there is along the postaxial margin a well-marked flow running distally.

Abnormality—The single male (C.590) of the series has bilateral gular whorls,
somewhat nearer to the medial line than usual, but otherwise normal. In addition,
there are irregularities in the disposition of the currents on the head, neck and thorax.

184 THE HAIR TRACTS IN MARSUPIALS. II,

A supernumerary clockwise whorl is present on the left side beneath the base of the
ear. There is a small counter-clockwise whorl on the dorsal neck immediately behind
the occiput and to the left of the middle line; an imperfect clockwise whorl occurs
dorsally on the thorax, to the left of the middle line and approximately opposite to the
middle of the attachment of the fore-limb; a further clockwise whorl is present on
the right side just behind the shoulder. These anomalous whorls and the interplay
of the territories associated with them are charted in Figs. 10 and 11. An additional
imperfect clockwise whorl (not shown in the figures) is formed laterally on the right
side of the thorax behind and somewhat above the level of the axillary fold; its field
of disturbance is very restricted.

EL) WSN
Ve S

Ze SSS

i iL ESD

AC
eS)

zd

— =<

~~

AT
a

SS
NV

Ws

4)

Sem
M(
NX

v

—.

We

Sa

—

-_—

Figs. 10-13.—Perameles gunnii. 10. Face and gular field of C.590.° 11. Dorsal view of head,
neck and thorax of C.590. 12. The ventral abdomen in the male. 13. The ventral abdomen in
the female.

MACROTIS MINOR MINOR Spencer.
Material—C.593—a female (length of head and body about 86 mm.);
Charlotte Waters, Central Australia; pres. W. B. Spencer, 23rd April, 1896.
R.12431—a male (length of head and body about 82 mm.), R.12432—a female
(length of head and body about 86 mm.); same locality; September, 1895.
Unfortunately these specimens have not the hair sufficiently developed for a full
account of the hair tracts to be given. Each of the females has a single clockwise

BY W. BOARDMAN. 185

gular whorl on the left side and a reversal of the usual type on the postaxial margin
of the forearm. A gular whorl and forearm reversal are absent in the male. The
disposition of the tracts in the inguinal region of the female is similar to the account
given of Perameles myosura notina (Boardman, 19430). In the male the inguinal
tracts are too obscure for comment.

Suborder DIPROTODONTIA.
Family PHALANGERIDAE.
Subfamily PHALANGERINAE.
ACROBATES PYGMAEUS Shaw.

Material.—R.8139—two males and a female, litter mates (crown-rump length
about 33 mm.); Meenyan, Victoria; February, 1895. R.12685—a male and a
female, litter mates (crown-rump length 40 mm.); Victoria; 14th May, 1906.

The primitive nature of the hair currents as recorded earlier (Boardman, 1943b)
is confirmed. The flying membrane, at this stage rudimentary, does not interfere with
the primitive ventro-caudal flow over the side of the body between the attachments
of the limbs.

The scrotum, which is flattened transversely, is attached to the body not by a
narrow stalk but by a broad base as wide as the scrotum’s greatest width. In front of
the sessile scrotum the caudally and medially flowing hairs on the ventral abdomen
stop short, leaving an almost naked triangular patch having a base as wide as the
scrotal base and a height about half the length of the base; the triangle constitutes
an area of reversal and is formed in the same manner as in the Phascogalinae.

CERCARTETUS NANUS NANUS Desmarest.

Material—R.13003—a male and two females, litter mates (crown-rump
length 26 mm.), R.13012—two males and two females, litter mates (crown-
rump length 34 mm.); Tasmania; 20th September, 1872.

The subspecies nanus is similar in every way to unicolor (v. infra).

CERCARTETUS NANUS UNICOLOR Krefft. Fig. 14.

Material.—_C.793-4—two males, litter mates (crown-rump length 33 mm.);
Mordialloc, Victoria; pres. A. Campbell, 5th October, 1887. C.791—three females,
litter mates (crown-rump length 30 mm.); Mallee, Victoria; pres. A. Mattingly,
28th September, 1910.

Head.—On the dorsum of the head and neck the hair current sweeps caudalwards
on to the back. The lateral aspect of the head shows a considerable disturbance of
the normal backward hair-flow which is centred round a quadrilateral divergent interval
lying immediately behind the genal papilla. A strong preauricular current sweeps
upwards and recurves round the base of the ear so that it is completely encircled.
Where the current sweeps round the base of the ear below and encounters the normal
caudalwards flow on the side of the face beneath the eye, the divergent interval referred
to above is formed. As a consequence, the stream on the side of the face above the
genal papilla bends so that its course is upwards and caudally and merges with the
preauricular stream; the hair deflection below the divergent interval results in a curved
flow towards the mid-ventral line. In the vicinity of the chin the hair flow is
caudalwards; further back towards the angle of the mouth the flow gets progressively
more and more towards the mid-ventral line and behind merges with the currents
sweeping round from the side of the face.

Neck.—Caudal of the recurvature round the base of the ear the arrangement of the
currents on the neck is similar to that described above for Dasycercus cristicauda.

Trunk.—The primitive arrangement prevails except in the axilla where a divergent
interval occurs just within its caudal fold, that is, opposite to the elbow. From this
divergent interval hair flows distally along the forearm and medially and caudalwards
on to the chest.

186 THE HAIR TRACTS IN MARSUPIALS. II,

It is worthy of record that the scrotum is similar to that described in Acrobates
pygmaeus and is similarly associated with a prescrotal reversed triangle.

Limbs.—The hair currents on the limbs are entirely primitive.

16

Fig. 14.—Cercartetus nanus unicolor. Head, neck and upper thorax in ventro-lateral view.
Fig. 15.—Petaurus breviceps. Head, neck and upper thorax in ventro-lateral view.

Fig. 16.—Petaurus papuanus. Lateral view of head and neck.

Fig. 17.—Pseudocheirus laniginosus. Lateral view of head and neck.

PETAURUS AUSTRALIS Shaw and Nodder.

Material—A male and a female (length of head and body 100 mm. and
96 mm. respectively); no data (Australian Institute of Anatomy Collection).

This species shows no differences from Petaurus breviceps (v. infra).

PETAURUS NORFOLCENSIS Kerr.

Material—M.3783 and M.3785—two males (crown-rump length about 57
mm.); Cooroomon, Rockhampton district, Queensland; pres. H. C. McCartney
(Australian Museum Collection).

The hair on these two specimens is not sufficiently developed for detailed charting
except on the head, where it corresponds point for point with the account given below
for Petaurus breviceps. On the rest of the body comparison with Petaurus breviceps
shows that it is unlikely that any major differences from that species are present.

PETAURUS BREVICEPS Waterhouse. Fig. 15.

Material—R.12536—7—two males (length of head and body 90 mm.); Cape
York, Queensland; June, 1897.

Head.—The hair tracts on the head are practically identical with those described
above for Cercartetus nanus. Minor differences are mostly due to a single factor—the
greater field of influence of the preauricular stream which, in this species, sweeps
forward beyond and above the lateral angle of the eye and forms with the caudally

BY W. BOARDMAN. 187

flowing current of the dorsum of the head a short hair-ridge with a divergent interval
at about its middle. Consequent upon the presence of a convergent interval on the
throat (v. infra) the currents on the ventral aspect of the head converge on a point in
the mid-ventral line and just caudal of the level of the angle of the mouth.

Neck.—Generally, the neck is reminiscent of that described and figured above for
Cercartetus nanus but differs in the presence of a convergent interval midventrally at
about its middle.

Trunk and Limbs.—The hair on the trunk follows the pattern described for
Schoinobates volans (Boardman, 1943b and Fig. 18 below). Little doubt exists that in
specimens with the hair more advanced a triangular prescrotal reversal would be
detectable.

PETAURUS PAPUANUS Thomas. Fig. 16.

Material—_M.6379-80—two males (crown-rump length about 55 mm.);
Bulolo, Morobe Division, Territory of New Guinea; pres. Dr. C. E. M. Gunther
(Austraiian Museum Collection).

The species agrees with the account of P. breviceps given above except in the
vicinity of the lateral angle of the eye. This difference will be apparent from a
comparison of Figures 15 and 16.

PSEUDOCHEIRUS LANIGINOSUS Gould. Fig. 17.

Material.—C.599—a male (crown-rump length 75 mm.); Gippsland, Victoria;
pres. K. Flatow, 1st November, 1886.

Head and Neck.—The condition of the hair tracts on the head and neck is
reminiscent of what has been described above in Petaurus breviceps. The presence on
the dorsum of the head, above and somewhat in front of the ears, of bilaterally paired
centrifugal coils immediately distinguishes the genus (see also P. convolutor below).
The relationship of the coil to the hair-ridge into the formation of which it enters and
which runs between it and the base of the ear is Shown in Fig. 17; there is a convergent
interval on the ridge at its middle.

Trunk.—The trunk currents are primitive in their arrangement. Midventrally a
convergent line is present between the inguinal folds and thighs in front of the scrotum.
A naked triangle which probably signifies the presence of a prescrotal reversal occurs
in the usual position in front of the scrotum.

Limbs.—No interruptions of the primitive disposition occur.

PSEUDOCHEIRUS CONVOLUTOR CONVOLUTOR Oken. Plate viii.

Material.—No. 11, Pearson Collection (Tasmanian Museum)—a male (crown-
rump length 74 mm.); no data.

The growth of hair in this specimen is somewhat short of ideal requirements but
the principal features are well defined.

Head and Neck.—The condition of the hair tracts on the head is very similar in
general to that described and figured for P. laniginosus (v. supra). The divergent
interval on the side of the face lies immediately behind the genal papilla. The recurved
current at the back of the base of the ear bends sharply round it ventrally, and where
the current encounters the lower part of the preauricular reversal, a short forwardly
directed hair-ridge is formed. Bilaterally paired centrifugal coils occur as in P.
laniginosus and have the same relationships to the ear base and associated hair-ridge.

The same arrangement as in Dactylopsila picata (Boardman, 19436) is found
immediately behind the rhinarium, but in this case the reversed area is more definite
and there is on the mid-dorsal line behind it a potential convergent interval caused
by the bending backwards of the hairs behind the level of the middle of the dorsal
border of the mystacial zone.

On the dorsum of the neck the hair-flow is caudalwards; laterally it follows the
plan which is general in the group when a recurved current runs round the back of

188 THE HAIR TRACTS IN MARSUPIALS. II,

the base of the ear. There is a convergent interval midventrally on the neck as figured
for P. laniginosus.

Trunk and Limbs.—No departures from the primitive arrangement are present. The
hair on the flanks is sparse, flows ventrally and caudally, but there is no mid-ventral
line of convergence developed at this stage. A prescrotal reversed triangle occurs in
association with a male pouch (Plate viii).

There is no axillary reversal; a divergence of the caudally flowing stream on the
ventral chest flows across the axilla and so on to the medial aspect of the arm.

Ze Wf N
if Hy" \S
ZAIN
Jay | ae
SN CZF
YW ZEN

a

LZ

in ean

ee
SY GEE
LU Bigs fig

zi \f USES
\
EZ

Fig. 18.—Schoinobates volans. Dorsal view of entire animal with flying membranes spread
out.

SCHOINOBATES VOLANS Kerr. Fig. 18.

“The previous account of this series (Boardman, 19430) is augmented by the
provision of a figure of the dorsal surface of the body. The drawing is from specimen
M.3036 (Australian Museum Collection).

TRICHOSURUS VULPECULA VULPECULA Kerr. Figs. 19 and 20.

Material.—A male (crown-rump length 101 mm.); pres. W. Eldridge, 18th
February, 1944. A male (crown-rump length 99 mm.); pres. R. N. Wardle,
20th March, 1944. A male (crown-rump length 109 mm.); pres. W. Eldridge,

BY W. BOARDMAN. 189

15th July, 1944. These three males were all collected in Canberra, Australian
Capital Territory. In addition, there is a female (crown-rump length 90 mm.)
and a male (length of head and body 220 mm.), both from Canberra but having
no further data. The series is in the Australian Institute of Anatomy
Collection.

Wood Jones (1920) has already provided an account of the hair tracts in this
species under the name T'richosurus vulpecula var. typicus. The material used by him
included specimens from both the South Australian and the Western Australian
Museums. It would appear probable that not all of his material is referable to the
typical form the distribution of which is defined in Iredale and Troughton’s check-list
(1934). The following account, based on the 220 mm. male, is to be regarded as
supplementary to that of Wood Jones.

Cy

<(e>
Wes
PEM SRW

——

es
ae

SS

g
V
35

Y

(
@

SSS

Figs. 19 and 20.—Trichosurus vulpecula vulpecula.. 19. Lateral view of head and neck.
20. Ventral aspect of the entire animal. Both figures are based on the 220 mm. male which
apparently early in life lost the right hind-limb.

The specimen is, in the arrangement of its hair tracts, very similar to what has
been recorded by Wood Jones. Differences in detail, however, are sufficient to merit
the provision of a comparable figure of the face in side-view. It will be noticed that
there is no clear demarcation in the form of a hair-ridge between the areas marked
A and B by Wood Jones; also, there is no hair parting between the areas B and C.
The reversal in front of the eye which contributes largely to the territory B takes its
origin in the present series in what might be described as a divergent centre at the

190 THE HAIR TRACTS IN MARSUPIALS. II,

medial angle of the eye (cf. Wood Jones’ Fig. 2). Hairs streaming forward from this
centre encounter along the upper portion of the caudal margin of the mystacial zone
a stream flowing upwards and caudally, with the result that a hair-ridge is formed with
_a divergent interval at about the middle of its length. Immediately above the divergent
interval which results from the impact of the sub-ocular stream on the reversed
current which flows from behind the ear there is on each side a small whorl.

The radiating centre recorded by Wood Jones as situated between the ears is in
all of the present series a clockwise whorl.

Wood Jones makes no reference to the disposition of the hair tracts on the ventral
surface. What he refers to as “the general line of convergent streams (which) runs
downwards and forwards over the cheek and to the lower jaw near to its angle’ is
continued towards the mid-ventral line where the convergent stream line (in this
specimen it should rather be called a ridge) joins its fellow mid-ventrally just behind
the interramal papilla, the point of meeting being marked by a small clockwise
centripetal whorl. In the axilla a whorl is present on each side situated just within
the free border of the caudal fold; it is clockwise on the left, counter-clockwise on the
right. As a result of the hair reversal which originates in the axillary whorl, a ridge
including a divergent interval is formed on the medial aspect of the upper arm. On
the side of the body between the limbs the current flows caudally and ventrally; a
mid-ventral convergent point is present in the form of a tuft with a suggestion of
clockwise centripetal whorling. As is usual, a well-defined convergent interval occurs
in the mid-ventral line just in front of the scrotum; there is no prescrotal reversed
triangle. The mid-ventral line is indicated by a hair-ridge only behind the abdominal
tuft.

Variation._In none of the remaining specimens is the sub-ocular divergent
interval and its associated whorl present. Sometimes a distinct mid-ventral hair-ridge
is formed on the upper chest especially along the area occupied by the sternal gland.

TRICHOSURUS VULPECULA HYPOLEUCUS Wagner.

Material.—C.600—a female (crown-rump length 80 mm.); Claremont,
Western Australia; pres. H. J. Coles, 3rd October, 1911.

The hair is at a too early stage of growth for detailed charting except on the head.
The condition on the face is similar to that described for individuals of the subspecies
vulpecula (v. supra) in which the divergent interval beneath the eye is not present;
in this respect the specimen differs also from Wood Jones’ Western Australian Museum
example which formed the basis of his description and was most probably from a
Western Australian locality. A divergent centre occurs on the crown between the ears
as described by Wood Jones.

TRICHOSURUS FULIGINOSUS Ogilby.

Material.—No. 10, Pearson Collection (Tasmanian Museum)—a male (length
of head and body 225 mm.); Mt. Wellington, Tasmania; 17th March, 1943.
C.602—a male (crown-rump length 83 mm.); Tasmania; purch. Castleman, 1863.

The Tasmanian Museum male is at a very satisfactory stage for describing the
hair tracts. Its only difference from Trichosurus vulpecula vulpecula (sub-ocular
divergent interval absent) lies in the arrangement of the hair streams on the crown.
Two counter-clockwise whorls (the whorling is not very marked) are present, one
behind the other in the mid-dorsal line; the more cranial of the two is situated at the
vertex some little distance in front of the line joining the bases of the ears in front
and is produced into a feathering before and behind; the second is situated directly
between the bases of the ears and is associated with similar featherings. Midway
between the two whorls a divergent interval is produced where the opposing streams
meet. In so far as the disposition of hair streams in their vicinity is concerned
these two whorls have together the same effect as the single whorl or divergent centre
described elsewhere in the genus.

BY W. BOARDMAN. 191

A convergent ridge occurs along a line running obliquely across the back of the
thigh from behind the knee to the mid-ventral line which it reaches at a point about
two-thirds of the distance from the base of the scrotum to the base of the cloacal
hillock. It is formed by the stream on the lateral aspect of the thigh bending round
its postaxial margin and recurving proximally with respect to the limb before merging
with the stream on the medial aspect of the thigh.

Variation.—The hair is only weakly developed on the specimen C.602, but as far
as can be ascertained the tracts agree with those of the example just described except

FE

Vg
=
Ye VEX

fof

el

ZZ
Hc

(4
WZ \ (( Ose

<~

(7)
(=)

KS

ss

SS

ye)

Y

iy

Ws

\ \
yy

=U
»)

LL)

i,
ILI EGS

me
SY)

WN

ee
sy

GE
=

<_™

We
SSS
“4

igs

hs

Figs. 21-27.—Phascolarctos cinereus. 21. Lateral view of head and neck (C.433). 22. Head
and neck from above (C.433). 23. Axilla, thorax and root of neck (stage H). 24 and 25.
Lateral view of head and neck from the left and right respectively (stage H). 26. The mental
zone (stage H). 27. Left hind-limb showing the whorl just above the ankle (stage H).

192 THE HAIR TRACTS IN MARSUPIALS. II,

in one particular—the absence of the caudal member of the pair of whorls on the
vertex.
Family PHASCOLARCTIDAE.
PHASCOLARCTOS CINEREUS Goldfuss. Figs. 21-27.

Material.—C.604—a male (crown-rump length 92 mm.); no data. C.603—a
female (crown-rump length 103 mm.); no data. C.433—a male (crown-rump
length 112 mm.); Victoria; pres. Zoology and Acclimatization Society of
Victoria, 5th June, 1901.

All three examples of the species appear to coincide generally with Wood Jones’
original account (1923a), but only C.433 has the hair sufficiently developed for detailed
charting. Each has a single counter-clockwise whorl between the shoulders.

C.433 differs from the individual examined by Wood Jones in that there is no
whorl on the front of the hind-limb just above the ankle. This specimen also displays
an abnormality on the right side of the head (see below and Figs. 21 and 22).

It cannot be said with certainty whether or no C.603 and C.604 possess a whorl
on the front of the hind-limb owing to the weak development and sparseness of the hair.
Considering, however, the irregularity of hair inclination on the site where the whorl
normally occurs, it would seem likely that the whorl would have developed in each case.

No reference has previously been made in the literature to the arrangement of the
hair streams on the ventral surface. The typical phalangerine arrangement prevails.
Axillary whorls are present (Fig. 23).

My previous supplementary account (Boardman, 1948a) of the species was based,
for the most part, on a specimen designated “H” in the collection of the Australian
Institute of Anatomy. As a result of re-examination of this individual emended figures
are submitted (Figs. 24 and 25) of the hair streams on the sides of the head. The
right side of the head of this specimen is not symmetrical with the left, an arrangement
which was previously dismissed as an artefact. A variation of similar type on the
right side of C.433 (Figs. 21 and 22), however, leaves little doubt that it is an
abnormality.

A figure is also submitted of the whorl on the hind-limb and the condition of the
hair streams on the chin; both drawings are from specimen “H”’.

Family VOMBATIDAE.
VOMBATUS URSINUS URSINUS Shaw. Fig. 28.

Material.—R.4949—a female (crown-rump length 155 mm.); Flinders Island,
Bass Strait; 11th November, 1909.

Head, Neck and Trunk.—Disturbances to the primitive flow of hair are, for the
most part, traceable to a pair of whorls one on each side of the mid-ventral line opposite
a point a little above the middle of the humerus. The left whorl is clockwise, the
right counter-clockwise. From the level of the centres of the whorls caudally for a
short distance a hair-ridge is formed lying somewhat obliquely with reference to the
longitudinal axis; it terminates in a small convergent interval. The effect of the
presence of the whorls on the hair flow of the gular region is to produce a reversal
in the form of an extensive mid-ventral feathering. From this feathering and the
associated whorls hair streams on a recurved course across the upper arm, the shoulder,
and the side of the neck and face. The feathering terminates in front of the interramal
papilla where it forms in the mid-ventral line a small divergent interval where contact
is made with the caudally and laterally flowing current on the chin. The chin current
merges insensibly with the recurved flow at the cranial limit of the feathering thus
contributing to the general caudalwards current on the side of the face. The facial
current is divided by the ear around which it sweeps above and below to rejoin behind,
thence to continue straight back along the short neck. Immediately in front of where
these streams unite behind the base of the ear a strong recurvature of hairs both
from the dorsal and ventral components provides the hairy covering on the medial
aspect of the auricle and produces a small convergent interval behind the ear base.

BY W. BOARDMAN. 193

Caudal of the centre of the bilateral thoracic whorls hair streams on to the medial
aspect of the arm, into the axilla, and towards the mid-ventral line of the chest, there
to initiate a parting which persists back as far as the pouch. From this mid-ventral
parting the hair trend is caudally and dorsally to merge with the currents on the side
of the body. Consequent upon the flow away from the medial line the currents in the
inguinal region diverge and sweep round the pouch almost meeting behind and then
diverge again to surround in similar manner the cloacal hillock. There is some
asymmetry behind the pouch (see Fig. 28).

ys
S))
y

D)
Ly)
SS
Wy

<
}

Fig. 28.—Vombatus ursinus ursinus. Ventral view of whole animal.

Limbs.—The fore-limb exhibits a peculiarity in that along the postaxial aspect of
the forearm the postaxially or postaxially and distally flowing current on the lateral
and medial faces of the limb bends sharply towards its extremity to produce a broad
distally flowing stream. A somewhat similar situation occurs on the shank of the
hind-limb, but since the current on the medial surface is already flowing distally, only
the flow on the lateral surface is involved in the distal bending. On the postaxial
margin of the thigh a hair-ridge occurs brought about by the meeting of the currents
from its medial and lateral surfaces; at about the middle of this ridge a small
convergent interval is produced by a hair disposition such that proximal of the interval

194 THE HAIR TRACTS IN MARSUPIALS. II,

a stream flows towards the root of the limb, and cn the opposite side of it a stream
flows distally.

VOMBATUS URSINUS TASMANIENSIS Spencer and Kershaw. Fig. 29.

Material—No. 22, Pearson Collection (Tasmanian Museum)—a female
(length of head and body about 340 mm.); HEaglehawk Neck, Tasmania;
27th October, 1939.

wWS\

aE

=<
}
SS
\V
)

Ly

Fig. 29.—Vombatus ursinus tasmaniensis. Ventral aspect of the head.
Fig. 30.—Vombatus hirsutus hirsutus. Ventral view of head, neck and thorax.

Wood Jones (1924) has discussed the hair tracts in the Tasmanian member of
the genus and recorded that they showed no deviation from the primitive arrange-
ment in any part. The single specimen available to me does not support this view.

Unfortunately this very fine individual from the Pearson Collection has been
extensively dissected from the ventral surface whereon the main disturbances of the
primitive arrangement are found in the genus. There does not seem to be any doubt,
however, that the subspecies would be described in similar terms to those used for
the subspecies ursinus. The region where the paired whorls occur on the upper thorax
in the subspecies wrsinus is in this subspecies damaged, especially on the left side. On
the right side a counter-clockwise whorl is present situated on a slightly lower level
than in ursinus; it would appear that the bilateral pair exists.

BY W. BOARDMAN. 195

Two details in which the subspecies under consideration differs from wursinus are
worthy of note. In tasmaniensis the mid-ventral feathering which runs along the gular
region stops short just behind the interramal papilla. The arrangement of hair round
the interramal papilla is curiously complicated; it is plotted in Fig. 29 (cf. Fig. 28).
A further difference lies in the nearness to the flexure behind the knee of the
convergent interval on the hair-ridge lying along the postaxial margin of the thigh.

VOMBATUS HIRSUTUS HIRSUTUS Perry. Fig. 30.
The previous account (Boardman, 1943a) of this form is extended by a figure of
the ventral aspect of the head, neck and thorax to facilitate comparison with ursinus.
The drawing is from specimen ‘G’’. ,

SS

WS
x

SS
2;
pee

SS
a
SN

G

K

SAN

xs
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Figs. 31 and 32.—Potorous tridactylus apicalis. Lateral and ventral views respectively of
the whole animal.

Family MACROPODIDAER.
Subfamily PoToRoInak.
POTOROUS TRIDACTYLUS APICALIS Gould. Figs. 31 and 32.

Material.—Zb2—a male (crown-rump length 105 mm.); Lunawanna, S.
Bruny Island, Tasmania; coll. A. W. G. Powell, February, 1936 (Tasmanian
Museum Collection).

The hair tracts of one other member of the subfamily Potoroinae have hitherto been
described, viz., Bettongia penicillata (Boardman, 19430). The present species is similar
in type but displays considerable differences in detail which serve clearly to separate the
two species.

The bilateral system of divergent centres and partings recorded as occurring
ventrally on the thorax and neck of B. penicillata is, in the present species, reduced in
extent. The divergent centres (they have the facies of diffuse imperfect whorls) are
more cranial in position, being situated ventro-laterally on the neck medial of, and
on about the same level as, the shoulders. Consequently, the feathering is practically

U

196 THE HAIR TRACTS IN MARSUPIALS. II,

eliminated since the forwardly directed hairs enter almost immediately into the forma-
tion of the divergent interval on the side of the face.

A convergent interval occurs immediately behind the base of the ear. It is
produced where the current sweeping caudally along the dorsum of the head between
the bases of the ears diverges to join with the caudally and dorsally directed current
on the side of the neck. Hach of these two major elements provides in front a recurved
stream which flows over the medial surface of the auricle, thus completing the interval.

Hair disposition on the trunk does not follow the almost completely primitive plan
found in B. penicillata. Instead, the currents from the axilla and adjacent lower thorax
recurve away from the mid-ventral line and flow dorsally and caudally on the side of
the body between the attachments of the limbs. The effects of this arrangement are
shown in Figs. 31 and 32. The tracts of the inguinal region are illustrated in Fig. 32:
a figure (33) of the inguinal region of B. penicillata drawn from the same specimen as
the previous figures of the species is added for comparison.

SSS

a7 MSS
v ESE
IS |\ SSAA
= ae
Y mh Se

34

Ka ee

Fig. 33.—Bettongia penicillata. Hair tracts of the groin.
Figs. 34 and 35.—Caloprymnus campestris. 34. Hair tracts of the groin. 35. Head, neck
and upper thorax in ventro-lateral view.

CALOPRYMNUS CAMPESTRIS Gould. Figs. 34 and 35.

Material.—R.13612—a female, R.136183—a male (both have the length of
head and body about 200 mm.); Mulka, South Australia; 6th August, 1932.

Ordinarily, hair development on these specimens would be considered too advanced
for charting, but the account available of Bettongia penicillata (Boardman, 1943b)
leaves no doubt of how the tracts should be interpreted; the same ground-plan is found
to occur in both genera.

The bilaterally disposed featherings or partings situated ventrally on the upper
thorax, neck and head run obliquely inwards from the level of the cranial limit of the
axilla towards the angle of the mouth. They extend much further forwards than in
Bettongia so that the divergent interval on each side of the face comes to be situated
just behind the angle of the mouth. The reversed flow within the axilla, which
originates as in Bettongia in the recurvature of the flow on the side of the thorax over

BY W. BOARDMAN. 197

the weakly developed caudal fold of the axilla, is continuous with the feathering in front.
However, younger specimens may approach closer to the picture given for B. penicillata
as there is a possibility that the length attained by the hair has obscured some of the
details of its arrangement in tracts.

Elsewhere on the trunk the hair currents follow the primitive plan. No reversal
occurs between the scrotum and the cloacal hillock as described in Bettongia; the female
groin displays no reversals of any kind.

=

Figs. 36-37.—Onychogalea fraenata. Ventral and dorsal views respectively of head, neck
and thorax.

Subfamily MAcROPODINAE.
ONYCHOGALEA FRAENATA Gould. Figs. 36-39.

Material.—R.3056—a female (length of head and body about 223 mm.) ;
bred in Melbourne Zoological Gardens; died 2nd July, 1909.

The hair tracts of this unique specimen are very complex. In view of the necessity
to figure from dorsal, lateral and ventral aspects, a minimum of verbal description is
submitted.

Head.—There are three intervals on each side of the head—(a) a divergent interval
in front of the medial canthus, (b) a divergent interval midway between the eye and
the base of the ear, and (c) a convergent interval on the lateral border of the interramal
zone just behind the level of the medial canthus. In front of the divergent interval a
the current on the bridge of the nose recurves to produce a reversal that is carried
forward on to the rhinarium. Along the crown a convergent hair line conspicuously
indicated by a hair-ridge extends from about the level of the divergent interval b forward
to the level of the divergent interval a. A convergent centre, not very clearly defined, is
placed approximately over the side of the mandible at the level of the lateral canthus.
Both the convergent centre and the convergent interval ¢ are on a weakly developed
convergent hair line that runs from the base of the ear diagonally across the face to
the mid-ventral line at the caudal margin of the submental zone. The presence of these
two structures involves changes in direction of the hair streams (Figs. 36 and 38).

Neck.—a counter-clockwise whorl is situated mid-dorsally on the neck immediately
behind the occiput. On the side of the neck somewhat below the level of the mid-lateral
line, a convergent ridge occurs where the currents flowing over the side of the neck from
the nuchal whorl meet those of the ventral neck that flow with a general cranial and
lateral inclination from the upper thorax; a convergent interval occurs on it at about

198 THE HAIR TRACTS IN MARSUPIALS. II,

SSSy) =
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))

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Set

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Figs. 38-39.—Onychogalea fraenata. 38. Lateral view of whole animal. 39. The groin.

the middle of the neck. Cranially this ridge extends almost to the convergent point on
the side of the face; caudally it proceeds over the shoulder and on to the upper arm on
which it runs just lateral of and parallel with the preaxial border almost to the flexure
of the elbow. Its dorsal component at the caudal end is contributed by the recurved
flow from the outer surface of the caudal axillary fold.

Trunk.—Other than in the inguinal region the distribution of the currents on the
trunk is largely determined by the presence of whorls (clockwise on the right, counter-
clockwise on the left) within the axilla. The whorled systems give rise in front to the
reversed stream on the upper thorax and neck that recurves in its lateral portions over
the root of the neck, shoulder and arm to participate in the hair-ridge at the side of the
neck as described above, and behind to a current that sweeps caudally and laterally
across the axilla and recurves on to the side of the body over the axillary fold and the
flank just caudal of it. This recurved field is distributed on the trunk, shoulder and
upper arm as illustrated in Fig. 38; where it meets the caudally flowing current from
the mid-dorsal nuchal whorl a divergent interval occurs on each side situated over
about the middle of the scapula and just within its vertebral border.

BY W. BOARDMAN. 199

The tracts of the inguinal region are complex and show considerable asymmetry;
they are delineated in Fig, 39. The recurvature of the current on the buttocks on to
the postaxial aspect of the thigh involves a centripetal coil reminiscent of that recorded
as occurring on the left side of Wallabia bicolor (Boardman, 19430), but differing in the
constitution of the surrounding fields (Fig. 38).

Limbs.—The fore-limb presents no unusual features. The hind-limb has a pair of
whorls on the distal portion of the shank similar to those described in Thylogale sp.
(Boardman, 194380).

WALLABIA AGILIS Gould. Figs. 40 and 41.

Material.—R.13138—a female (length of head and body about 305 mm.);
E. Alligator River, Northern Territory; 11th July, 1912.

Head.—On the whole, the head resembles that of Onychogalea fraenata (v. supra)
but differs in having a convergent centre (whorled counter-clockwise) on the crown in
place of the extensive hair-ridge recorded in that species and in possessing a small whorl
with its centre immediately behind and just above the lateral angle of the eye. The
left side of the head shows differences from the right; these appear to be artefacts.

Neck.—Ventro-laterally on the neck at about the middle of its length a convergent
interval is present; it is associated with only a very short hair-ridge in front and
behind, but this may be due to the fact that the fur is not far developed in this the only
specimen available. The nuchal whorl is clockwise; it is situated just caudal of the
occiput and has its central portion longitudinally elongate. Ventrally on the neck the
hair reversal is the same as in Onychogalea.

Figs. 40 and 41.—Wallabia ‘agilis. 40. Lateral view of head. 41. Lateral view of hind end
of body.

Trunk.—The hair tracts of the trunk are similar to those of W. bicolor (v. infra
and Boardman, 1943b) with the proviso that, owing to extensive damage to the
inguinal region, it is not possible to give an account of that part other than to say
that it appears to be laid down along similar lines. The recurvature over the buttocks’
on to the postaxial aspect of the thigh shows less complications than in bicolor; there
are not present any supernumerary ridges or whorls, and there is no asymmetry
between the right and left limbs in the disposition of the tracts (Fig. 41).

WALLABIA BICOLOR Desmarest.

Under this name in a previous contribution (Boardman, 19430) a single male
specimen (930 part—Australian Museum Collection) was examined. The opportunity
to record further members of the subfamily including other species of the genus calls
for supplementary details.

Head and Neck.—The hair tracts of the head resemble the condition in Onychogalea
fraenata (v. supra) more than the condition in Thylogale sp. The convergent centre
on the crown has behind and continuous with it a short hair-ridge where the converging

200 THE HAIR TRACTS IN MARSUPIALS. II,

streams on the crown meet at the mid-dorsal line behind the centre. The hair-ridge
running obliquely across the face from the base of the ear terminates somewhat further
behind the interramal papilla than in agilis; as in agilis the ridge is practically
non-existent below the convergent interval. The specimen shows the convergent
interval on the side of the neck particularly well.

Trunk.—Like Onychogalea fraenata the arrangement on the ventral thorax, neck
and throat is caused by a divergent centre in each axilla which does not, in this
species, show any tendency to be whorled. From the centre a stream flows caudally
and laterally over the free border of the caudal fold of the axilla on to its lateral
surface where it encounters the caudal and ventral flow over the side of the thorax
and forms with it a convergent interval.

The presence of a centripetal coil on the left buttock has already been mentioned.
The condition occurs on both sides of W. dorsalis (v. infra).

WALLABIA DORSALIS Gray.

Material—M.5391—a male (length of head and body about 295 mm.);
North Coast district of New South Wales; coll. Noel Burnet, 22nd September,
1933 (Australian Museum Collection).

The hair characters of the head and neck are practically identical with those of
bicolor (v. supra) except that the nuchal whorled system is bilaterally doubled. The
left member of the pair is more cranially situated than that on the right side and is
also larger. The left whorl is counter-clockwise, the right clockwise. <A short hair-
ridge with an imperfectly formed convergent interval occurs in the mid-dorsal line
behind the right whorl.

On the trunk and limbs generally, no significant differences from bicolor could
be defined. Although the inguinal region is extensively damaged by incision and
folding, there seems to be close agreement with the account and figure given for bicolor
(Boardman, 19430, Fig. 25). The centripetal coil recorded as occurring on the left
buttock of bicolor is present on both sides in dorsalis.

WALLABIA Sp.

Material_—993—a male (length of head and body 230 mm.); South-Hastern
Queensland (Australian Museum Collection).

The specimen shows no differences except in detail from the accounts given of
other members of the genus. As in dorsalis (v. supra) a bilateral pair of whorls occurs
dorsally, but in this species they are somewhat in front of the cranial limit of the
ear base, and are, therefore, more accurately described as occipital whorls. The whorls
are symmetrical for size and position; that on the left is counter-clockwise, that on the
right clockwise and there is the usual short ridge and convergent interval between them
on the mid-dorsal line. The convergent point on the crown is well developed and shows
a tendency to clockwise whoriing.

While the tract arrangement on the side of the face is that of a Wallabia as at
present known, it is of specific interest that the convergent centre on the side of the
face is replaced by a centripetal whorl. The convergent interval on the side of the
neck is further forward than usual and is separated from the whorl on the face by
only a short distance.

OSPHRANTER ROBUSTUS Gould.

In the specimen of Osphranter robustus that formed the subject of the previous
communication on the species (Boardman, 19430), it should be added that the con-
vergent interval on the side of the neck is ventrolaterally placed on the root of the
neck immediately in front of the shoulder. In this respect Osphranter robustus
approaches closer to the conditions described for Thylogale sp. than any other of the
Macropodinae described herein.

BY W. BOARDMAN.

Macropus MAsgor Shaw. Figs. 42 and 43.

201

Material—R.1531—a young male (length of head and body 400 mm.);

Victoria; 9th July, 1906.

Head.—The head has the hairs arranged on the plan set down for Thylogale sp.
(Boardman, 19430). There is a convergent interval above the medial angle of the

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202 THE HAIR TRACTS IN MARSUPIALS. II,

eye and immediately in front of the supraorbital papilla. The divergent interval
situated between the eye and the ear occurs about opposite the middle of the base of
the ear but above the level of the lateral angle. The convergent centre on the crown
has a slight tendency to be whorled in a clockwise fashion. A. divergent centre is
situated between the ears; just in front of its level on the dorsum of the head and
medial of the uppermost limit of the base of the ears, the hairs flowing cranially and
laterally from the centre meet an upward flow from the front of the base of the ear
to produce a convergent interval. The stream sweeping round from behind the ear
forms with the current on the face a convergent ridge which runs obliquely across
the angle of the mandible towards the mid-ventral line; the identity of this ridge is,
however, lost over the angle of the mandible. Formation of a mid-dorsal occipital
divergent interval will be considered below.

Neck and Trunk.—Macropus major has a divergent centre in the mid-dorsal line
at about the centre of the back. From it a parting extends along the middle line
cranially to a position approximately above the occipital region of the skull at which
level the backward flowing current from the divergent centre between the ears is
encountered and a divergent interval produced. Caudally from the mid-back divergent
centre a similar feathering occurs and this persists to about the cranial limit of the
ilium. The normal direction of hair on the side of the trunk is not greatly affected by
the reversal mid-dorsally.

As in Thylogale sp. the arrangement of the tracts on the ventral thorax is dependent
on the presence of bilaterally arranged centres of dispersal, in this case, in the form
of whorls on the medial aspect of the upper arm somewhat distal of the middle of the
humerus; on the left arm the whorl! is clockwise, on the right counter-clockwise. At
the root of the neck in front of and. Slightly ventral of the shoulder, a convergent
interval is formed similar to that shown as a somewhat imperfect structure in
Thylogale (Boardman, 19436, Fig. 24).

Tract disposition in the inguinal region is charted in Fig. 43.

Limbs.—Except for the presence of the whorl described above on the medial aspect
of the upper arm, both fore- and hind-limbs may be said to have the primitive
arrangement. ;

LITERATURE CITED.
BoaRDMAN, W., 1943a.—On the External Characters of the Pouch Young of Some Australian
Marsupials. Aust. Zool., 10: 138-160. :
, 1943b.—The Hair Tracts in Marsupials. Part i. Description of Species. Proc. LINN.
Soc. N.S.W., 68: 95-113. —
IREDALE, T., and TROUGHTON, HE. Le G., 1934.—A Check-list of the Mammals recorded from
Australia. Aust. Mus. Mem., 6: 101 pp.
JONES, F. Woop, 1920.—The External Characters of Pouch Embryos of Marsupials. No. 1.
Trichosurus vulpecula var. typicus. Trans. Roy. Soc. S. Aust., 44: 360-373.
, 1923a.—Id., No. 5. Phascolarctus cinereus. Ibid., 47: 129-135.
, 1923b.—Id., No. 6. Dasycercus cristicauda. Ibid., 47: 136-141.
, 1924.—Id., No. 9. Phascolomys tasmaniensis. Ibid., 48: 145-148.

EXPLANATION OF PLATE VIII.

Pseudocheirus convolutor convolutor. The groin of the Pearson Collection male drawn to
show the relationships between hair arrangement and the scrotum and its pouch.

203

NITROGEN FIXATION IN LEGUMINOUS PLANTS. VI.
FURTHER OBSERVATIONS ON THE EFFECT OF MOLYBDENUM ON SYMBIOTIC NITROGEN FIXATION.

By H. L. Jensen, Macleay Bacteriologist to the Society.
(From the Department of Bacteriology, University of Sydney.)

(Plate ix; one Text-figure. )

[Read 31st October, 1945.]

INTRODUCTION.

It is now generally recognized that molybdenum is one of the several ‘‘minor” or
“trace” elements required for normal! growth of plants, and the investigations of recent .
years have shown that soils more or less deficient in molybdenum occur in several
parts of Australia. Such soils were found in South Australia by Anderson (1942), in
Tasmania by Stephens and Oertel (1943) and by Fricke (1944), in New South Wales by
Shaw et al. (1944), and in Western Australia by Teakle (1944). These experimental
data refer chiefly to pasture legumes, especially lucerne and subterranean clover; Fricke
(1944) states that legumes in field experiments, upon the whole, appear more responsive
than grasses to molybdenum fertilizers. Previous experiments (Jensen and Betty, 1943)
suggested that molybdenum is needed especially for the process of symbiotic nitrogen
fixation in the root nodules of the legumes, as it has long been Known to be for the
corresponding process of non-symbiotic nitrogen fixation by Azotobacter and apparently
also certain blue-green algae (Bortels, 1938). The present paper gives some additional
evidence derived from experiments which temporarily had to be discontinued owing to
the difficulty of obtaining sufficiently molybdenum-free sand for the pot experiments.

EXPERIMENTAL.

In the first series of experiments, lucerne was grown in sand which had already
been used for similar cultures until signs of a definite molybdenum-deficiency began
to appear (Jensen and Betty, 1943). As in the previous experiments, the plants were
grown in glazed earthenware pots holding 3 kgm. of sand with addition of a basal
fertilizer mixture consisting of 0-25 gm. K.HPO,, 0:25 gm. KH.PO,, 0:25 gm. CaCl, and
0-1 gm. MgSO, After 150 days, when two cuts of the tops had been taken, the same
quantity of salts was again added, together with 3 mgm. of MnSO,, ZnSO,, CuSO, and
Na.B,0O, per pot. The experiment included three treatments:

(1). No addition of molybdenum (in the following called ‘0 Mo’).

(Aj) al mem. Na,.MoO, per kgm. of sand (“+ 1 Mo’), added when the first crop was

sown (Jensen and Betty, 1943).

(3). 5 mgm. Na,MoO, per kgm. of sand (‘t+ 5 Mo”).

To make up for the removal of molybdenum in the previous crops, each molybdenum
pot was given an extra dose of 1 mgm. Na.MoO, per kgm. of sand. Each treatment
included six replicate pots with eight plants in each. Lucerne seeds of the variety
“Giant Upright’, inoculated with an effective strain of Rhizobium Meliloti, were sown
on 23rd March, 1943. The pots were kept in a greenhouse and watered with distilled
water throughout the growth period. The tops were cut four times, after 94, 150, 185
and 216 days. After 150 days, three pots from each treatment were given a supply
of combined nitrogen in the form of 0:607 gm. sodium nitrate (= 100 mgm. N) which
by analysis was found to contain less than 0:06 part per million of molybdenum.

Vv

204 NITROGEN FIXATION IN LEGUMINOUS PLANTS. VI,

At the conclusion of the experiment (216 days) the roots were also collected, and
the root nodules of the plants that had not received nitrate were separated from the
root-substance proper. Nitrogen and molybdenum were then determined in the dried
and finely-ground substance of tops, roots, and nodules. The tops of the first two and
the last two cuts were bulked for analysis. Nitrogen was determined by the Kjeldahl
method, with selenium as a catalyst in the digestion, and molybdenum by Marmoy’s
thiocyanate method, as described by Piper (1942). All figures are calculated on the
basis of material dried at 96—-98°C.

Dry matter, gm.

0 50 100 150 200
Days. ; :
Fig. 1.—Average yield per pot of dry matter in tops of lucerne harvested at four successive
stages (94, 150, 185 and 216 days).

TABLE 1.
Influence of Molybdenum on Growth of Lucerne in Sand Culture.

0 Mo. 1 Mo. 5 Mo.
Mean. 8.D. Mean. S.D. Mean. §8.D.
Tops 7°65 +0°66 10:75 +0-89 11:62 +0-50
Dry Matter, Roots 2-70 +0-32 .. 2°97 +0-13 2°87 +0:23
gm. per Pot. Nodules 0-251 +0-032 0-167 40-035 0-191+0-066
Nodules in % of Total Dry Matter. 2-38 +0-41 Theil), SEO)outs) 1-29 +0-12
Tops, cuts 1+2 3-79 4-48 4-29
Percentage N in 59 9 Oaret 3°22 3°34 3-20
Dry Matter. Roots 1-49 2-84 3°04
Nodules 5-02 5:76 5°56
Total Nitrogen, mgm. =e nt a 4 319:0 +19-7 506-0 +41:7 529°0 +26-6
Uptake of N, mgm. per gm. Dry Nodule- 1285 +194 3085 +459 2799 +288
substance.
Tops, cuts 1+2 .. ae 0-45 6-0 PALO
Mo-content of 3 53 (OA ee ne 0:48 7:8 15:5
Dry Matter, Roots .. te ee 0:48 8:6 2523
p.p.m. Nodules ik i aa 2% 29°3 62°8

The yield and composition of the crops (not including the pots receiving nitrate)
are shown in Text-fig. 1 and Table 1. The yields of dry matter in tops are seen to be
increased by some 40 to 50%, due to the addition of molybdenum, over the treatment
“Q Mo”, and this influence is fairly constant all through the growth period. The
difference between the two doses of molybdenum is not significant. No special symptoms

BY H. L. JENSEN. 205

of deficiency were noticed in the plants not given molybdenum, nor was there any
visible difference in colour, as observed by Stephens and Oertel (1943) in subterranean
clover; the molybdenum-deficient plants merely differed from the others by being
smaller in size. The analyses show that the percentage of nitrogen in the tops of the
first two cuts is also considerably increased where molybdenum is supplied. As to the
roots, their actual weight is not significantly influenced, but the decrease in nitrogen
percentage in the molybdenum-deficient plants is even more pronounced in the roots
than in the tops, as also found in the previous experiments (Jensen and Betty, 1943).
The total yields of nitrogen in the crops therefore show an even stronger effect of
molybdenum than do the weights of the tops, being some 60 to 66% higher than in the
set of pots without molybdenum, but also, in this respect, the difference between the
high and the low dose of molybdenum is insignificant. The nodules show a most
interesting phenomenon: not only is there no decrease in the mass of nodule-substance
of the molybdenum-deficient plants, but on the contrary, their weight is actually a little
higher, and the proportional weight of nodules in percentage of total dry matter is
roughly twice as high as in the plants supplied with molybdenum. The increase appears
to be due to a larger. average size of individual nodules and not to a larger number;
although no counts were made, the type of nodule formation looked the same in all
three treatments. This stronger development of nodule-tissue and smaller yield of
nitrogen indicates a very conspicuous reduction in the nitrogen-fixing efficiency of the
nodules in the molybdenum-deficient plants, as shown by caiculating the uptake of
nitrogen per unit weight of dry nodule-substance; this figure is seen to be nearly
2% times higher in the plants given molybdenum than in the molybdenum-deficient ones,
but again there is no significant difference between the two doses of molybdenum.

The molybdenum determinations show, as in the previous experiments (Jensen and
Betty, 1943), that the nodules are constantly richer in molybdenum than the rest of
the plants, and this difference is most pronounced in the series “0 Mo” where the
plants have still been able to obtain a quantity of approximately 5y of molybdenum per
pot, possibly from the water and the nutrient salts, but perhaps more likely from the
gravel that was used for equalizing the weights of the empty pots and as a surface
mulch to reduce evaporation. The fact that the tops of the molybdenum-deficient plants
contain approximately 0:5 part per million of molybdenum, agrees well with the
observation of Stephens and Oertel (1943) that a molybdenum content of about 1 p.p.m.
in the tops is necessary for normal growth of white and subterranean clover, and that
0-5 p.p.m. is suboptimal. Furthermore, the present results show that a considerably
higher concentration of molybdenum in the nodule-tissue is necessary for an optimal
rate of nitrogen fixation. In lucerne, this optimal Mo-concentration seems to lie between
approximately 3 and 30 parts per million of dry matter. Most samples of nodule-
material taken from plants growing in ordinary soils have shown a molybdenum content
of this order of magnitude (Bertrand, 1940; Jensen and Betty, 1943); it would be of
great interest to extend these observations to the nodules of plants in molybdenum-
deficient soils under field conditions.

The fact that the molybdenum-deficient plants develop a larger proportion of
nodule-substance than plants with an adequate supply of molybdenum is interesting in
view of the observation of a similar phenomenon in lucerne and clover grown in sand
or soil of different reaction, where the relative weight of the nodules tended to decrease
in alkaline medium (Jensen, 1943). This might indeed be due to molybdenum rendered
available by the addition of lime to the acid substrate (cf., Stephens and Oertel, 1943),
but the phenomenon has also been observed, though rarely so markedly, in later
experiments where an adequate supply of molybdenum was given. Still another example
of the same phenomenon can be found in the data of Jones and Tisdale (1921) on the
rate of nodule growth and nitrogen fixation in soy beans at different temperatures.
These authors observed that the largest proportion of nodule-substance developed at
21-24°C., but the strongest nitrogen fixation took place at 27-30°C., at which latter
range of temperature the uptake of nitrogen (atmospheric plus combined nitrogen from

206 NITROGEN FIXATION IN LEGUMINOUS PLANTS. VI,

the soil) per unit of dry nodule substance appears roughly twice as high as at 21—24°C.
(data from Jones and Tisdale’s Tables III and IV). f

It thus seems that the legumes possess a mechanism of adaptation by means of
which they are able, within certain limits, to compensate for the lower nitrogen-fixing
efficiency of the nodules by developing a larger proportion of nodule-tissue in response
to unfavourable environmental factors like molybdenum deficiency, acid soil reaction,
or suboptimal temperature. ;

The three pots of each molybdenum-treatment, to which sodium nitrate was added,
gave the yields shown in Table 2.

TABLE 2.
Influence of Molybdenum on Growth of Lucerne supplied with Sodium Nitrate.

0 Mo. 1 Mo. 5 Mo.
Mean. §.D. Mean. §S.D. Mean. S.D.

Dry Matter, gm.—

Tops, cuts 3+4 ‘ eA 0: ae 5-752 0-52 6:68+ 0°37 7:20+ 0-05

Roots + Nodules a ae oe a 3:°51+ 0:14 4-05+ 0:23 3:87 0-32
Percentage N in Dry Matter—

‘RODS ar ie 2°54 3°17 3-15

Roots + Nodules 1:94 2-09 2-19
Total Nitrogen, mgm. ae ia yn ae 215:0+16°5 296:0+11-4 310-0+ 8:2
(Do. in Pots—NaNO,) 188:0+18:2 285:0+ 4:0 287-0+18-8

It appears at once that the increase in weight of tops due to molybdenum is much
less pronounced than in the corresponding pots without nitrate, where the combined
weights of the third and fourth cut in the three treatments were, respectively, 4-20,
5:72, and 5:93 gm. This levelling effect of the nitrate was most pronounced in the
third cut, the first after the addition of nitrate, which gave the following average yields
in gm. per pot:

5 Mo.

0 Mo. 1 Mo.
Pots+NaNO, ve 2-84 3°44 3°69
°% increase due to Mo — 20 30
Pots—NaNO, ts 1:94 2-95 3:02
% increase due to Mo — 52 56

The appearance of the plants immediately before taking the third cut is shown in

Plate ix.

In the fourth cut, after which all the added nitrate had disappeared from

the sand, the effect of the nitrate was still noticeable but much less pronounced; the
yields of dry matter in this cut were as follows:

0 Mo. 1 Mo. 5 Mo.

Pots +NaNO; ae 2-91 3:24 Siow
% increase due to Mo — 11 26
Pots—NaNO, Me 2:26 PACU ey 2-91
% increase due to Mo — 23 29

It is further seen in Table 2 that the percentage of nitrogen in the molybdenum-

deficient plants is much lower than in the corresponding plants growing with free
nitrogen (Table 1). In the plants with the two molybdenum treatments there is a
similar reduction in the percentage of nitrogen in the roots (+ nodules), and in all
three treatments there is no significant difference in the total nitrogen contents of
plants grown with free nitrogen alone and with supply of nitrate.

It thus appears that the production of dry matter under conditions of molybdenum
deficiency is less strongly affected when combined nitrogen is supplied than when the
plants have to depend on fixed nitrogen only. This, together with the fact that the

BY H. L. JENSEN. 207

nodules require for optimal rate of nitrogen fixation a concentration of molybdenum
higher than the rest of the plant tissues, seems to show conclusively that molybdenum
is essential not only for the general plant metabolism, but also for the specific process
of nitrogen fixation. This strongly supports the view that the biochemical mechanism
of nitrogen fixation is essentially the same in the leguminous plants and in Azotobacter,
perhaps also the nitrogen-fixing blue-green algae.

An attempt was made to grow a new (fourth) crop of lucerne in the same sand,
this time with addition of sodium nitrate to half the pots from the beginning, but
this attempt, unfortunately, failed because the sand appeared in some way to have
become unsuitable for further growth of lucerne. The seedlings in this crop showed
a heavy mortality, and the surviving plants made only a very poor growth in three
months. Further tests were therefore made with two other sands, but these appeared
too rich in molybdenum to show any direct response.

The first of these was a faintly acid river sand of medium fineness, containing a
fair amount of silt and, according to a previous analysis, 0:02 p.p.m. of molybdenum
(Jensen and Betty, 1943). Lucerne was grown in an experiment comprising com-
_ binations of the following treatments, in addition to a basal fertilizer mixture similar
to the one used in the previous experiment:

(a). 0:2% calcium carbonate, to give approximately pH 7.

(0). 0-600 gm. ammonium nitrate per pot (= 210 mgm. N).

(c). 6:0 mgm. sodium molybdate (2 p.p.m.) per pot.

The first crop of lucerne was sown on 7th March, 1944, in four replicate pots of
each treatment, with eight plants in each. This crop grew only very slowly and poorly,
agreement between replicate pots was unsatisfactory, and no influence of the molybdenum
was discernible when the plants were harvested after 20 weeks. The only significant
effect was on the relative weight of the nodule-substance in the pots without ammonium
nitrate, which in the acid sand (pH 4:8—5:3) averaged 2-28% of total dry matter, against
1:52% in the alkaline sand (pH 7:0-7:5); the corresponding figures for uptake of
nitrogen in mgm. per gm. dry nodule-substance were 1,380 and 2,550 mgm. respectively.
The main reason why the experiment is recorded here is that analysis of the crop
showed a lower molybdenum content of plants grown with combined nitrogen, as also
observed in an earlier experiment in the same sand (Jensen and Betty, 1943). Nodules
were practically absent on the plants given ammonium nitrate, and from those with
free nitrogen, the amount left over after nitrogen determination was insufficient for
separate molybdenum determination. The molybdenum content of the tops and roots
is seen in Table 3.

TABLE 3. : 2
Molybdenum Content, p.p.m. of Dry Matter, of Lucerne Plants grown in Sand with Free and Combined Nitrogen.

Addition of Na,MoO, sa nt Ae None. 2 p.p.m.

Source of Nitrogen NED NH,NO3. No. NH,NO3.
Sand —CaCO, Tops Ais oC 8e/7, 0-4 9:5 BEB}
(pH 4:8-5-3). Roots A Ae 0:3 (lost) 12-3 7-4
Sand +CaCO,; Tops ats a 0:7 0:7 16°3 10:8
(pH 6-6-7°-5). Roots .. ae 4-0 2-0 Dealt il eng

The tendency to reduction in the uptake of molybdenum when combined nitrogen
has been provided is obvious, although it is not completely constant, and it is a
remarkable fact that the figures for tops and roots are almost reversed in acid and
alkaline sand without molybdenum and combined nitrogen. More consistent results
were found when molybdenum was determined in lucerne grown for three months in
the same sand at two ranges of pH, approximately 5 and 7, with addition of 0-5 p.p.m.
of sodium molybdate and combined nitrogen (240 mgm. N) as alternating doses of
sodium nitrate and ammonium sulphate. This was one of several experiments, to be
discussed in detail later, which were designed to test the influence of varying reaction

208 NITROGEN FIXATION IN LEGUMINOUS PLANTS. VI,

and nitrogen supply on the nitrogen-fixing efficiency of the root nodules. In this
instance the growth of the lucerne was excellent, and sufficient material was available
for determination of molybdenum in the nodules, even the small amount of almost
ineffective nodule-tissue formed in the presence of combined nitrogen.

TABLE 4.

Molybdenum Content, p.p.m. of Dry Matter, of Lucerne Plants grown with Free and Combined Nitrogen in Sand with
Addition of 0-5 p.p.m. Na,Mo0,.

Sand —CaCO;,. Sand +CaCOs.
(pH 5-1-5-4). (pH 7-0-7-4).
NaNO, and NaNO, and
Source of Nitrogen. Ne. (NH,)2504. Ni: (NH,).SO,.
P.p.m. Mo in Tops Ais a ae 22-5 15:4 19-8 19-4
Be ue VOOtSmes a ee He 16°3 Bo 21-6 BBO
,. Nodules rid ae a 136-2 3° iiky/ee) 93-1

” 2”?

The molybdenum content of this crop is unusually high (a calculation showed that
roughly 25% of the added molybdenum had been assimilated), but at acid reaction
the higher concentration of molybdenum in plants dependent on free nitrogen is quite
unmistakable, especially in the nodule-substance. At neutral reaction there is no such
difference, but it is noteworthy that even in plants with such an abnormally high
molybdenum content, its accumulation in the nodules is still very marked. There is
little difference between the tops and the roots; the tops from acid sand with free
nitrogen are even significantly richer in molybdenum than the roots (cf., Table 1,
1st column). The change of reaction produced by the addition of lime has had little
influence, except that it has somewhat increased the uptake of molybdenum in the
presence of combined nitrogen. Generally the figures form a remarkable contrast to
those given by Stephens and Oertel (1943) who found only 2 parts per million of
molybdenum in tops of subterranean clover grown in a clay soil with an even higher
dose of molybdenum (2-5 mgm. ammonium molybdate per 4 kgm. moist soil) than
was used in the present experiment. Apparently the molybdenum-deficient soil used
by Stephens and Oertel must have been able to immobilize the added molybdenum.

Incidentally figures like those in Table 4 convey a warning against an indiscriminate
use of readily available molybdenum compounds as fertilizers for soils that already
contain an adequate supply of this element. The addition of 0-5 mgm. sodium molybdate
to each kgm. of sand corresponds roughly to 1 lb. per acre, an amount which is often
used-under field conditions (Anderson, 1942; Fricke, 1944), and yet the application of
this small dose to a sand by no means rich in molybdenum has been sufficient to raise
the molybdenum content of the lucerne tops to roughly 20 parts per million. This
concentration borders on the limit at which herbage becomes dangerous to domestic
animals. Muir (1941), in a discussion of the properties of so-called “teart’’ pastures in
certain areas of England, where an excessive molybdenum content of soil and herbage
causes chronic poisoning in grazing cattle, reports that plant material from affected
areas contains from 20 to 100 p.p.m. molybdenum in dry matter, against usually less
than 5 p.p.m. in healthy areas. He also called attention to the paradoxical fact that
“improvement” of such pastures may actually aggravate the condition of cattle, owing
to the fact that clovers take up more molybdenum than the grasses, and that application
of lime and other basic fertilizers increases the uptake of molybdenum.

After the experiment recorded in Table 3, another crop of lucerne was grown in
the alkaline sand without combined nitrogen. This crop, which was sown on 23rd
August, 1944, and harvested after 112 days, grew very well, but the addition of
molybdenum had no effect on the yields of nitrogen or the efficiency of the nodules,
as shown in Table 5. The concentration of 27-6 p.p.m. of molybdenum in the nodules
has thus been fully sufficient for their activity (cf., Table 1, treatment “+ 1 Mo”). The
only apparent effect of the molybdenum is a somewhat higher percentage of nitrogen

BY H. L. JENSEN. 209

in the tops, but this is not sufficient to cause any significant increase in the actual
nitrogen content of the crop.

TABLE 5.
Composition of Plants grown in Two Sands with and without Extra Addition of Molybdenum.

Addition of Na,MoQ,. None. 2 p.p.m.

Mean. 8.D. Mean. S.D.

1. Lucerne in River Sand—

Total N in Plants, mgm. .. is 230 +. 32'-0 255 + 32-2
Per cent. N in Tops (d.m.) ze 2-40 7 Peel
Uptake of N, mgm. per gm. Dry
Nodule-substance .. Ae ae 2004 +433 2289 +149
Mo in Nodules, p.p.m. as ne 27-6 150-0
2. Subterranean Clover in Hill Sand, pH
4-8-5 -0—
Total N in Plants, mgm. Se 248 + 57-5 299 + 48-0
Per cent. N in Tops (d.m.) oN 3°05 3°29
Uptake of N, mgm. per gm. Dry
Nodule-substance .. a oe 882 +135 910 +137
Mo in Nodules, p.p.m. an = 11-6 3y7/ 35)
3. Same, Sand+0-2% CaCO;, pH 7:5-7°8
Total N in Plants, mgm. a 282 + 28-2 251 + 27-1
Per cent. N in Tops (d.m.) ae 2°65 3:29
Uptake of N, mgm. per gm. Dry
Nodule-substance : 774 = 51-2 915 + 54-5

Mo in Nodules, p.p.m. ak ae 20:6 109-0

Also a fine, yellow, hill sand of acid reaction was-tried for possible response to
molybdenum. The sand was given doses of 2 p.p.m. of sodium molybdate and 0:2%
calcium carbonate, in addition to a basal fertilizer consisting of 1-2 gm. CaHPO,, 0°3 gm.
MegSO,, 0:3 gm. KCI, 0-1 gm. FeCl,, and minor elements as in the previous experiments,
all per 3 kgm. sand. Subterranean clover, of the variety ‘Mount Barker’, inoculated
with effective root-nodule bacteria, was sown in triplicate pots of each treatment, with
eight plants in each, on 12th May, 1944, and harvested after 129 days. The results of
this experiment are also seen in Table 5. The crop made a very good growth, but the
harvest yields do not indicate any beneficial effect of the molybdenum. Its addition
has resulted in a somewhat higher percentage of nitrogen in the tops, especially at
alkaline reaction, but as in the previous experiment, this is not reflected in a significantly
higher return of total nitrogen; at alkaline reaction the weight of the tops was actually
somewhat, although not significantly, decreased by the molybdenum addition. The
content of 11-6 and 20-6 p.p.m. of molybdenum in the nodules of plants from acid and
alkaline sand, respectively, appears to have been fully sufficient; in the alkaline sand
the uptake of nitrogen per gm. of dry nodule-substance is actually raised a little by
the addition of molybdenum, but although the difference appears significant (n = 4,
t = 3-267, P: 0:05-0:02), it is due entirely to depression in the weight of nodules and
not to an increase in the yield of fixed nitrogen. It is further seen that in this sand
the addition of lime has considerably increased the availability of the molybdenum;
not only is the molybdenum content of the nodules approximately doubled at alkaline
reaction in the pots not supplied with molybdenum, but the same was the case with the
tops which contained 3-0 and 5:3 p.p.m. respectively. The remarkably small increase in
molybdenum content of nodules in acid sand plus molybdenum, together with the large
increase that results when lime is also added, suggests that the sand possesses some
mechanism that renders the added molybdenum unavailable, but that this is counter-
acted by the addition of lime (cf., Stephens and Oertel, 1943).

No further experiments were undertaken with these sands which evidently contained
too large a reserve of molybdenum, but it is hoped to repeat and extend the observations
on more molybdenum-deficient growth-media.

210 NITROGEN FIXATION IN LEGUMINOUS PLANTS. VI,

SUMMARY.

Pot experiments with lucerne in sand of very low molybdenum content showed a
considerable decrease in yield of dry matter and particularly of nitrogen by molybdenum-
deficient plants containing only approximately 0:5 part per million of molybdenum in
dry matter of tops and roots and 2:7 parts per million in the nodules. Such plants
developed a larger mass of nodule-substance than plants with an adequate supply of
molybdenum. The gain of nitrogen per unit weight of nodule-substance was approxi-
mately 23 times higher in normal than in molybdenum-deficient nodules. Increasing
the molybdenum content of the nodule-tissue beyond 28-30 parts per million had no
effect on either the actual or the relative gain of nitrogen. When given a supply of
sodium nitrate, molybdenum-deficient plants made a better growth than when dependent
on free nitrogen alone; the nitrogen content of the dry matter, however, was greatly
reduced. Plants supplied with combined nitrogen generally contained less molybdenum
than plants living on free nitrogen; exceptions to this rule were sometimes seen when
large doses of molybdenum were given, particularly together with calcium carbonate.
An experiment with subterranean clover showed no significant effect when molybdenum
was added to a sand from which the root nodules could obtain some 10 to 20 parts per
million of molybdenum.

Generally it appears that. molybdenum is essential for the specific process of nitrogen
fixation, and that the root nodules, in order to carry out this process at an optimal
rate, must contain more than 3 parts per million of molybdenum in dry matter, while
an increase beyond 20-30 p.p.m. has no additional stimulating effect.

REFERENCES.

ANDERSON, A. J., 1942.—Molybdenum Deficiency on a South Australian Ironstone Soil. J. Aust.
Inst. Agric. Sci., 8: 73-75.

BERTRAND, D., 1940.—Sur le molybdéne des nodosités des légumineuses. C. R. Acad. Sci. Paris,
211: 670-672.

Borte ts, H., 1938.—Entwicklung und _ Stickstoffbindung bestimmter Mikroorganismen in
Abhangigkeit von Spurenelementen und vom Wetter. Ber. dewts. bot. Ges., 56: 153-160.
FRICKE, HE. F., 1944.—Molybdenum Deficiency. Field Experiments at Cressy, Longford and

North Mottonts slasmindie Agricy,. b= 365= 10:

JENSEN, H. L., and Betty, R. C., 1943.—Nitrogen Fixation in Leguminous Plants. iii. Proc.
LINN. Soc. N.S.W., 68: 1-8.

——__——., 1943.— Nitrogen Fixation in Leguminous Plants. iv. Ibid., 68: 207-220.

JONES, FE. R., and TISDALE, W. B., 1921.—EHffect of Soil Temperature upon the Development of
Nodules on the Roots of Certain Legumes.. J. Agric. Res., 22: 17-32.

Muir, R. W., 1941.—The “Teart” Pastures of Somerset. Vet. J., 97: 387-400.

PipgrR, C. S., 1942.—Soil and Plant Analysis. Adelaide.

SHAW, N. H., Barriz, N., and Kipps, BH. H., 1944.—The Effect of Lime, Phosphate and
Molybdenum on the Growth of Lucerne in Duntroon Loam. J. Cowne. Sci. Industr. Res.,
Melbourne, 17: 233-241. :

STEPHENS, C. G., and OERTEL, A. C., 1943.—Responses of Plants to Molybdenum in Pot
Experiments on the Cressey Shaley Clay Loam. Ibid., 16: 69-73.

TEAKLE, L. J. H., 1944.—Molybdenum as a Fertilizer in Western Australia. J. Dept. Agric.
W. Aust., 21: 335-339.

EXPLANATION OF PLATE IX.

Appearance of lucerne plants in sand with three molybdenum treatments, after 185 days
(first crop after addition of combined nitrogen). Above: pots with free nitrogen only. Below:
pots with addition of 100 mgm. nitrogen as NaNoO,.

(S. Woodward-Smith photos. )

211

CONTRIBUTIONS TO A KNOWLEDGE OF AUSTRALIAN CULICIDAE.
No. VIII.*

By FraAnK H. Taytor, F.R.E.S., F.Z.S., School of Public Health and Tropical
Medicine, University of Sydney.

(One Text-figure. )

{Read 28th November, 1945.]

This paper places on record new localities for three species. One new species of
Finlaya is described.

ANOPHELES (ANOPHELES) ATRATIPES Skuse.

Proc. Linn. Soc. N.S.W., xiii, 1889, 1755.
This species was taken, fairly common, on two occasions biting in full sunlight.
Habitat: New South Wales: Byron Bay (F. H. Taylor).

URANOTAENIA PYGMAEA Theobald.

Monogr. Cul., ii, 1901, 254. :
This genus has not previously been found in New South Wales.
Habitat: New South Wales: Tweed Heads; Nelson’s Bay (F. H. Taylor).

TAENORHYNCHUS (COQUILLETTIDIA) XANTHOGASTER Edwards.

Bull. ent. Res., xiv, 1924, 366.

This species, so far as my knowledge goes, has not been previously recorded from
New South Wales.

Habitat: New South Wales: Maclean (F. H. Taylor).

Aé&pDES (FINLAYA) KOCHI Donitz.

Insektenborse, v, 1901, 38 (Culex).

This species is of considerable importance since it is an efficient intermediary host
of Wuchereria bancrofti Cobbold. It extends for some distance down the north coast
of New South Wales.

Habitat: New South Wales: Murwillumbah, Byron Bay (F. H. Taylor).

AEDES (FINLAYA) CLINTONI, 0. Sp.

Head: Proboscis dusky-brown, no banding or pale scales present; palpi slightly
shorter than proboscis, dusky-brown, slightly hairy apically, segments three to five
inclusive with conspicuous white basal banding; antennae brown, plumes brown, torus
covered with narrow white scales; a narrow border of flat white scales round the eyes
except in front where they are much narrower, rest of head with narrow white ones
and black upright-forked ones.

Thorax dusky-brown covered with brown narrow-curved scales, posterior half, except
the extreme lateral border, unfortunately rubbed, a broad stripe of narrow silvery-white
scales, occupying about one-third the width of the thorax, posterior extent of white
scaling undefined due to the abraded thorax; scutellum brown, lobes covered with
narrow white scales; pleurae: pronotum and postpronotum with a patch of silvery-white
flat scales, postspiracular area with a small patch of silvery-white flat scales, propleuron

* Continued from these PROCEEDINGS, Ixix, 1944, 120.

W

22, AUSTRALIAN CULICIDAE. VIII.

covered with silvery-white flat scales, postspiracular area and sternopleura covered with
patches of white flat scales, a large elongate patch below the wing base, coxae with
prominent patches of white scales.

Wings with brown scales, mostly denuded; base of posterior forked cell nearer the
base of the wing than that of the anterior forked cell, stem of the latter about the
length of its cell, stem of the former slightly longer than its cell, cross-vein 3—4 barely
its own length from cross-vein 2-3.

Legs dusky-brown; fore and mid legs with a small white knee spot, tarsals i-ii
with narrow white basal banding, femora with a narrow white line of scales beneath,
femora of mid legs white scaled beneath except about apical quarter; hind legs with
femora white scaled beneath except about apical quarter; all tarsi with broad white
basal banding.

Abdomen black scaled, segments three to six with white basal banding which
broadens out a little on the lateral margin, all segments of venter with white basal
banding, terminalia as illustrated.

Fig. 1.—Aédes (Finlaya) clintoni, n. sp. co terminalia.

Length: 4:0 mm. (vix), wing, 3:0 mm.

Habitat: Territory of New Guinea: Lae (K. J. Clinton).

A very distinct species clearly separated from other known species. _I have much
pleasure in naming it after its discoverer.

213

NOTES ON SOME FIJIAN MOSSES.
By WILLIAM GREENWOOD.

{Read 26th September, 1945.]

Mr. H. N. Dixon, the noted authority on mosses, died in May, 1944, ending a
correspondence with the author which had lasted twenty-five years.

Since “The Mosses of Fiji’ appeared (these PROCEEDINGS, lv, 1930, 261-302) Mr.
HE. B. Bartram has published two important papers on Fijian mosses (Contribution to the
Mosses of Fiji, Occasional Papers, Bishop Museum, Vol. xi, No. 20, 1936, and Additions
to the Mosses of Fiji, The Bryologist, Vol. xlvii, 1944). These papers brought the number
of mosses known from Fiji to about 250.

During the last few years the late Mr. H. N. Dixon has named numerous Fijian
mosses for the author and these include 12 species not previously known from Fiji.
Also a number of previously known species have been found in new localities in Fiji.
This paper is prepared so as to make this information available. All the numbers
given have been collected by the author on the island of Viti Levu and have been
seen and named by the late H. N. Dixon.

In “The Mosses of Fiji” the probable number of Fijian mosses was given as about
415. Mr. E. B. Bartram (loc. cit.) thinks this number slightly optimistic and with
his opinion the author now agrees.

I have put Tanithelium in Sematophyllaceae (not in Hypnaceae, as in “The Mosses
of Fiji’), following more recent papers.

DICRANACEAE.
DICRANOLOMA BRAUNII (C.M.) Par.
Near summit of Loma Lega Mt., Nadarivatu, c. 3,800 ft., May, 1941; (674).

DICRANELLA FLACCIDULA Mitt.
On clay banks, hills, Navua to Suva road, Namosi, c. 600 ft., May, 1943; (952).

DICRANELLA PYCNOGLOSSA (Broth.) Par.

On clay banks, Loma Lega Mt., Nadarivatu, c. 3,700 ft., May, 1941; (778). On clay
banks, hills, Navua to Suva road, Namosi, c. 600 ft., May, 1943; (904). On clay banks at
roadside, Serua hills, Serua, c. 700 ft., May, 1943; (994).

In “The Mosses of Fiji’ this species was ascribed to (Broth.) Broth. in error.

CAMPYLOPODIUM INTEGRUM (C.M.) Par.
On damp ground, Nadarivatu, c. 3,000 ft., May, 1941; (638). On clay banks, hills,
Navua to Suva road, Namosi, c. 700 ft., May, 1943; (949).

LEUCOLOMA TENUIFOLIUM Mitt.
On trees, Nadarivatu, c. 3,000 ft., May, 1941; (684). On trees, hills, Navua to Suva,
Namosi, c. 600 ft., May, 1943; (916).

CAMPYLOPUS SAMOANUS Broth.
New to Fiji. On wet rock in creek, Nadarivatu, c. 2,700 ft., May, 1941; (637). Only
known hitherto from Samoa.

LEUCOBRYACEAE.
LEUCOBRYUM SANCTUM Hampe.
On rotten wood, hills, Navua to Suva, Namosi, c. 800 ft., May, 1943; (900).

214 NOTES ON SOME FIJIAN MOSSES,

LEUCOBRYUM TAHITENSE Aongstr.
On trees, Mt. Evans, Lautoka, c. 3,600 ft., Dec., 1942; (876).

LEUCOBRYUM PUNGENS C.M.
On dead wood, Nadarivatu, c. 3,000 ft., May, 1941; (657, 658).

‘ LEUCOBRYUM SAMOANUM Fleisch. ined.

On dead wood, Nadarivatu, c. 2,800 ft., May, 1941; (655, 656). On rotten wood,
hills, Navua to Suva, Namosi, c. 800 ft., May, 1943; (950, 970).

LEUCOPHANES PUNGENS Fleisch. ined.
On rotten logs, Nadarivatu, c. 3,000 ft., May, 1941; (662). On dead wood, hills,
Navua to Suva, Namosi, c. 700 ft., May, 1943; (905). On dead wood, Serua hills, Serua,
c. 700 ft., May, 1948; (1008).

EXoODICTYON DENTATUM (Mitt.) Card.
On rotten logs, hills, Navua to Suva, Namosi, c. 700 ft., May, 1948; (965, 983). On
rotten wood, Serua hills, Serua, c. 700 ft., May, 1943; (1009).

FISSIDENTACEAE.
FISSIDENS DALTONIAEFOLIUS C.M.
On ground in forest, Sigatoka River, Sigatoka, c. 300 ft., Dec., 1940; (577).

FISSIDENS GLOSSO-BRYOIDES Dix.
On ground, Nadarivatu, c. 3,000 ft., May, 1941; (795).

FISSIDENS LAGENARIUS Mitt.
On wet rock, Mts., Lautoka, c. 1,800 ft., Sept., 1941; (827).

FISSIDENS LAUTOKENSIS Dix.

Mr. H. N. Dixon recently re-examined all his material of F. lautokensis and came
to the conclusion that, of the eleven numbers given under F. lautokensis in ‘The
Mosses of Fiji’, only two belong there—No. 184 (type) and No. 150. The remainder
belong to the following two species.

FISSIDENS ABBREVIATUS Mitt.
New to Fiji. Nos. 4, 133, 134, 147, 148, 149.

FISSIDENS PHILONOTULUS Besch.*
New to Fiji. Nos. 363, 458, 495. Previously only known from Tahiti.

FISSIDENS SYLVATICUS Griff.

New to Fiji. On wet rock, Nadarivatu, c. 2,700 ft., May, 1941; (791, 796). On wet
rock, Mt. Evans, Lautoka, c. 3,000 ft., Oct., 1942; a form with leaves strongly falcate,
moist and dry; (854). On trunks of tree-ferns, hills, Navua to Suva, Namosi, ec. 700 ft.,
May, 1943; (924, 962). On wet rock, Serua hills, Serua, c. 700 ft., May, 1943; (1004).

CALYMPERACEAE.
SYRRHOPODON MAMILLATUS C.M.
On rotten logs, Serua hills, Serua, c. 700 ft., May, 1943; (1000).

SYRRHOPODON LUTEUS (Mitt.) Jaeg.

On tree trunks, hills, Navua to Suva, Namosi, c. 800 ft., May, 1943; (902, 978).
Milne’s specimen from Ovalau was collected on stones—an unusual habitat.

SYRRHOPODON LONGIFOLIUS (Mitt.) Dix.
On tree trunks, Mt. Evans, Lautoka, ec. 3,000 ft., Oct., 1942; (868).

SYRRHOPODON LAEVIGATUS Mitt.
On dead wood, hills, Navua to Suva, Namosi, ec. 800 ft., May, 1943; (971).

BY WILLIAM GREENWOOD. 215

CALYMPERES TENERUM C.M.
On rotten wood, hills, Navua to Suva, Namosi, c. 800 ft., May, 1943; (956).

CALYMPERES MOLLUCCENSE Schwaeer.
On dead wood, hills, Navua to Suva, Namosi, ec. 700 ft., May, 1943; (901, 903).

CALYMPERES LORIFOLIUM Mitt.
On rotten wood, hills, Navua to Suva, Namosi, c. 800 ft., May, 1943; (964).

CALYMPERES MARGINATUM Dix.
On rotten wood, hills, Navua to Suva, Namosi, c. 800 ft., May, 1943; (967).

POTTIACEAE.
RHAMPHIDIUM VEITCHII Dix.
On clay bank, Loma Lega Mt., Nadarivatu, c. 3,800 ft., May, 1941; (603). On clay
benk, Serua hills, Serua, c. 700 ft., May, 1943; (1003).

HYOPHILA BERUENSIS Dix.
New to Fiji. On rocks, Mt. Evans, Lautoka, c. 3,000 ft., Oct., 1942; (859b). Only
known previously from the Gilbert Islands, on coral rocks near sea-level.

BRYACEAE.
BRACHYMENIUM NEPALENSE Hook.
New to Fiji. On rotten stump, Loma Lega Mt., Nadarivatu, c. 3,900 ft., May, 1941;
(786). In 1943 it was recorded for the first time from New Guinea and up till then
had not been known east of the Philippines.

BRYUM VITIANUM Dix.
On rocks, Nadarivatu, c. 2,700 ft., May, 1941; (605, 624). On dry banks, hills, Navua
to Suva, Namosi, c. 700 ft., May, 1943; (896).

BrRYUM GREENWOODII Dix.
On wet rocks, Nadarivatu, c. 2,800 ft., May, 1941; (789).

BRYUM WEBERACEUM Besch.
New to Fiji. On rocks in shade, Mts., Lautoka, c. 1,800 ft., May, 1942; (842, 846).
Previously known from Tahiti.

RHIZOGONIACEAE.
RHIZOGONIUM SETOSUM Mitt.
On tree trunks, hills, Navua to Suva, Namosi, c. 800 ft., May, 1943. A few stems
mixed with Syrrhopodon luteus (902). Evidently rare, as this is only the second
collection in Fiji.

BARTRAMIACEAE.
PHILONOTIS ETESSEI Broth. and Par.
Nadarivatu, c. 3,000 ft., May, 1941; (790).

CYRTOPODACEAE.
BESCHERELLEA CRYPHAEOIDES (C.M.) Fleisch.
On tree trunks, Nadarivatu, c. 3,000 ft., May, 1941; (677).

PTEROBRYACEAE.
EWNDOTRICHELLA GRAEFFEANA C.M.
On trees, Nadarivatu, c. 3,000 ft., May, 1941; (664).

EUPTYCHIUM SETIGERUM (Sull.), Broth.
On trees, hills, Navua to Suva, Namosi, c. 800 ft., May, 1943; (910).

216 NOTES ON SOME FIJIAN MOSSES, ~

SYMPHYSODON VITIANUS (Sull.) Broth.
On trees, hills, Navua to Suva, Namosi, c. 700 ft., May, 1943; (914, 953).

METEORIACEAE.
PAPILLARIA CROCEA (Hampe) Jaeg.
On trees, Nadarivatu, c. 3,000 ft., May, 1941; (749), a form with dense, short

branching.

PAPILLARIA INTRICATA (Mitt.) Jaeg.
On trees, Nadarivatu, c. 3,000 ft., May, 1941; (589, 751).

PAPILLARIA PELLUCIDA Broth. and Watts.
On trees, Nadarivatu, c. 2,700 ft., May, 1941; (773), a colour variation.

MeETEORIUM MIQUELIANUM (C.M.) Fleisch.
On trees, Nadarivatu, c. 3,000 ft., May, 1941; (743, 800).

FLORIBUNDARIA FLORIBUNDA (Doz. and Molk.) Fleisch.
On trees, Nadarivatu, c. 3,000 ft., May, 1941; (799).

FLORIBUNDARIA AERUGINOSA (Mitt.) Fleisch.
On trees, Nadarivatu, c. 3,000 ft., May, 1941; (628).

AEROBRYOPSIS VITIANA (Sull.) Fleisch.
On trees, Nadarivatu, c. 3,000 ft., May, 1941; (631).

AEROBRYOPSIS LONGISSIMA (Doz. and Molk.) Fleisch.
On dead wood, Nadarivatu, c. 3,000 ft., May, 1941; (630, 679).

AEROBRYOPIS STRIATULA (Mitt.) Broth.
New to Fiji. On dead wood, Nadarivatu, c. 2,700 ft., May, 1941; (782). On humus,
Loma Lega Mt., c. 3,900 ft., May, 1941; (588). Known previously from the Austral
Islands and New Hebrides.

NECKERACEAE.
NECKEROPSIS LEPINEANA (Mont.) Fleisch.
On trunks of trees, hills, Navua to Suva, Namosi, c. 700 ft., May, 1943; (957, 928 a
slender form).

HIMANTOCLADIUM IMPLANUM (Mitt.) Fleisch. ‘
On wet rocks in creek, Sigatoka River valley, Sigatoka, c. 500 ft., Dec., 1940; (586).

HOOKERIACEAE.
DISTICHOPHYLLUM VITIANUM (Sull.) Besch.
On decomposing rock, hills Navua to Suva, Namosi, c. 700 ft., May, 1943; (980).
On dead wood, Serua hills, Serua, c. 700 ft., May, 1943; (993).

DISTICHOPHYLLUM FLAVESCENS (Mitt.) Par.
On rotten wood, Nadarivatu, c. 3,000 ft., May, 1941; (816).

HYPOPTERYGIACEAE.
HYPOPTERYGIUM TAHITENSE Aongstr.
On dead wood, Nadarivatu, c. 3,000 ft., May, 1941; (643, 676).

HYPOPTERYGIUM SEMIMARGINATULUM C.M.
The specific name of this moss is incorrectly given in “The Mosses of Fiji’ and
“Paris Index’’.

RHACOPILACEAE.
RHACOPILUM PACIFICUM Besch.
On wet rocks, Nadarivatu, c. 3,000 ft., May, 1941; (693).

BY WILLIAM. GREENWOOD. 217

THUIDIACEAE.
CLAOPODIUM NERVOSUM (Harv.) Fleisch. (C. AMBLYSTEGIOIDES Dix.).
On dead wood, Nadarivatu, c. 3,000 ft., May, 1941; (620, 767). In naming No. 767
Mr. Dixon wrote: “This is a form of the plant which I named C. amblystegioides; but
it varies so much that I think it must be considered a form of the Indian species.”

THUIDIUM CYMBIFOLIUM (Doz. and Molk.) Bry. Jav.
On dead wood, Nadarivatu, c. 3,000 ft., May, 1941; (651, 653).

THUIDIUM MEYENIANUM (Hampe) Bry. Jay.
On dead wood, Nadarivatu, c. 2,700 ft., May, 1941; (808b, with Vesicularia
calodictyon).

THUIDIUM GLAUCINOIDES Broth.
On dead wood, Nadarivatu, c. 2,700 ft., May, 1941; (780).

HYPNACEAE.
_ ETROPOTHECIUM PERCOMPLANATUM Broth.
On wet bank, Nadarivatu, c. 3,000 ft., May, 1941; (592).

ECTROPOTHECIUM PERCOMPLANATUM Broth. var. FALCATUM Dix.
On dead wood, Nadarivatu, c. 3,000 ft., May, 1941; (698). On dead wood, hills,
Navua to Suva, Namosi, c. 700 ft., May, 1943; (930, 943).

ECTROPOTHECIUM INCUBANS (Reinw. and Hornsch.) Jaeg.
New to Fiji. On rotten wood, Serua hills, Serua, c. 700 ft., May, 1943; (1012).
Known from Ceylon, Java and Sumatra.

ECTROPOTHECIUM SODALE (Sull.) Mitt.
On rotten wood, Nadarivatu, c. 3,000 ft., May, 1941; (594).

ECTROPOTHECIUM PACIFICUM Mitt.
On rotten log, Serua hills, Serua, c. 700 ft., May, 1943; (986).

ECTROPOTHECIUM TUTUILUM (Sull.) Mitt.
On rotten log, hills Navua to Suva, Namosi, ec. 800 ft., May, 19438; (912).

ECTROPOTHECIUM ADNATUM Broth.
On rotten wood, Sigatoka River valley, Sigatoka, c. 500 ft., Dec., 1940; (576). On
rotten wood, Nadarivatu, c. 2,700 ft., May, 1941; (752).

ECTROPOTHECIUM MALACOBLASTUM (C.M.) Par.
On wet rocks, Serua hills, Serua, c. 700 ft., May, 19438; (1001).

VESICULARIA CALODICTYON (C.M.) Broth.
On submerged rocks in creek, Sigatoka River valley, Sigatoka, c. 500 ft., Dec.,
1940; (584).

LEUCOMIACEAE.
LEUCOMIUM ANEURODICTYON (C.M.) Jaeg. (L. DEBILE (Sull.) Mitt.).
In a letter (Sept., 1941), Mr. Dixon wrote: ‘‘The plants which I have referred to
Leucomium debile must be named L. aneurodictyon (C.M.) Jaeg. as Fleischer found
that L. debile was only a synonym of this.”

SEMATOPHYLLACEAE.
MEIOTHECIUM RECHINGERI Broth.
New to Fiji. On rotten wood, hills, Navua to Suva, Namosi, c. 800 ft., May,
1943; (894).
In naming this species Mr. Dixon wrote: “This is certainly the plant referred by
Bartram to this species (Polynesian Mosses), and has the two forms of leaf very
marked, while the colour is a bright green, as in Brotherus’ original plant.”

218 NOTES ON SOME FIJIAN MOSSES.

MEIOTHECIUM MICROCARPUM (Harv.) Mitt.
On dead wood, hills, Navua to Suva, Namosi, ec. 700 ft., May, 1943; (931). On
trees, Navua, Serua district, near sea-level, May, 1943; (945, a robust, deep green form).

ACROPORIUM SUBULATUM (Hampe) Fleisch.
New to Fiji. On trees, mts., Lautoka, c. 1,800 ft., Aug., 1941; (818).
Known previously from Borneo, Siam, Malay Peninsula and Philippines.

RHAPHIDOSTICHUM THELIPORUM (C.M.) Broth.
On dead wood, mts., Lautoka, c. 2,000 ft., May, 1942; (845).

TRICHOSTELEUM HAMATUM (Doz.-and Molk.) Jaeg.
On rotten wood, hills, Navua to Suva, Namosi, c. 800 ft., May, 1943; (898).

TRICHOSTELEUM Boscuit (Doz. and Molk.) Jaeg.
On rotten logs, hills, Navua to Suva, Namosi, c. 900 ft., May, 1943; (906, 974).

TAXITHELIUM PAPILLATUM (Harv.) Broth.

On trees, hills, Navua to Suva, Namosi, ec. 700 ft., May, 1943; (975, with Taxithelium
samoanum ). °

TAXITHELIUM SAMOANUM (Mitt.) Broth.
On trees, hills, Navua to Suva, Namosi, c. 700 ft., May, 1943; (975, with Taxithelium
papillatum). On dead wood, Serua hills, Serua, c. 700 ft., May, 1943; (992).

TAXITHELIUM LINDBERGII (Bry. Jav.) Ren. and Card.
On rotten wood, Mt. Evans, Lautoka, c. 3,600 ft., Dec., 1942; (872, a small form).
On dead wood, hills, Navua to Suva, Namosi, c. 800 ft., May, 1943; (954).

TAXITHELIUM TENUISETUM (Sull.) Mitt.
On trees, Serua hills, Serua, e. 700 ft., May, 1943; (999).

TAXITHELIUM PROTENSUM Dix.
On dead wood, hills Navua to Suva, Namosi, c. 800 ft., May, 1943; (959).

BRACHYTHECIACEAE.
RHYNCHOSTEGIUM JAVANICUM (Bel.) Besch.
New to Fiji. On dead wood, Nadarivatu, c. 3,000 ft., May, 1941; (6138, with
Vesicularia calodictyon Broth).

HYPNODENDRACEAE.
HYPNODENDRON VITIENSE (C.M.) Mitt.
On trees, Nadarivatu, c. 3,000 ft., May, 1941; (680, a dark green form).

MNIODENDRON TAHITICUM Besch.
On rotten wood, Nadarivatu, c. 3,000 ft., May, 1941; (601, 758).

POLYTRICHACEAE.
POGONATUM GRAEFFEANUM (C.M.) Jaeg.

On wet soil, Nadarivatu, c. 2,700 ft., May, 1941; (619, a form with short, crowded
leaves). On clay bank, Loma Lega Mt., Nadarivatu, c. 3,900 ft., May, 1941; (681). On
clay banks, hills, Navua to Suva road, Namosi, ec. 600 ft., May, 1948; (9384). On clay
banks at roadside, Serua hills, Serua, ec. 600 ft., May, 1943; (995).

PSEUDORHACELOPUS PHILIPPINENSIS Broth.
On clay banks, hills, Navua to Suva road, Namosi, c. 600 ft., May, 1943; (895). On
clay banks at roadside, Serua hills, Serua, c. 700 ft., May, 1943; (1005, 1006).

219

NOTES ON AUSTRALIAN MOSQUITOES (DIPTERA, CULICIDAE).
PART VI. THE GENUS TRIPTEROIDES IN THE AUSTRALASIAN REGION.
By Davip J. Les, B.Sc.
(Plates xii—xiii; sixty-nine Text-figures. )

[Read 31st October, 1945.]

Contents.
Page.
Introduction Ore. Sess Tei Je) tle (cad ed Eee ae Me RN ALi tA eo an A A ae nM AT 219
Genus Tripteroides.
Synonymy EES Ee PVM LE cea yeni vies tw PeAche 4 Ole React, CH Moi ate OES ahs besb ed © eee) Cites: Aeeepore Hordes view MAE 222
Characters of the genus ae gay eames ee een CN ss UME itn Baten Waeeat a YD Sr hemes | ENC ma 222
Relation to other genera .. ret Sade Se ENS on ae Aad ones oe weaten ian EIA Ray i) ice 223
Subgeneric divisions Btls ieee a 20 ote EA eigen hea cit Net cant det ee aU a SE CO OL aR 224
List of Australasian species He SPATE Arh foon Lee Eh etn ERLE SA ob eneas dare VE ea 225
Ixeys to Australasian species of TMT eROId es:
Adults Rar Meee eee acerca Satin ce ig yy ce ivan Maen a hin Set On tape Tea et Bu 226
Larvae os RCM eS arsed OS Sele ert ee ee aS oD ee nemo ily eed eae Rese, Oe Cie aE A a 228
Descriptions of species.
Ornamented species GS SEULs cals Sete Oot OTE Lea ee 8b Clee Tea Migs AOL Cee Dee ease erg Re Mh AC 229
SUDSCHUSeMAOTGOCLEAI Gates «cat okie pee RUNES ckeh ko ee pee eee Hore oaceg Vee Boe 229
SUI emu S Helen tDtCiNOULE Sencar re arn eR) usyue fsa vce ae hee teen te oe ac ae 230
INDULGIU ER CEN SLOW P raceme tren eed ee hese mitt, s Uae Now toe DONE MI EIN) el) ete Pa Ate 230
WmornannentecdlspeClesiyant murine ciatn Seen ena tar Mer SA DERN eid tM RSE AUER en > ek 7 Pas enee 242
SUI EMU Swe CIHISOUGG@ arin Miter duets cadet pest haa ore ty. bo tokea wee, als bed Eee aie 242
LD ES- SOUP Mary dest halts, EE Yee hehe. steam these Bese TASB cpu aba Phe sap taut Sete pipes 242
Vanleewweni-group ssi Bs SS eee Pann escent iby TONG eRe RAR! aoe ae MERI 252
Species not placed in any SIrasanS TAT AN pea Bee phe hes. Fa ae Nao Hea tse ena 260
Subgenus Mimeteomyia BGcva case esa ot er Ue Rane Brn mre Miter clay it ae LATE REET eae ween, 261
Atripes-group .. .. Serle PONE Beek Cds See tel cone sil Ant = aa an Ab Oprtl 261
Caledonica-group.. i, “fe 4 Ee ee Ley aa gs 5% AR bP IED tae 264
Obscura-group Soa aN ys SOON UE TS URES wT RE ite VORA Net a ney ae ed Mena Peas Se 273
Acknowledgements AGS, Obs LAS RCRA COREE EAI: trans ARR oat an De yi em RUE EEL aoe 274
References Pa SI EAvE Nance seas trope HCA ings armen en 274
Index to species ae Tripter Sides Rentioned in ie eee SRD ACLs kc en WA Ren ate aagS Gane a a7 ERE E 275
INTRODUCTION.

The present paper comprises a revision of the Australasian members of the genus
Tripteroides, a discussion of subgeneric divisions, the description of fourteen new
species, and keys for the identification of both adults and larvae of species known to
- occur within the region. As much information as is available on the biology of the
various species is also included.

The known distribution of the genus extends from India and Ceylon through the
Oriental and Australasian Regions into the Pacific as far as New Zealand and Fiji.
Southward it extends to Tasmania (one species) and northward to Japan (also only
one species recorded). Of the 60 or so species or varieties so far described (including
those described as new in this paper) 40 are recorded from the Australasian Region,
some five or six species occur in India or Ceylon, five in the Philippines, and nine in
the Sunda Islands. Vertically the genus is known to occur from sea-level to a height
of at least 2,400 metres (Kdwards, 1927).

The circumstances surrounding the collection of most of the New Guinea species
are noteworthy. In 1932, a half-day’s collecting by Dr. de Rook and Dr. S. L. Brug

x

220 NOTES ON AUSTRALIAN MOSQUITOES. VI,

at Tanah Merah yielded five new species of nepenthicolous Tripteroides, later described
by Brug in 1934. Previously a collection made by Dr. W. Docters van Leeuwin near the
Rouffaer R. and on the Nassau Mts., both areas in the heart of Dutch New Guinea, also
yielded five new species of Tripteroides, but even more interesting is the fact that
only one other known species of Tripteroides and one of Armigeres were included in
this collection. Recently Lieut.-Col. W. V. King and his assistants at Hollandia (U.S.
Army 19th Medical General Laboratory) have, over a period of some four months, made
an extensive collection of Tripteroides and have thus provided the majority of the
species herein described as new. ;

‘It seems not unlikely, then, that further collecting, perhaps particularly in the
interior of New Guinea, should bring to light many more as yet unknown species of
this genus. Borneo also, since it is the centre of distribution of the plant genus
Nepenthes (Lioyd, 1942, p. 51), may possibly be more fruitful of nepenthicolous species
than present records indicate.

In habitat the many species conform to a common type. Most species breed in the
pitchers of the insectivorous pitcher plants, Nepenthes spp. (Brug, 1934, and others),
one has been recorded from the floral bracts of Curcuma sp. (Zingiberaceae) (Brug,
1934), another in the leaf axils of an Aroid (Paine and Edwards, 1929), others in
bamboo stumps (Brug, 1934, and others), and others in treeholes, and as is often the
case with mosquitoes whose breeding specializations are towards small accumulations
of water in plants as contrasted with ground pools, a few species are to be found in
coconut husks (Paine and Edwards, 1929; Lee, 1944) and a few have adopted a
semi-domestic habitat and are found in small artificial containers (tins) or even barrels
and large tanks (Graham, 1929; Lee, 1944). One of the Australian species has also
been recorded from rock pools (Lee, 1944).

Since so many of the known species are nepenthicolous, the question arises as to
how they are able to go through the whole of their larval development in a fluid
capable of digesting most insects. This problem has been reviewed by Lloyd (1942,
p. 79), and at present it appears that some authors consider that such larvae are
capable of forming an antipepsin or that peptic digestion is retarded by the presence of
neutral salts in the larvae. Others consider that the evidence for the presence of an
antipepsin is not conclusive and cannot see that any special problem is involved, since
internal parasites of animals are capable of existing in digestive fluids. One may
assume, however, that any dead Tripteroides larvae would be subject to digestive action.
Possibly the apparently predatory larvae of many of the nepenthicolous species may also
assist in providing material for digestion by the plant. ,

However, whether or not any problem is involved, it must be pointed out that
little if anything has been recorded as to the nature of the association of Tripteroides
to particular species of Nepenthes. Seldom is the species of Nepenthes recorded in
which larvae have been found. Naturally we do not expect to find larvae in the fluid
of the pitcher before that organ has opened, but we do not know how long it is before
larvae do appear. The length of time a pitcher has been open may affect the digestive
capabilities of the fluid enclosed. Some species have the pitcher hooded in such a way
that rain cannot possibly enter, others are at least partially open to rain, but whether
Tripteroides larvae are restricted to any particular type or are to be found in all types
is not yet clear. So far no Tvripteroides, nor indeed any mosquitoes, have been found
in the West Australian Cephalotus. In any case some interesting field observations
are still to be made.*

Quite a number of species are known to bite man, and biting records are almost
invariably diurnal; one (probably 7. punctolateralis) on occasion assumes the proportion
of a household pest, particularly in isolated settlements in the north-west of Australia.
Others occur in such out of the way places that it seems likely that their attacks on
man are purely fortuitous and that other animals are involved as the source of their

* Other genera of mosquitoes have also been recorded from Nepenthes, particularly
Megarhinus and certain species of Culex, and certain other Diptera, including several Phorids
and a Chironomid, are among the list of nepenthebionts.

BY DAVID J. LEE. 22!

blood meal. In New Guinea, 7. bimaculipes at least is reasonably common in the bush
and bites fairly freely. It was collected in some numbers at Lae by D. O. Atherton
in 1944 with Aédes scutellaris hebrideus* Edw., Armigeres milnensis Lee, Armigeres
breinli Tayl., and Aédes aurimargo Edw. when experiments were planned to disclose
the vector of dengue fever in areas where Aédes aegypti Linn. was absent. Had
T. bimaculipes not been a common sylvan species at the time it would not have been
so well represented in these collections. Nevertheless there is no existing evidence to
suggest that any member of the genus Tvripteroides is in any way concerned in the
transmission of diseases of man.

Some of the members of the genus are strikingly ornamented species, the others
are almost equally conspicuous for their lack of ornamentation. Both ornamented and
unornamented species are closely similar in adult structural characters, although wider
differentiation is found in larval characters. Certain subdivisions of subgeneric value
appear obvious from a study of New Guinea species, but the occurrence of annectant
forms in the Oriental Region and on islands of the Pacific interfere with what appear
clearly defined natural divisions in the former region. Specific identification within
any particular group depends largely on characters which would be considered obscure
in many other genera, but although the structure of the male terminalia conforms to a
simple pattern in all species, the form of the ninth tergite is subject to valuable specific
variations. The range of variation in larval characters within a species is rather great
but the major modifications of larval structure are perhaps the most useful as a basis
for subgeneric divisions. Also in the adults the form of the wing scaling follows several
distinct patterns, which are of considerable use in the grouping of species.

Specifically, amongst the ornamented species, one looks in particular to the colour
of the thoracic integument, the colour of the scutal scales, the form of the scaling on
the anterior and posterior pronota, and the abdominal adornment, in the identification
of individual species. Amongst the unornamented species the length of the proboscis
and palpi, the form of wing scaling and scutal scaling and the presence or absence of
dorsocentral, prescutellar, posterior pronotal, upper sternopleural and the number of
spiracular bristles are characters of importance. The shape of the pale lateral abdominal
patches and the colour of the scales of the posterior pronota are also useful. As
indicated above, reference to hypopygial characters and to those of the larvae is always
helpful in establishing the identity of a species.

Although the larvae of all species have the thoracic and abdominal hairs mostly -
developed as stellate tufts, there is quite a wide variation in the degree of development.
This, however, is seldom of specific importance. Some species have the clypeal spines.
rather strongly thickened, most have the dorsal head hairs simple, but if branched
hairs are present they are usually useful diagnostically. The most striking character
of the larval head is found in those species in which the maxillay is modified to a
large, strongly-developed, apparently predatory, clasping organ. Other species have
hair 7 of the mesothorax and metathorax modified as a prominent thickened spine.
On the terminal segments the development of the lateral comb is particularly useful
(although the variation within a species is at times misleading), the branching of the
pentad hairs is occasionally important, and the development of the pecten, the branching

*T have used the subspecies name hebrideus to indicate that the New Guinea form is the
same as that subspecies. Further work, however, will probably disclose that either zonatipes
(Walk.) or scutellaris (Walk.) will have priority.

7 The function of this modified maxilla is not yet clear. It seems obviously an organ
associated with predation, and Paine and Edwards (1929, p. 305) note that the larvae of the
species called by them T. filipes (see below, page 246), were predacious and attacked each
other in captivity; and Brug (1934) has suggested that the maxillae may serve to destroy the
bodies of metallic green ants which are found in numbers in Nepenthes bowls. On the other
hand an obviously similar structure in some species of Trichoprosopon, e.g., T. rapax (D. & K.)
from Brazil, is associated with predation on other sabethine species and species of Culex living
in the leaf bases of Bromeliaceae. It would be most interesting to disclose the use to which this
structure is put, particularly as it is so different from the usual modifications for predation
found in other culicines, as, for example, the pectinate mouth-brushes of Aédes alternans West.,
Culex halifaxii Theo., and more particularly of Megarhinus spp.

222 NOTES ON AUSTRALIAN MOSQUITOES. VI,
of the siphonal hairs and the saddle hair are also on occasion used in specific
identification. Quite useful at times is the long hair on the ventral siphonal valves.

Finally, difficulty in the identification of species has probably hitherto militated
against the collection of biological information concerning individual species. It is
hoped that more may now be contributed along these lines and field studies- of the
genus, particularly relevant to distribution and specializations in habitat, should yield
most interesting information. It seems not unlikely that a high degree of specialization
in the choice of larval habitat will actually be found; at least the evidence so far
available does suggest this. It will be interesting to see how far correlations between
species and their larval habitat are able to be carried, for at present it appears that
the most widespread species are those whose breeding places are similarly ubiquitous,
namely, treeholes and semi-domestic situations.

The present paper is the result of the examination of some 500 adult specimens
and a comparable number of larvae or associated larval skins. On the basis of the
proposed classification it has been possible to identify almost all the specimens available.
Some of the unnamed material constitutes apparently new species, but the
specimens available have not been satisfactory for adequate description although a
brief account of them is included in the text. Again, when female specimens,
unassociated with males or larvae, have originated from isolated localities, such as
small islands, it has been deemed wise to withhold identification pending the receipt
of material of greater diagnostic value.

Some corrections of my interpretations of inadequately described species are almost
inevitable, but it is hoped that the present revision will materially assist in arriving at
a satisfactory classification of this previously neglected genus.

Genus TrRIPTEROIDES Giles.
SYNONYMY.

TRIPTEROIDES Giles 1904. J. Trop. Med., 7: 369. Genotype: Runchomyia philippinensis Giles
(= nitidiventer Giles).

RACHIONOTOMYIA Theobald 1905. J. Bombay Nat. Hist. Soc., 16: 248. Genotype: R. ceylonensis
Theo. (= aranoides Theo.).

POLYLEPIDOMYIA Theobald 1905. Ann. Mus. Nat. Hung., 3: 118. Genotype: P. argenteiventris
Theo..

COLONEMYIA Leicester 1908. Culicidae of Malaya in Stud. Inst Fed. Malay Stat., 3: 273.
Genotype: C. coeruleocephala Leic.

SKBIROMYIA Leicester 1908. Loc. cit.: 248. Genotype: S. fusca Leic. (= aranoides Theo.).

SeuAMOMYIA Theobald 1910. Rec. Ind. Mus., 4: 28. Genotype: S. inornata Theo. (= aranoidés

Theo. ).

RACHISOURA Theobald 1910. Monogr. Cul., 5: 207. Genotype: R. sylvestris Theo. (= filipes
Walk.).

MIMETEOMYIA Theobald 1910. Loe. cit.: 210. Genotype: M. apicotriangulata Theo. (= atripes
Sk.). : ;

TRICHOLEPTOMYIA Dyar & Shannon 1925. Insec. Inscit. menst., 13: 72. Genotype: Wyeomyia
nepenthicola Banks.
MAORIGOELDIA Hdwards 1930. Bull. ent. Res., 21: 302. Genotype: Culex argyropus Walk.

CHARACTERS OF THE GENUS.
Adult.

Head: The scales on the vertex are all broad and flat and there is a row of. upright
forked scales on the nape. The eyes touch for a long space above the antennae and
there is a pair of strong occipital bristles widely separated from the smaller lower
orbital ones. The proboscis varies from short and thick to very long and slender,
the mouthparts are normal, the palpi variable in length in both sexes. The male
antennae are distinctly plumose with the last two segments elongated. In the female
the flagellar segments are subequal and the verticillate hairs rather long.

Thorax: The pronotal lobes are widely separated, the scutum has central and
dorsocentral bristles in one species, dorsocentrals only in others and others again have
no dorsocentrals. Most species have the prescutellar group represented by one or more
pairs of bristles. A single posterior pronotal may be present (three or four in one
species) but others have no such bristle. Spiracular bristles are always present, but

BY DAVID J. LEER. 223

no postspiraculars and at most one or two upper sternopleurals, no lower mesepimerals
and a few upper mesepimerals and subalars. The scutal scaling varies from sparse
narrow scales to a dense covering of broad spindle-shaped scales. The pleura are
largely clothed with broad appressed scales, silvery in some species, white or creamy
in others. The scutellar scales are always broad and flat. The postnotum is usually
furrowed slightly on either side of the midline, the furrows approaching the midline
distally. Except in a very few species the postnotum is bare (minute hairs or even
bristles are occasionally present).

Legs: The legs are slender with a few short tibial bristles, the hind tibiae are
usually shorter than those of the fore or mid leg, and the first- hind tarsal segment is
usually longer than the tibia. The fore claws of the male are unequal and all claws
of the female are simple; the hind tarsi have only one claw in some species. No
pulvilli are present.

Wings: The upper fork cell is always longer than its stem and narrowed apically.
The posterior crossvein is usually situated well before the middle crossvein and the
anal vein extends well beyond the base of m-cu on Cu,.* There are no hairs on either
surface of the stem vein and at least a partial fringe is present on the squame.

Abdomen: The abdomen appears sleek with few hairs except at the tip, both the
eighth tergite and sternite of the female being broad and very bristly. The male hypo-
pygium is prominent in most species. The coxite has a slight basal lobe bearing
bristles, and the style is simple, long and slender, with a small terminal spine. The
ninth tergite is prominent, usually divided into two distinct lobes (in some species a
complete fusion of these lobes has occurred) bearing a variable number of strong
bristles. The paraprocts are strongly chitinized at the tip, usually with several teeth,
and the phallosome is a simple incomplete tube sometimes bearing weakly chitinized
internal teeth:

Larva.

The head is small, the antennae always short and bare with the shaft hair small
and placed beyond the middle. The mouthparts are usually unmodified except in the
subgenus Rachisoura wherein the maxilla is strongly modified. The thorax and
abdomen are usually covered with stellate hairs. The metathorax has in most groups
a long and strong dorsolateral spine inserted on a plate which itself bears a small
spine. The lateral comb consists of a row of teeth (except JT. argyropus) which may
or may not be attached to a plate. The first pentad hair is more strongly developed
than the third; the siphon is variable in length, without an acus, and with numerous
scattered hairs and spines, the latter forming a rather irregular “false pecten’. The
anal segment has only one pair of ventral tufts, there is usually a fringe of spines on
the distal margin of the saddle and the dorsal subcaudal hair is branched and the
ventral single. The anal papillae are well developed, often long.

RELATION TO OTHER GENERA.

The genus Tripteroides comes closest to the American genus Tvrichoprosopon (as
defined by Lane and Cerqueira, 1942) and indeed there is little to separate the two.
The range of variation of adult characters appears to be such that all variations found
in Tripteroides are within the range of those of Trichoprosopon. Of particular
importance in linking the two are the characters of the male terminalia which are
identical in structure, except that somewhat more diversity in the basal lobe of the
coxite is to be found in Trichoprosopon. In larval characters there is again a decided
parallel in the development, in some species of both genera, of enlarged, toothed
maxillae. On the other hand there does appear to be a constant difference in that a
pecten is always present in Tvripteroides and absent in Trichoprosopon. This does
provide an argument for separating the two genera, and apart from this it is always
desirable, for the sake of simplified regional taxonomy, to keep separate, genera

*In conformity with the terminology proposal by Lee and Woodhill (1944). Other authors
usually refer to this vein as Cu,.

224 NOTES ON AUSTRALIAN MOSQUITOES. VI,

which are quite definitely geographically discontinuous and which have no species
common to both regions. Were the two genera combined, the subgeneric divisions of
either would not be satisfactory for the species from the other region. This is at least
an argument against such a union and it seems likely that if the union were made
then the primary separation within the composite genus would be into two geographical
rather than morphological groups, with each group then subdivided in a manner
following the present subgeneric divisions. If this is really the result that would be
obtained, then it serves to emphasize that the evolutionary processes involved in the
two regions have followed different paths although the closeness of the relationship of
the two genera has inevitably resulted in considerable parallelism. To unite the two
genera, then, would be to unite the present ends of two diverging, but not greatly
divergent, evolutionary trends which are quite unable to come together again by any
natural means and which have enjoyed a considerable period of geographic isolation.

SUBGENERIC DIVISIONS.

Edwards (1932) has divided the genus Tripteroides into four subgenera, namely,
Maorigoeldia, Tripteroides, Rachisoura and Mimeteomyia. There is no doubt that his
divisions of the genus are useful, although his definitions do not apply in their entirety
on the data now available..

The subgenus Maorigoeldia may be retained as originally defined, since it still only
contains the one species from New Zealand. This is 7. argyropa, which is an ornate
species with three to four posterior pronotal bristles, both central and dorsocentral
bristles, and larvae without any modifications of maxilla or thoracic hairs and the
lateral comb a patch of many scales.

In the subgenus Tripteroides, Edwards included all species in which the palpi
is very short in both sexes, usually less than one-sixth that of the proboscis which in
its turn is slender and usually longer than the abdomen. There is one posterior
pronotal bristle and some of the outstanding wing scales are narrow. Most species
have dorsocentral bristles and most (all Australasian members of the subgenus) are
ornate with silvery markings on thorax, legs and abdomen and azure-blue on the head.
The larval maxillae are not specially modified and there is a modified spine on the
metathorax and sometimes also on the mesothorax. Two groups are recognized by
Edwards:

(a) aranoides-group. No dorsocentral bristles are present, the scutal scales are
often broad; the white scales of the thorax and abdomen are without silvery reflections
an‘ the femora are unspotted. No member of this group is as yet recorded from the
Australasian Region. (Hdwards included 7. argenteiventris and T. atra in this group
but the first is obviously a Mimeteomyia and the second merely a synonym of T.
bimaculipes.)

(0) nitidiventer-group. At least one pair of dorsocentral bristles are present and
the scutal scales are all narrow. The white scales of the thorax and abdomen have a
pronounced silvery lustre and the femora are usually spotted. Most species also have
the anterior part of the head azure-blue.

Rachisoura appears to be a valid subgenus based particularly on the modified larval
maxilla (much enlarged with one very strong terminal tooth and often subsidiary
smaller teeth as well). The adults correlated with such larvae have in common their
lack of ornate ornamentation together with the outstanding wing scales, at least of
veins R, to R,, dense and broad. Two groups within the subgenus are also recognizable
on adult characters:

(a) filipes-group. The members of this group have all the outstanding scales of all
veins broad and a posterior pronotal bristle is present. The length of the palpi is very
variable and there are often small postnotal hairs.

(0) vanleeuweni-group. This is characterized particularly by the wing scaling
which is broad and dense on veins R, to R, (as in the filipes-group) but on the rest
of the veins the outstanding scales are rather sparse and narrow. No posterior pronotal
bristle is present.

BY DAVID J. LEE. 225

The subgenus Mimeteomyia requires some modification if 7. obscura and T.
subobscura are to be included. It would seem best to base this subgenus on the wing
sealing particularly, and define it as follows: The outstanding wing scales are all long
and narrow, the wing scaling generally is fairly dense; the male palpi are usually almost
as long as the proboscis but may be reduced, those of the female are from 0-1 to 0:25
the length of the proboscis, which itself is variable in length. Most species have a
posterior pronotal bristle; all are dull coloured species without silvery markings. The
larvae lack modified maxillae but have mesothoracic spines in most species, both
metathoracic and mesothoracic in some, and a few have neither.

Again, the subgenus is best divided into three groups, two of which.remain almost
as defined by Edwards:

(a) atripes-group. The male palpi are long and the proboscis is not longer than
the abdomen. The mesothorax of the larya has a strong spine similar to that of the
metathorax. The lateral comb teeth are prominent and at least some of them arise from
a lateral chitinous plate.

(ob) caledonica-group. The male palpi are also long but the proboscis is distinctly
longer than the abdomen and very slender. The mesothoracic spine is not developed
and the metathoracic spine may be present or absent. The lateral comb teeth do not
arise from a chitinous plate.

(c) obscura-group. The male palpi are only one-third the length of the proboscis;
the latter is short and stout. The larval characters are not known.

All the Australasian species so far described can be relegated to one or other of the
above subgenera and groups with the exception of two, namely, 7. swbnudipennis and
T. concinna. These two species are Known from females only, and as they do not
readily fit into any of the groups as defined above, it remains to be seen, when males
and larvae of these have been discovered, whether they are deserving of a grouping of
their own and in which subgenus they should be placed.: It is not unlikely that they
will prove to be members of the aranoides-group of Tripteroides, but on the circum-
stantial evidence of their distribution it is still possible that they will prove to be
members of Rachisoura.

LIST OF AUSTRALASIAN SPECIES.

The following is a list of the described species (including those described as new
in the present paper) in their arrangement according to subgenera and groupings.

Subgenus Maorigoeldia. Subgenus Rachisoura.
T. argyropa. vanleeuweni-group.
Subgenus Tripteroides. T. kingi.
nitidiventer-group. T. pallida.
T. alboscutellata. T. pilosa.
T. bimaculipes. T. simplex.
T. brevipalpis. T. vanleeuweni.
T. distigma. Subgenus Mimeteomyia.
T. elegans. atripes-group.
T. littlechildi. T. atripes.
T. magnesiana. T. digoelensis.
T. nissanensis. T. punctolateralis.
T. purpurata. T. solomonis.
T. quasiornata. caledonica-group.
T. splendens. T. argenteiventris.
Subgenus Rachisoura. Tatra:
filipes-group. T. caledonica.
T. confusa. T. collessi.
T. filipes. T. microlepis.
T. fuliginosa. T. rotumana.
T. fuscipleura. T. tasmaniensis.
T. latisquama. obscura-group.
T. longipalpata. T. obscura,
T. papua. T. subobscura.
vanleeuweni-group. Species not placed in any subgenus.
T. bisquamata. T. concinna.

T. brevirhynchus T. subnudipennis.

226

10.

vale

12.

14.

15.

NOTES ON AUSTRALIAN MOSQUITOES. VI,

Krys to AUSTRALASIAN SPECIES OF TRIPTEROIDES.
(@) ADULTS.

Three or four posterior pronotal bristles present; wings very densely clothed
with broad scales; palpi about one-quarter the length of the proboscis
with white ring at middle; silvery markings on thorax, legs and abdomen

Ree eee oy SRO ee inne mG OL CRMC NU CoN I MyM atl lars Aes ia ig Sia cba te ae argyropa
No more than one posterior pronotal bristle; other characters various but wings
never as densely scaled and palpi without white ring at middle ....... 2

Ornate species with azure-blue band anteriorly on head, silver scaling on pleura
and abdomen, and silvery or white spots or bands on femora; palpi very
9

Shontitini Doth: (Sexes iyi es Gas BOE Pe AAC es SCA a on eed 3
Drab species with only white or creamy ornamentation, no blue scaling on head,
no silvery markings elsewhere; palpi variable ....................... 13

Scutal integument black or at least dark distally; scutal scales of uniform
colour with integument at least on distal half ........................ 4
Scutal integument pale, from yellow to light orange; in some species the scutal
scales are dark and in strong contrast to the integument, in others they are

VellowiISheSreei «PALES AL EE ED RRs Sires Ue ui Se SS re RCE NTR Mey Ua AAC IED ea 8
Pronotal lobes clothed with broad black scales only ....................... 5
Pronotal lobes clothed with narrow black scales or if some broad ones are

PLESEMt NaLEROws .Ones _pLedOminiate ease ieee ner cere 7

Posterior pronota with broad flat black scales; mesepimeron very dark but
with a small patch of silvery scales; abdominal tergite III with a very small
apical sublateral patch of silver; lobes of ninth tergite long and narrow with

four spines arising well below the distal margin ................ splendens
Posterior pronotal with: narrows Scales soniliyess ee ee ee ene eee 6
Mesepimeron shining black and bare; abdominal tergites with lateral dull black
MAT KMS. A EE ee TA ee, DR ee ER ie Ly Suedee ane een ate ae Spek eee littlechildi

Mesepimeron dark but at least partly covered with flat silvery scales: no lateral
dull black markings on abdominal tergites; lobes of ninth tergite with nine
or more irregularly arranged spines of which about three are longer and

stroniser thant the rest. tac cece chee saeice ene eo One ee TEER Cn Te Cea bimaculipes
Scutellum dark scaled; lobes of ninth tergite long and narrow with four or five
long spines arising well before the apex ..........2..9.2...5-5..... elegans
Scutellum pale scaled; lobes of ninth tergite not separated, bearing a pair of
unequalespinesonyealchwsiG eas ease ne Peniie ie nana alboscutellata
Pronotal lebes clothed with fine narrow black scaies; abdominal tergites with
lateral d whl, black Patel Sve osctsaccs snes ce oierow ie ce ie OO Dee magnesiand
Pronotal lobes clothed with broad black scales; abdominal tergites without
lateral dull. blackepatches aaais os ue ed ee oan a ea Oh Ry 9
Posterior pronota clothed with broad flat scales; scutal scaling light greenish;
outstanding wing scales rather long and narrow ................. purpurata
Posterior pronota with narrow scales (if some broad ones are present they are
THEN Thy TOUbDaaoyere, I=) SOUL KRIS WENOIOMIS sooo occaacouncavoonsuyanuove 10

Seutum clothed with greenish scales; a roundish dark brown spot above each
wing root covered with flat black scales; no silvery markings on abdomen

RR eer Eee ea RCC rs ty hates cme Heer Garner Me Crecente aig eaaeny bunt" { distigma
Seutal scales dark brown or black; abdomen with silvery markings ........ 11
Bases of fork cells level; no broad scales on posterior pronota ...... brevipalpis
Bases of fork cells not level, that of the posterior cell being distinctly nearer

base of the wing; a few broad scales on the posterior pronota ......... 12
Lobes of ninth tergite long, deeply separated, with a group of about 10 spines

distally, of which a few are longer and stronger than the rest ... quasiornata
Lobes of ninth tergite not deeply separated, rather broad, with 20 or more

fainly, -evenspinesi distally: sissy sss ew ee Hee: Ce eae nissanensis
Outstanding scales on all wing veins large and broad; wings densely scaled;

One Posterior Pronotalebristles presenta sweeter | cece ces ee eee ~14
At least some narrow outstanding scales on the posterior veins; posterior

pLonotalporistle presemt> OmPeabSenite ms saree ree teeter eats ee ornare 19
Pleural integument light in colour (yellowish-brown to medium brown) .... 15
Pleural integument dark brown or blackish-brown ........................ 18
Prescutellar bristles numerous (about 7 pairs) .................... latisquama

No more than 2 pairs of prescutellar bristles .......................-000-. 16

16.

UL.

18.

1g).

20.

ile

22.

23.

24.

25.

26.

27.

28.

29.

BY DAVID J. LEE. 227

Hind tibia about 75% length of mid; male palpi one-third length of proboscis
ER ONE RS Laser asl Arete tt at TARE APA Te a aN Ie. a Sea a tat ateee Mranar ent, cl Slama e ena rs filipes*
Hind tibia 88% or more length of mid; male palpi two-thirds length of proboscis
35GB 1S Bron ME. ONG GUS Gt RUAN lot Oy Gah GF Diptiers err Has Berlin ts aan RE ie nee eee en EE anes Kanteen ot a ee 17

Pleural integument yellowish-brown; hind tibia 94% or more length of mid;
lobes of ninth tergite long, narrow, with 7 or 8 spines increasing in length
TPO, NE) DUTIES Wo) HE) Cbisie TNEWREN Loeobdeounedtenoouanaunnadoocdc confusa

Pleural integument medium brown; hind tibia 88% length of mid; lobes of
ninth tergite of only moderate length with 8 or 9 spines of even length
mR Reem rt eae tehs are oray seu eyes Te ceang ear ear al GMA INE os RICE ee cca kee SE SEE longipalpata

Proboscis equal in length to fore femur; palpi one-sixth to one-eighth length of
proboscis; lobes of ninth tergite short with 4 or 5 long bristles distally
5 6:0 616. bE SOL Cate Oe BeOHOCs B ERORS CUORCRO EN ee RON cic es ent Pinar APR na ert ses arse fuscipleura
Proboscis slightly longer than fore femur; palpi (of male) one-quarter length
of proboscis; lobes of ninth tergite of medium length with 8 or 9 bristles
distally which are not markedly longer than the lobe itself ...... fuliginosa
Veins R,, R. and R, with broad outstanding scales; the distal portions of the
rest of the veins with narrow outstanding scales; no posterior pronotal

DIGI SEL MDLESC IN te eeearunen Get ua seat iene Arar Wiad aaudia at ep eee none Ua Rae cae Vea Te, 20
Wing scaling not as above; posterior pronotal bristle present or absent .... 25
Clypeus without scales (male palpi long in brevirhynchus) .............. Zi
Clypeus with a few scattered broad flat scales; male palpi short ......... 23
Scales on upper part of posterior pronota curved and hairlike ......... pilosa
Scales on upper part OL posterior spronotasbroadwands flat ss nme cer. cco ae 22

Female palpi one-quarter length of proboscis (male palpi three-quarters); usually
2-3 spiraculars, no prescutellars; posterior pronota dark scaled above

MR arg sear teers erate taTa Lanne ero lgs rey Sue oe lence Si Lie ior ao soins ne cE ay Saubcied enone, ase See vane bisquamata
Female palpi slightly longer; 6—8 spiraculars; one pair of prescutellars; posterior
pronotamdarksSCaledrawOviel chase ales care casio he w) ebesevenenesticue, stele seats vanleeuweni

Female palpi less than one-quarter length of proboscis; 5-6 spiraculars; 3 pairs
of prescutellars; posterior pronota entirely pale scaled
MERE ai essere aye 2) USE nue rains: ieee aed Gee Saboe pallida (females lacking clypeal scales)

Scales of posterior pronota pale below, brown on upper part; proboscis longer
than fore femur; lobes of ninth tergite moderately large, not widely separated,

with 6-8 spines rather irregularly arranged ....................... kingi
Scales of posterior pronota all pale or at most some greyish ones above; proboscis
SHVOGUE TRC ATe LONS aL STIMNU yy eeackc rene teuc seb ecs enor ei omer secenieGn aluveia mp cvencieaciors. sieici sehen enol 24

Lateral abdominal markings distinctly serrate; lobes of ninth tergite undivided
& 6b: Gus BPA Cu hc SRT MeN IS Vie MAN le BRN a Pe Pe brevirhynchus yt
Lateral abdominal markings with an indefinite margin; lobes of ninth tergite
SOWA ee Cugaiatecrsryijes, claus ies etoechecsushnuenaeeley dy ace sce acelleay = (alten suthepe onetime parecer Wels leneaes pallida

Wing scales small but broad; no narrow outstanding scales on R, and R;; some
narrow outstanding scales distally on veins R,,; to Cu,;; a posterior pronotal

WISTS RP RESECMIERE Seed SON oem ieey atest of cy Aad fonds pete oe) meee fel eared caliathetaten a Arctan ees hid 26
Wing scaling fairly dense, all outstanding scales on all veins long and narrow
HINO LUC Stage MCU TU CdS,» coh kU We oka aUte cess RCA Rinn, cube abet tdeeds sayaicnccnm: Anessa sea Atos aunts ors 27
Wprishtitorked scales of head all-yellow <).223.22....-2..5-.-... subnudipennis
Elcadmscalesisentinely: dark “at. Wap eiyscscicicie oc steret elects thls Grete cue ieth deta co elas concinna

Proboscis shorter than abdomen (usually less than the fore femur, but if, as it
may be in 7. atripes and closely related species, it is longer than the fore
femur, it still appears quite short since the fore femur itself is unusually

SHO 1st) eee eke ora ate LIM Ren HOUNRME 0 LINO elon thy Sasi ciet ses afaia) stbaneen enone 28
Proboscis longer than the abdomen and distinctly longer than the fore femur,
SISTING STE 5s ho RSet MSR RSE SCP CASIPL-O C717 TEL CNR CCEENREIC CUE Tae) i ACU ce: etre RnR OU URINE Ceres 3
Palpimormbovh sexes one-third, lenethivor PTrODOSCIS tac ee a cise ache omic cro cee ene 29
Palpi of. female less than one-third; those of male more than half length of
OG. DOS Gi Summer pemetee real te 8) re Lr cape betty Jet ani Ua cia or caren ARENT Che vat calige SRaNeaS Taea ets ary Tre i SITES oie 30

No posterior pronotal bristle; abdominal markings strongly serrate .. subobscura
Posterior pronotal bristle present; abdominal markings not serrate ..... obscura

* 7. simplex would probably key out here but it is not sufficiently described for inclusion.
+ T. papua would probably come out here but the description is not sufficiently precise for
it to be separated in the key.

al.

vo
bo

NOTES ON AUSTRALIAN MOSQUITOES. VI,

Dorsocentral bristles present at least on distal portion of scutum ........... 31
Dorsocentral bristles absent; spiracular bristles yellow; upper fork cell 3-5 x
length of its stem; border of abdominal markings straight ..... digoelensis
Dorsal head scaling very dark or black; no pale scaling at base of proboscis
(oy ieg F214 Ot ene ET cee ore On AAA Ce eGR NGAI etn AU vO MEAN aloia'e Lita ey 32
Dorsal head scaling light fawn; pale scaling at base of proboscis and palpi,
particularly \in® male eee eek eas eiaiapeseee ayerapaedeis eae teed pert ae punctolateralis
Scuitaliscalins wba ke = ein weh GRP OSe OSE URON aC eetke Urke lat 5 ieee es tN id Pe errr solomonis
Scutalscalhinig: sonomzy ta Pe isc shoe IE lis AES Ue en DU eee Sepa oa ee norte eam as atripes
Pleural integument with scaling confined to longitudinal bands ............ 34
Sealing of pleura forming an almost complete cover with no tendency to be
confined ito Joneitudinalsbands steeper 36
Seales of pleura in two longitudinal stripes on either side of a longitudinal bare
strip. Hind tarsi largely white apically .................... tasmaniensis
Seales of pleura confined to a median longitudinal band .................... at
Pale apical bands on abdomen conspicuous across dorsum ......... caledonica
Abdominal pale markings basal and lateral only ........ Ses Mae RR aa tre 3 rotumana
Venter distinethy: banded ee ois aie tiene tere iene, caste heeeuei os ee eee seedy eee collessi
Venter pales not Dan degenerates c UnRS at A/a, Ave 0 SDC eal agi CoN ape a re 37
’ Seales of scutum very small and marrow ..2....2..5....2.......... microlepis
Scales of "scutum) moderatenorn danse “Aaa ety sceneries eicrer hearer 38
Palpi of female one-sixth length of proboscis, extending beyond the clypeus for
twice its length; upper fork cell 2-5—3-0 the length of. its stem ....... atra

Palpi of female one-tenth the length of the proboscis, extending beyond clypeus
for less than twice its length; palpi of male almost as long as proboscis;
upper fork cell about 3-5 the length of its stem ............ argenteiventris

(0) LARVAE.

Since only half of the described species are Known in the larval stage this key is
of limited value and should always be used in conjunction with the larval descriptions.

il

bo

eo

10.

Ant,

Lateral comb of eighth abdominal segment a patch of over 100 scales; no

Chora cle: SPINES TDLESE Mtr hoy caies: use ice ace ee sO Cae argyropa
Lateral comb a row of scales, never arranged in a patch .................. 2
Maxillae strongly developed with one or more strong apical spines; no thoracic

SPINES: WTESESMC!? esses a Sisycesee ere veua ate Oe Gat ach i PeWaest alan Ps Gea so be ee 3
Maxillae not greatly enlarged, without strong apical spines; thoracic spines

PHESeN Lt MN M OSE SPECIES Misies Guta at ue come mene ay Scene pe per ee a sur an pe 10
Apical spine of maxilla over twice as long as body of maxilla itself, slender and.

rounded at tip; ventral portion of anal segment spinose ...... longipalpata
Apical spine of maxilla less than or scarcely longer than body of maxilla,

tapering and pointed; ventral portion of anal segment not spinose ..... 4
lateral combi comprisine onlyaay Sinelewspinewany eee etnias 5
Lateral combiot twoyon MOTZe SpINeSW aera tomer ane ti ene di
Siphonal index over 2:0; saddle hair 3—4-branched; ventral siphonal hairs scanty

MOEA SN Pear MeN a UCR T Ane iy Mate E aaa ee SUS Bly aa ey cic ese ey Giese 9 OO, 1 brevirhynchus
Siphonal index less than 2:0; saddle hair with fewer than 4 branches ...... 6
Siphonal index 1:5; 4-5 dorsal hairs on siphon; 6—9 ventral tufts; saddle hair

Wath 2) Dranchesind. amet nie Gl ONES ie oes i hla TLS AOE gate Garp a Sema PNA kingi
Siphonal index 1:0; 2-4 dorsal hairs on siphon; 6 ventral tufts; saddle hair

With Sbramchege pee cuanto toa aa suns SNE INGE oO aa Re FOR Gun pallida
Pecten\contined toy distal phialiqromasiphonery eerie eee ete ee aa a ene 8
Pecten either basal or extending from near base to apex .................. 9
Ventral siphonal valve hair with 4—5 branches; 3-5 lateral comb seales.. filipes
Ventral siphonal ‘valve: hair sine lei ace eee eek hoe oe Ric ae eee confusa

Pecten of 5-6 spines extending from base to apex; ventral siphonal valve hair
with 4 branches; lateral comb with the uppermost spine larger than the rest

Bh ea ac ee iPr a oANE aARR ETH, CML ons pe aaa al toy cba Me etn REE OR Sa ren cane EC bisquamata
Pecten of 5-9 spines on basal portion of the siphon; ventral siphonal valve with
2 branches; lateral comb spines of even size ..................-. fuscipleura
Bateralicombyarisinics tromeaachitinousuplatcwmnner tte eee een iene ist
No lateral chitinous plate on eighth abdominal segment ................... 15
Lateral plates of eighth segment fused dorsally .................... purpurata

Lateral plates! ot: fused VaWOVIE™ 6 ce ek Smears he Sete ALAN tea Seo ea 12

——E———

BY DAVID J. LEE. 229

12. Both mesothoracic and metathoracic spines present ...................05- 13
OnlvamMetathoracicuspinempneSemtG lek rleiere -nleleresed-ne) siciep elie) leslie alboscutellata

13. Lateral comb extending ventrally below the lateral chitinous plate, the spines
SEO UM Cae saa een OLIN arte ner atdtes: mens kcetet meclcrobiagley tet h eiaitel ur atueleeigelteuleyaglenslca (el wumiarey reesrel eshvests 14
Lateral comb not extending below the lateral chitinous plate, the spines sharply

HD OUMES Comer cia seat sits ete n nie cue tara och aee Los nahrd lates a glebaincurtardiesh opal eters meets solomonis
IAPeCadmhaig Aubin dss) 3-branched sna... cscs cee ce cece se cias punctolateralis
eadehaineAw4-branchedi; 3) 6-branched! shir ce sete lela) schacle ciceg ete set etene atripes

15. Hither mesothoracie or both mesothoracic and metathoracic spines present .. 16
Neither mesothoracic nor metathoracic spines present .................... 22

16. Both mesothoracic and metathoracic spines present ...... ENESCO LURID) ISA epee 17
Only a metathoracic spine present; mesothoracic hairs not obviously modified 20

time onsal hairs or siphon swath 5-6 branches) sf 52... 225.5...t66 0500086 quasiornata
Dorsal Mawes Cir Siolncm DRL joie! ~~ bscceavccocusdosenusububdonucoobuode 18

see Donsalehead hairs bifid: saddle hain bifid) (55 -hi. oe s ee ee brevipalpis
Dorsal head hairs simple; saddle hair with more than two branches ....... 19

19. Branches of stellate tufts ending in divergent points ............ bimaculipes
Branches of stellate tufts ending in non-divergent points .......... nissanensis

20. Siphon clothed with pecten-like spines over greater part of surface .... collessi
SiolaGia Wriwoa momo) jWercvusa Howes SosoccagsuonevcaveccnguuvocdcuuouOUanDE 21

Zi SiDhonalwindex 2-05 about a0) ventral) hair tuts: 2030.21... ows rotumana
SipMNo Mew wWovle< SANE Ale ats) Asie ONNES! “BG gos eooodocoDboguooGbeaG caledonica

22. All or most of the lateral comb spines sharply pointed ........ argenteiventris
Lateral comb scales all blunt and fringed ....................... tasmaniensis

DESCRIPTIONS OF SPECIES.
ORNAMENTED SPECIES.

The ornate species of Tripteroides with their bright colouration and particularly the
silver scaling of the thorax and abdomen are immediately distinguishable from the
unornamented species, which lack all ornamentation except white or creamy scaling
laterally on the thorax and abdomen. One ornate species belongs to the subgenus
Maorigoeldia and the rest to the subgenus Tvripteroides.

Subgenus Maoricortp1a Hdwards.

EDWARDS, F. W., 1930.—Bull. ent. Res., 21: 302.
, 1932.—In Wystman’s Genera Insectorum, Diptera, Culicidae, Fasc., 192: 75.

This subgenus is represented by the single species 7. argyropa (Walk.), known
only from New Zealand.

The palpi are about one-quarter the length of the proboscis in both sexes and have
a white ring at the middle. The proboscis is rather stout but longer than the abdomen.
There are three or four strong posterior pronotal bristles and the wings are densely
clothed with broad scales. Silvery markings are present on the thorax, legs and
abdomen. The larval maxilla is unmodified and there are no distinct spines on the
metathorax. The lateral comb of the eighth abdominal segment comprises a large
number (more than a hundred) of small scales forming a semi-circular patch. The
anal papillae are very large and well tracheated.

TRIPTEROIDES (MAORIGOELDIA) ARGYROPA* (Walker).

WALKER, F., 1848.—List Dipt. Brit. Mus., 1: 2 (Culex).
THEOBALD, F. V., 1901.—Monogr. Cul., 2: 264 (Uranotaenia?).
EDWARDS, F. W., 1924.—Bull. ent. Res., 14: 360 (Rachionotomyia).
GRAHAM, D. H., 1929.—Trans. N.Z. Inst.. 60: 227 (Rachionotomyia—includes larval description).
IepDWARDS, F. W., 1930.—Bull. ent. Res., 21: 302 (Rachionotomyia).
GRAHAM, D. H., 1939.—Trans. Roy. Soc. N.Z., 69: 213 (Rachionotomyia).
Type: Female in British Museum.
Type Locality: New Zealand.
Synonymy: Culex argyropus Walker 1848. Loc. cit. Uvranotaenia? argyropus Theobald
1901. Loc. cit. Rachionotomyia argyropus Edwards 1924. Loc. cit.

* Since it is obvious that the intention of authors has been for Tripteroides to be regarded
2s feminine, the change in termination is necessary for agreement with the gender of the genus.

230 NOTES ON AUSTRALIAN MOSQUITOES. VI,

DISTINCTIVE CHARACTERS.

The head is black with a conspicuous silvery pale blue band bordering the eyes.
There are fine dark hairs on the brown pedicels, the clypeus is almost black; the palpi
are black with a white band at the middle, they extend beyond the clypeus almost three
times its length and are one-fifth the length of the proboscis. The latter is rather stout,
black, distinctly longer than the abdomen and slightly longer than the fore femur.

The scutum is dark brown, scantily clothed with dark bronzy narrow scales but
with lateral borders of large silvery scales. There are numerous central and dorso-
central bristles. The pleura are dark brown, there are two to four posterior pronotal
bristles, two spiraculars and one upper sternopleural. The pronotal lobes are white
scaled and from them continues a diagonal white scaled line to the base of the
mesepimeron. A further patch of broad white scales is present on the upper part of
the mesepimeron amongst the subalar hairs.

The femora are dark with silvery white patches at about two-thirds from the base
and also at the apex. The tibiae are aiso tipped with white and the apical tarsi are
largely white.

The wings (Plate xii, a@) are very densely clothed with broad brown scales. The
fork cells are very long with the base of the anterior cell slightly nearer the base of
the wing than that of the posterior cell.

The male agrees closely with the female, the palpi are 0-225 the length of the
proboscis. The genitalia are prominent with the lobes of the ninth tergite (Text-fig. 1)
fused and bearing a very large number of dark spines.

The larva has been described by Graham (1929) and would appear to be distinct
from all other species for which the larvae are known because of the very numerous
lateral comb scales forming an irregular patch.

Biology: This species has adopted at least a semi-domestic habitat, having been
found breeding in house tanks around bush cottages, but its native habitat has not
been disclosed. It has been recorded biting man but is unusual in being nocturnal.

Distribution: The exact locality from which the type specimen came is not Known. I have
seen specimens from Auckland (J. T. Salmon) and Titirangi (S. L. Bisset) but Graham also
records it from Nelson, Wellington, Okahune and Waitakere Hills.

Subgenus TRIPTEROIDES Giles.

Epwarps, F. W., 1932.—Culicidae in Wystman’s Genera Insectorum, Fasc. 194: 77.

NITIDIVENTER-GROUP.

As indicated above; this subgenus has been divided by Edwards into two groups,
only one of which is known to occur in the Australasian Region. The group which is
our direct concern has been ealled the nitidiventer-group and includes all the ornate
species with metallic silvery scaling on the thorax and usually on the abdomen as well,
and usually with white or silvery femoral markings. All the Australasian species have
an azure-blue band at the front of the head and at least partial development of dorso-
central bristles. The wing scaling is not very dense but some of the outstanding scales
are narrow. The larvae have either a metathoracic spine or both mesothoracic and
metathoracic spines and the lateral comb teeth are numerous, long and close-set. The
palpi are very short in both sexes (usually no longer than the clypeus), the proboscis
is longer than the abdomen and slender, and a posterior pronotal bristle is present.

The subgenus Tripteroides is the most widespread within the genus and is found
in almost all areas whence the genus itself has been recorded. The nitidiventer-group
is best represented in the region from Malaya (including the Philippines) eastward.

In the arrangement of the species following below, 7. bimaculipes, T. splendens,
T. littlechildi, T. elegans and T. alboscutellata have in common the dark or even black
scutal integument and the rest of the species are distinguished by their largely light
orange or yellow scutal integument. Of these, 7. magnesiana, T. quasiornata, T.
nissanensis and T. brevipalpis have the scutal scales brown or black while the remaining
two, T. distigma and T. purpurata, have the scutal seales light greenish-yellow.

BY DAVID J. LEER. 231

TRIPTEROIDES (TRIPTEROIDES) BIMACULIPES (Theobald).
‘THEOBALD, F. V., 1905.—Ann. Mus. Nat. Hung., 3: 114 (Phoniomyia).
, 1907.—Monogr. Cul., 4: 660 (Phoniomyia).
EpWarpbs, F. W., 1924.—Buwll. ent. Res., 14: 360 (Rachionotomyia).
Brue, S. L., 19384.—Ibid., 25: 503, 509. (Figures of male genitalia and larval description from
unspecified specimens and localities. )
Lee, D. J., 1944.—Atlas of Mosquito Larvae of the Australasian Region. Australian Military

‘Forces (Restricted) : 32. (Figures of larva.)

Types: Described from three females lodged in National Museum of Hungary, Budapest.

Type locality: The type specimens came from two localities. The one listed first is Moroka
(altitude about 1,250 metres, just west of Mt. Victoria in Papua—9° 15’ S. by 147° 40’ E.). The
second is Friederich-Wilhelmshaven (now Madang). :

Note.—It does not appear that the types of this species have ever been re-examined. There
is little doubt, however, that the species commonly considered to be T. bimaculipes, by Edwards
(1924) and others following his definition of the species, has been correctly identified. It is
the most widespread ornamented species in New Guinea, a very common species and rather
catholic in the selection of breeding habitats, and occurs, in the material examined by me, from
sea-level to an altitude of at least 800 metres.

Synonymy: Phoniomyia bimaculipes Theobald 1905. Loc. cit. Rachionotomyia bimaculipes
Edwards 1924. Loc. cit. Stegomyia ornata Taylor, F. H., 1914. Trans. ent. Soc. Lond., 1914:
189. Mimeteomyia ornata Taylor, F. H., 1916. Proc. LINN. Soc. N.S.W., 41: 565. Rachiono-
tomyia ornata Edwards, F. W., 1924. Bull. ent. Res., 14: 361. Tripteroides ornata Edwards,
F. W., 1932. In Wystman’s Genera Insectorum, Diptera, Culicidae, Fase. 194: 78. T. ornata
(Tayl.) has been included in the synonymy since there seems little doubt that the type is
actually JT. bimaculipes. T. ornata came from Milne Bay, a place where TJ. bimaculipes is
commonly found, and the description of 7. ornata fits the present conception of 7. bimaculipes
quite well, the major discrepancies actually being inaccuracies in the description. The type
specimen is now badly damaged (head, wings and three legs missing, thorax and abdomen
denuded) but sufficient is present to show that the femora have silvery spots and the venter,
described as black, is actually pale scaled. Even were T. ornata actually a distinet form, it
would no longer be possible, either from the description or the type, to recognize it as such. The
type is a male, not a female as stated by Taylor (1916).

DISTINCTIVE CHARACTERS.

The rather dark scutal integument (particularly distally), the broad scaled pronotal
lobes and narrow scaled posterior pronota and the silvery scales on the mesepimeron,
together with the apical lateral silver bands on the tergites of segments III-VII
distinguish this species from its closest allies. The bent, thickened clypeal spines and
single dorsal head hairs of the larva, together with the form of the lateral comb, the
bifid dorsal siphonal hairs and the two- to three-branched saddle hair serve to distinguish
the larva.

DESCRIPTION.
Male and Female. y

Head: There is a band of azure-blue scales in front covering slightly more than
half the depth of the head. The pedicels are brown and the clypeus is dark brown, the
palpi are black, shorter than the clypeus and the proboscis is black, very long and
slender.

Thorax: The scutal integument varies from brown to almost black but is usually
dark with a lighter anterior area and clothed with fine narrow black scales. Some
broad black scales are also present among the bristles above the wing roots but they
are often difficult to discern. The pronotal lobes are yellowish-brown and clothed with
broad flat black scales and the posterior pronota are light or yellowish-brown bearing
narrow black appressed scales. There are from one to four spiracular bristles, but
usually three or four. The pleura are brown, the sternopleuron being largely covered
with appressed silver scales and the mesepimeron in specimens from eastern New
Guinea is usually about half covered with rounded fiat pale scales with silvery reflections.
The amount of mesepimeral scaling may be considerably reduced and this is common in
specimens from Hollandia. The scutellum has a patch of flat black scales on each lobe.

Legs: The legs are black except for the coxae which are yellowish with silvery
scales laterally, and the femora which have bluish-white or silvery markings. There is
a silvery white spot on the fore and mid femora just beyond the middle and a similar

232 NOTES ON AUSTRALIAN MOSQUITOES. VI,

one preapically. Anteriorly the mid femora have a silvery line from the base to almost
midway. On the hind legs this line extends beyond the middle of the femora and
there is also a silver preapical spot. ;

Wings (Plate xiii, c): The veins are clothed with small broad dark scales. The
base of the lower fork cell is closer to the base of the wing than that of the upper
fork cell which itself is about equal to its stem or even up to twice the length of its stem.

Abdomen: The abdomen is black dorsally and the venter is golden. There is a
large silver lateral area extending from base to apex of tergite II and apical lateral
silver bands on tergites III-VII.

Male Genitalia: Genitalia examined on New Guinea and northern Queensland
specimens conform generally to Brug’s figure k (1934, p. 503) rather than figure h.
It is unfortunate that localities of the specimens examined by Brug were not stated
as it is conceivable that the form represented in figure h may be distinct. (Note also
that it is not possible to reconcile his description of the larva of T. bimaculipes with
the description given by me.) The lobes of the ninth tergite (Text-fig. 8) are narrow,
deeply emarginate and with some nine to twelve spines irregularly arranged apically:
usually about three of these are longer and stronger than the rest.

Larva.

The head (Text-fig. 9) is round with strongly bent, thick clypeal spines and all the
dorsal hairs single and smooth. The thorax and abdomen are clothed with strongly
developed black stellate tufts in striking contrast to the general white body colour.
Hair 7 of the mesothorax (Text-fig. 11) is a thickened spine and hair 7 of the metathorax
a strongly chitinized stout spine with three unequal branches. The dorsal chaetotaxy
of the thorax is illustrated in Text-figure 10. The individual spines of the stellate tufts
may end in two or three divergent points. The lateral comb of the eighth abdominal
segment comprises some 18-25 strong teeth, of which the more dorsal are the largest,
the rest gradually decreasing in size ventrally. The shape of the individual teeth
(Text-fig. 13) is irregular, the more dorsal having a rather squarish basal plaque
surmounted by a strong pointed spine. Those towards the ventral surface are finely
spinose, particularly at the apex. Of the pentad hairs the first is a strongly branched
stellate tuft, the second is simple, the third three- to six-branched, the fourth simple and
the fifth bifid. The siphon is rather swollen basally and narrow apically, with ten to
twelve bifid ventral tufts and about ten pecten spines (Text-fig. 14), and there is an
irregular series of dorsal and dorsolateral hairs which are practically always bifid
(occasionally a single hair is seen). The saddle of the anal segment is strongly
chitinized but not complete ventrally. Its distal margin is invested with a row of
spines, the saddle hair is bifid or trifid and the ventral beard is a single four- or
five-branched hair. The dorsal subcaudal tuft is usually five-branched and the ventral
subcaudal single. The terminal segments are illustrated in Text-figure 12.

Note.—Brug describes the larva of this species as having the dorsal hairs of the
siphon two- to six-branched. The many larvae examined by me from such widely
separated places as Cairns, Milne Bay and Hollandia are remarkably constant in
having these hairs bifid only. Other minor details in Brug’s description are also at
variance with the material before me. it seems likely, then, that the larvae actually
described by Brug belong to some other species. Unfortunately he does not indicate
from which particular locality his specimens came.

Biology: This species bites quite freely throughout the day in scrub and around
jungle margins. The larvae are found breeding in hollow logs, treeholes and the like
in rain forest and in sago swamps; also in artificial containers such as tin cans but
particularly in old coconut shells. The many records before me indicate that such
habitats are favoured by this species and this is no doubt associated with its wide
distribution and relative commonness.

Distribution: Widely distributed throughout New Guinea and in northern Queensland.

xecords from the Moluccas and New Britain may be valid but I have had no means of checking
them. Specimens have been examined from Hollandia, 15.ix.44; 10.x.44; 18.x.44; 1.1.45; 2.1.45:

BY DAVID J. LEE. 233

9145+; 15.145; 24.31.45; 261.45; 6:11:45; 12.11.45; 22.11.45; 23.11.45; 26.11.45; 4.111.45; 20,111.45;
14.iv.45; Nakasawa, 12.11.45; Sapari, 26.11.45; Mt. Dafonsero, 4.iii.45 (460 metres); Mt.
Dafonsero, 4.iii.45 (770 metres) ; Doromena, 3.ii.45. (All the above collected by 19th Medical
General Laboratory, United States Army.) Milne Bay (Allman, .43; Cameron, .43; Ratcliffe,
1x.43; W. V. King, ix.43); Dobodura (Ratcliffe, x.43); Buna (Ratcliffe, xi.43); Tsili Tsili
(W. V. King, ix.43); Lae (Ratcliffe, xi.43; D. O. Atherton) ; Lalapipi (Ratcliffe, ix.43); Bulldog
(Atherton, 4.ix.43); Cape Endaiadere (K. G. Clinton, viii.43). Apart from the above localities
in New Guinea I have seen specimens from northern Queensland, namely, Wongabel (D. H.
Colless, 13.i.45) and Upper Barron (D. H. Colless, 17.i.45), and Miss E. N. Marks has informed
me that she has examined larvae from the-Blackall Range which were identical with those of
this species.

TRIPTEROIDES (TRIPTEROIDES) SPLENDENS, Nn. SDp-

Types: Holotype female from Dobodura (F. N. Ratcliffe, Oct., 1943), allotype male from
Buna (in scrub, July, 1942), and seven female paratypes in the Museum of the Division of
Economic Entomology, Council for Scientific and Industrial Research, Canberra, A.C.T. Two
female paratypes in Macleay Museum, University of Sydney, two female paratypes in the British
Museum and two female paratypes in the National Museum, Washington. All paratypes from
Dobodura (F. N. Ratcliffe, Oct., 1943).

Type Locality: Dobodura, New Guinea, approx. 7° 30’ S. by 148° 15’ E.

DISTINCTIVE CHARACTERS.

Particularly distinctive of this species are the fairly numerous broad scales. of the
posterior pronota, the dark and almost bare mesepimeron, the very small silvery spot on
abdominal tergite II and pronounced lateral silver triangles on the succeeding segments.

DESCRIPTION.
Female.

Head: There is an azure-blue band (appearing black in some lights) in front
extending from the eyes almost to the nape. The upright forked scales of the head are
entirely black but laterally the head is silver scaled. The palpi are black, a little shorter
than the clypeus and the proboscis is very long, slender and black. The pedicels are
yellowish-brown.

Thorax: The scutum is light brown anteriorly but almost black on the distal two-
thirds. It is clothed with fine black scales and dorsocentral bristles are present. The
pronotal lobes are clothed with broad black scales mingled with some narrow ones. The
posterior pronota are clothed with broad appressed black scales at times mingled with some
narrow ones. Except for the light brown anterior and posterior pronota the pleura are
dark; the sternopleuron is largely clothed with flat silver scales but the mesepimeron is
bare except for a very small patch of silvery scales on the upper third next to the sterno-
pleuron. The scutellum is yellowish-brown with a patch of flat black scales on each lobe
and the postnotum is dark brown.

Legs: The coxae are yellow with silvery scales laterally. The fore and hind femora
have silvery white spots at just beyond the middle and preapically. The mid femora
are similar but there is an additional spot on the basal third.

Wings (Plate xiii, b): The scales are small, dark and broad. The base of the lower
fork cell is slightly nearer the base of the wing than that of the upper and the upper
fork cell is almost twice the length of its stem.

Abdomen: This is black scaled dorsally with a large lateral silver area extending
from base to apex of segment II, a very small apical silver spot on III well removed from
the lateral margin of the tergite, large basal lateral silver triangles on IV—V, and VI-VII
with apical lateral bands expanding at their dorsal ends.

Male.

The male agrees in all respects with the female. On the genitalia the lobes of the
ninth tergite (Text-fig. 7) are long and narrow, straight for most of their length but
bluntly pointed apically. The four strong bristles arise from well below the apex. The
style (Text-fig. 7a) is modified with a slight spur at about three-quarters from the base.
Altogether the genitalia are very close to those of 7. elegans.

The larval stages have not yet been isolated.

ho
(5%)
NS

NOTES ON AUSTRALIAN MOSQUITOES. VI,

Biology: Most of the specimens collected have been taken in scrub, one in a tent and
the one from Tsili Tsili was biting at time of capture.

Distribution: So far this species has only been found at Dobodura, Buna, Lae (Ratcliffe,
Nov., 43) and Tsili Tsili (W. V. King, Sept., 43).

duu

AS ey
F am tg

Text-figs. 1-8.—Terminalia of ornamented species of Tripteroides. All figures x 190 approx.

1. Complete ninth tergite of T. argyropa. 2. Complete ninth tergite of T. alboscutellata.
2a. Style of JT. alboscutellata. 3. Lobe of ninth tergite of 7. nissanensis. 4. Lobe of ninth
tergite of T. quasiornata. 5. Lobe of ninth tergite of T. elegans. 6. Lobe of ninth tergite of
species close to 7. brevipalpis (specimen from Oro Bay, see page 241). 7. Lobe of ninth tergite
of T. splendens. Ta. Style of T. splendens. 8. Lobe of ninth tergite of T. bimaculipes.

TRIPTEROIDES (TRIPTEROIDES) LITTLECHILDI (Edwards).

Epwarps, F. W., 1930.—Bull. ent. Res., 21: 544 (Rachionotomyia).

Types: Holotype female and six other females in British Museum.

Type Locality: Tauri R., Tapala, between Yule I. and Port Romilly, Papua. ‘The Tauri R.
is also Known as the Heate, 8° S. by 146° EH.

Synonymy: Rachionotomyia littlechildi, Edwards 1930. Loc. cit.

DISTINCTIVE CHARACTERS.

T. littlechildi is close to both T. bimaculipes and T. splendens. It is rather darker
than both, the scutum except for its anterior margin being shining dark brownish-black
and the pleura are similarly dark. The completely bare, shining black mesepimeron
distinguishes it from both; the narrow black scales of the posterior pronota are similar
to those of 7. bimacuiipes but distinct from T. splendens. The abdominal adornment is
quite different from that of J. bimaculipes, there being the usual large silver lateral
area on segment II, there is apparently no silvery marking on III, IV and V have apical
lateral silver triangles and VI and VII have apical lateral bands. There are also dull
black patches laterally on the tergites rather similar to those of J. magnesiana. The
above description of the abdominal pattern is based on the specimen before me and is
much the same as that given by Brug (1934, p. 508) for J. littiechildi. It does not
conform with the original description given by Edwards (loc. cit.) but as Brug was

BY DAVID J. LEE. 235

almost certainly making his observations on the type series it may be assumed that his
notes are accurate, particularly as Edwards remarks that “the leaden-grey markings of
the abdomen might perhaps be more silvery in fresh examples’. The adornment of the
femora on the specimen before me comprises a pale silvery stripe on the basal half and
a preapical pale spot, proximally produced into a line on the fore femora; silvery spots
on the basal third, about the middle, and preapically on the mid femora, and a basal
silvery stripe to just beyond midway and a fairly large subapical spot on the hind
femora.

Neither the male nor the larva of this species is yet known.

Distribution: Known only from the type locality and Bulldog (on the Lakekamu R.) in
Papua (Atherton, viii.43). :

TRIPTEROIDES (TRIPTEROIDES) ELEGANS Brug.

Brue, 8S. L., 1934.—Bull. ent. Res., 25: 507.
Type: Described from a single female lodged in the British Museum.
Type Locality: Torpedoboot R., south coast of Dutch New Guinea.

DISTINCTIVE CHARACTERS.

The scutum is almost black and even the anterior margin is dark reddish-brown.
The pronotal lobes are yellowish-brown and clothed with narrow black scales and the
posterior pronota are dark brown and almost bare of scales (if any are present they are
narrow). ‘There are large areas of silver scales over most of the sternopleura and
mesepimera. The scutellum is brown with a patch of flat black scales on each lobe.

In the specimens before me the fore femora have a fine silvery line running from
the base to the middle and expanded there to a spot, and another spot is situated
preapically. The mid femora are similarly marked and also the hind femora except that
the preapical spot is somewhat more conspicuous.

There is a large lateral silver patch extending from base to apex of segment II of
the abdomen, a very small apical lateral patch on III (well removed from the lateral
margin), large apical lateral patches on segments IV—-VI and a smaller one on VII.
Male.

The male resembles the female in all details. The genitalia are very similar to
those of JT. splendens. The lobes of the ninth tergite (Text-fig. 5) are long and straight
but tapering apically and the four or five spines on each lobe are strong, broad, and
arise well behind the apex of the lobe. There is also a slight modification of the style,
there being a distinct spur at about two-thirds from the base.

The larva of this species has not yet been found.

Distribution: The type locality, and I have examined two female and one male specimens
from Hollandia (19th Med. Gen. Lab.). These specimens were caught in a light trap by
W. V. King.

TRIPTEROIDES (TRIPTEROIDES) ALBOSCUTELLATA, Nl. Sp.

Types: Holotype male, allotype female (both C. J. Steinhauer, 19th Med. Gen. Lab.,
15.xi.44) and two male paratypes (one as above, the other 22.i1.45), together with cast larval
and pupal skins of a paratype, lodged in the C.S.I.R. Collection. Paratype male (19th Med.
Gen. Lab., 22.145) and cast larval and pupal skin in National Museum, Washington, U.S.A.

Type Locality: Hollandia, Dutch New Guinea.

DISTINCTIVE CHARACTERS.

Particularly characteristic of this new species are the narrow scales of the pronotal
lobes and the silvery scaled scutellum. The dark scutal integument and the largely
silver scaled mesepimeron resemble T. bimaculipes but the characters above are distinc-
‘tive. The male terminalia differ from those of any other ornamented species for which
the male is known, although an undivided ninth tergite is also found in 7. argyropa and
occasional unornamented species.

DESCRIPTION.
Male.

Head: The head has a band of azure-blue scales in front for over half its depth.
The pedicels of the antennae are dull black, the clypeus and palpi black, the latter

Y

236 NOTES ON AUSTRALIAN MOSQUITOES. VI,

exceeding the clypeus by its length. The proboscis is very long, slender and black
scaled.

Thorax: The scutum is almost black except for the yellowish-brown anterior
margin, and is clothed with narrow black scales. The scutellar scales are silvery,
particularly when viewed from above; the postnotum is dark brown. The pronotal
lobes are yellowish-brown, clothed mainly with narrow black scales, but there may be
some broad scales intermingled. The posterior pronota bear few or no scales (if present
they are narrow) and a strong bristle. There are about three spiracular bristles. The
pleura are largely dark brown but yellow beneath the wing and the sternopleura and
mesepimera are largely clothed with silver scales.

Legs: The coxae are yellow with silvery scales laterally.. On the allotype the fore
femora have a golden line on the outer surface becoming silver distally, extending from
the base to about three-quarters. There is also a preapical silver patch. The mid
femora are similar and the hind femora have a preapical silvery patch which may be
indistinct.

Wings: The scaling is small but broad, the anterior fork cell is slightly shorter than
its stem and the base of the posterior one is nearer to the base of the wing.

Abdomen: This is black dorsally with a large lateral silver area on segment II,
lateral apical silver patches somewhat expanded dorsally on III and IV, lateral apical
patches on V and VI and a variable patch on VII. The venter is golden.

Genitalia: The ninth tergite (Text-fig. 2) of this species is very distinctive. It is
not divided but comprises a single plate with four prominent spines, the outer pair
being longer than the inner pair and nearly as long as the coxite. The basal lobe of
the coxite is unusual in having a curved row of about fifteen long and equal thickened
hairs and the style is distinctly swollen in the middle (Text-fig. 2a).

Female.

The female agrees with the male in all essential details.
Larva.

This differs from T. bimaculipes in having head hairs A and f three- to four-
branched and the clypeal spine evenly tapering and only moderately curved. Hair 7
of the mesothorax (Text-fig. 22) is simple but unmodified and hair 7 of the metathorax
usually has three spines as in TJ. bimaculipes but occasionally has four or five. A
striking feature of the eighth abdominal segment is the chitinous lateral plate from
which the lateral comb arises. There are some eighteen lateral comb teeth, each bluntly
pointed. The third pentad hair is usually nine-branched. On the siphon the pecten is
reduced to one to three teeth, there are ten to twelve bifid ventral tufts and the dorsal
hairs are bifid. The saddle hair has two to three branches and the ventral beard is a
single tuft of eight to twelve branches. The terminal segments are illustrated in Text-
figure 23.

Biology: The larvae have been found breeding in a log hole in a sago swamp and in
treeholes in rain forest.

Distribution: Apart from the type locality, I have examined both larval and adult specimens
from Lalapipi in Papua.

TRIPTEROIDES (TRIPTEROIDES) MAGNESIANA (Edwards).

EDWARDS, F. W., 1924.—Bull. ent. Res., 14: 361 (Rachionotomyia).
Type: Single female in British Museum.
Type Locality: Magnetic Island, north Queensland.
Synonymy: Rachionotomyia magnesiana, Edwards 1924. Loe. cit.
Rachionotomyia quasiornata (nec Taylor) of Edwards 1921. Bull. ent. Res., 12: 80.

DISTINCTIVE CHARACTERS.

The thoracic integument is shining yellowish-brown. From similarly coloured species
it is distinct in having the scales of the pronotal lobes entirely narrow.

There is a band of deep blue scales, covering more than the anterior half of the
head (in some lights, of course, this appears rather grey). The pedicels are yellowish-
brown with some fine hairs, the clypeus is dark brown and the palpi black and scarcely
longer than the clypeus. The black proboscis is long and slender, considerably longer
than the fore femur and longer than the abdomen.

BY DAVID J. LEE. 237

The integument of the scutum, the anterior and posterior pronota are yellowish-
brown, that of the pleura, including the mesepimeron, dark brown and largely clothed
with flat silver scales. The scales of the pronotal lobes are narrow and black and the
posterior pronota may also have a few fine black hair-like scales. The scutal scaling is
black, the individual scales being very fine, almost hair-like. Dorsocentral bristles are
present on the anterior half of the scutum and there is a pair of prescutellars on either
side. There is a single posterior pronotal and two or three spiracular bristles. The
scutellum is clothed with flat dark scales but those at the base of the mid lobe may
appear silvery in some lights.

There is the usual subapical silver patch on each femur and a variable amount of
silver scaling from the base to the middle.

The wings are scaled as in 7. bimaculipes, the upper fork cell is about equal to its
stem and the base of the lower cell is nearer to the base of the wing.

The abdomen is black dorsally with lateral dull black as well as silvery patches;
the silvery patch on segment II is large and of even width from base to apex of the
segment; those on segments III—-VII take the form of apical triangles, the base being
on the distal margin of each segment.

Neither the male nor the larva has yet been recorded for this species.

Biology: The breeding habitat is unknown but this species has been observed biting:
man in rain forest in the Northern Territory (A. R. Woodhill).

Distribution: Known from northern Queensland and the Northern Territory. Specimens:
have been examined from Adelaide R. (A. R. Woodhill) and Groote Eylandt (N. B. Tindale)
and Cairns (D. H. Colless, 27.iv.44).

TRIPTEROIDES (TRIPTEROIDES) QUASIORNATA (Taylor).
TAYLOR, F. H., 1915.—Proc. LINN. Soc. N.S.W., 40: 177 (Stegomyia).
EDWARDS, F. W., 1924.—Bull. ent. Res., 14: 361 (Rachionotomyia).

Type: Female type in School of Public Health and Tropical Medicine, Sydney.

Type Locality: Innisfail, north Queensland. ;

Synonymy: Stegomyia quasiornata, Taylor 1915. Loc. cit. Mimeteomyia quasiornata,
Taylor, F. H., 1916. Proc. Linn. Soc. N.S.W., 41: 566. Rachionotomyia quasiornata, Edwards
1924. Loc. cit. [Rachionotomyia quasiornata (nec Taylor) of Edwards, F. W., 1921. Bull. ent.
Res., 12: 80 is really T. magnesiana, subsequently described by Edwards 1924 (Rachionotomyia
magnesiana).]

DISTINCTIVE CHARACTERS.

These are discussed for the adult in the corresponding section under T. nissanensis.
The larva is distinguished by the number of branches in the dorsal hairs of the siphon
(usually five to six) and the rather regular lateral comb.

DESCRIPTION.
Female.

Head: There is a broad band of brilliant blue scales at the front of the head covering
less than half its depth. The pedicels are yellowish, the clypeus dark brown and the
palpi black and equal in length to the clypeus. The proboscis is black, slender, very
long, decidedly longer than the fore femur.

Thorax: The scutal integument is yellowish anteriorly and laterally and indefinitely
brownish medially and posteriorly; the scaling is sparse and the individual scales dark
and narrow curved with a very few broad, flat, black scales above the wing roots. The
seutellar scales are black. The pronotal lobes are clothed with flat black scales but the
posterior pronota have some narrow curved black scales and a few small broad, rather
triangular black ones. In the type, only one such scale is present on one side but on
two specimens (in Macleay Museum, University of Sydney) collected by Taylor from the
type locality, and presumably part of the original series, from two to three such scales
are present. There is a strong posterior pronotal bristle, two brown spiracular bristles
but no upper sternopleural. The central area of the pleura is brown, but the distal
half of the sternopleuron and the meron are rather darker. The darker portion of the
sternopleuron and the upper part of the mesepimeron are clothed with flat silvery scales.
The postnotum is also brown.

VI,

NOTES ON AUSTRALIAN MOSQUITOES.

238

<Q Pr

Mi

(IN |
‘< Xi

BY DAVID J. LEE. 239

Legs: All the femora have distinct pale or silvery patches just beyond midway and
preapically. There is a narrow pale-scaled line on the outer surface of the fore femora
from the base to about midway and there may be some indefinite pale scaling at the
base of the mid femora as well.

Wings: The wings are scaled as in T. bimaculipes, the upper fork cell is about equal
to its stem in length and the base of the lower cell is nearer to the base of the wing than
that of the upper cell.

Abdomen: There is a lateral silvery scaled area covering the depth of tergite II, the
succeeding segments have apical lateral silvery bands (not continuous across the dorsum)
and the venter is yellowish.

Male.

I have seen no males from the type locality but a specimen before me from Mt.
Glorious, near Brisbane, 13.11.45, agrees well with the Innisfail specimens but has six
to seven broad scales on the posterior pronota. The genitalia are similar to those of
T. bimaculipes and the ninth tergite is figured in Text-figure 4.

Larva.

The head and thorax are closely similar to JT. bimaculipes, the clypeal spines, the
dorsal head hairs and the modified hairs of the thorax being identical. The lateral comb
teeth (Text-fig. 16) are similar but rather more regular in size. There are ten to twelve
branches in the third pentad hair and four in the fifth. The pecten spines (Text-fig. 17)
are longer than in 7. bimaculipes and the dorsal hairs of the siphon are two- to seven-
branched with the majority with five to six branches. Both the saddle hair and the
ventral beard are more strongly branched than in TJ. bimaculipes, the former having
seven branches and the latter about twelve. The terminal segments are figured in
Text-figure 15. It will be noted that this description does not entirely tally with the figure
of the terminal segments of JT. quasiornata larva published by Hill (1925, Plate vi,
fig. 10) and reproduced by Lee (1944, p. 25), but see, however, remarks following
description of 7. nissanensis (page 240).

Biology: The larva has been found breeding in treeholes by J. L. Wassell on Mt.
Glorious.

Distribution: Specimens have been examined from Innisfail (F. H. Taylor) and Mt.
Glorious in Queensland.

TRIPTEROIDES (TRIPTEROIDES) NISSANENSIS, nN. Sp.

Types: Holotype male, allotype female and morphotype larvae in the C.S.1I.R. Collection.
Paratype female and morphotype larvae in the Cawthron Institute, Nelson, New Zealand.
Type Locality: Nissan in the Green Islands just east of New Ireland, 154° E. by 4° 45’ S.

DISTINCTIVE CHARACTERS.

The most obviously differential characters are to be found in the form of the ninth
tergite of the male and in the larva. However, the female is only likely to be confused
with T. brevipalpis, with the specimens noted under that species as being probably
distinct, and with 7. quasiornata. From T. brevipalpis it is distinct in having the base
of the posterior fork cell decidedly nearer the base of the wing, from the specimens from
Oro Bay and Biak by the darker central part of the pleura (particularly at the border of
the meron) and brown metanotum, but from 7. quasiornata there does not appear to be
any satisfactory way of separating the females. The larva is very similar to that of
T. bimaculipes but may be distinguished by the simple or non-divergent terminations
of the stellate hairs.

Text-figs. 9-23.—Larval structures of ornamented species of Tripteroides. 9 and 12 x 45
approx.; 10, 11, 15, 18, 19, 22 and 23 x 40 approx.; 13, 14, 16, 17, 20 and 21 x 100 approx.

9-14. T. bimaculipes. 9. Dorsal view of head. 10. Thoracic chaetotaxy (dorsal). 11.
Mesothoracic hairs 2-7. 12. Terminal segments. 13. Lateral comb teeth. 14. Pecten spines.
15-17. T. quasiornata. 15. Terminal segments. 16. Lateral comb teeth. 17. Pecten spines.
18-21. T. nissanensis. 18. Mesothoracic hairs 2-7. 19. Terminal segments. 20. Lateral comb
teeth. 21. Pecten spines. 22-23. T. alboscutellata. 22. Mesothoracic hairs 2-7. 23. Terminal
segments.

240 NOTES ON AUSTRALIAN MOSQUITOES. VI,

DESCRIPTION.
Male.

Head: There is a band of azure-blue in front covering half the depth of the head.
The pedicels of the antennae are brown, the palpi black and not as long as the clypeus.
The proboscis is black, very long and slender.

Thorax: The scutum is shining yellowish-brown and clothed with narrow black
scales. The pronotal lobes are yellowish with broad flat black scales and the posterior
pronota similarly coloured but bearing narrow curved black scales with a very few
broad ones intermingled, and a strong bristle. There are two to three black spiracular
bristles. The sternopleura and mesepimera are largely clothed with pale scales with
silvery reflections. The pleura are yellowish-brown but the central area is slightly
darker. The scutellum has patches of flat black scales on each lobe and the postnotum
is dark brown.

Legs: The coxae are yellow with silvery scales laterally. The rest of the legs,
except for the femoral markings, are black. All the femora have a white patch just
beyond the mid point and another one preapically.

Wings: The scales are small and broad. The upper fork cell is not quite as long
as its stem and the base of the lower fork cell is distinctly nearer the base of the wing
than that of the upper cell.

Abdomen: This is black dorsally with a large apical lateral silver triangle on
segment II and apical lateral silver bands on III to VII. The venter is golden.

Genitalia: The lobes of the ninth tergite (Text-fig. 3) are shallow, broad and not
deeply indented. Each lobe carries at least twenty setae rather irregularly disposed on
the distal margin. The coxites are short being no longer than their width at the base.
Female.

This sex is similar in all essentials to the male.

Larva.

The head and thorax closely resemble those of 7. bimaculipes. The clypeal spines
are stout and very strongly elbowed at the base, the dorsal head hairs are all simple and
the modified hairs of the thorax are similar to those of 7. bimaculipes. The mesothoracic
hairs are shown in Text-figure 18. On the eighth segment the lateral comb is more
strongly developed than in 7. bimaculipes, the combs on each side meeting or almost
meeting ventrally. (See Text-fig. 20 for details of lateral comb teeth.) The third
pentad hair usually has six branches and the fifth four. The siphon has about seven
pecten spines (Text-fig. 21), ten to twelve ventral tufts, of which the two basal ones
are trifid, the rest bifid, and the dorsal hairs are all bifid. The saddle hair is three- to
four-branched and the ventral tuft comprises some seven branches of varied lengths.
Text-figure 19 illustrates the terminal segments. The stellate tufts of the thorax and
abdomen have the individual branches terminating in a single or double point, but if
the latter then the points are not divergent. In the specimens before me the general
colouration of the integument is rather darker than is usual in the genus.

Biology: The larvae were found in treeholes by L. J. Dumbleton.

Distribution: The type series was collected on Nissan by L. J. Dumbleton. Apart from
these specimens, a male and a female in the Macleay Museum, University of Sydney, from
Rabaul, New Britain, are rather similar to JT. nissanensis but the genitalia preparation of the
male is rather too distorted for a determination to be made with confidence. These specimens
also differ in having the blue band on the head more than half its depth (this is particularly
so in the male) and another specimen from Toma, New Britain, agrees with those from Rabaul.
These specimens, together with the figure published by Hill (1925—called T. quasiornata) of a
larva from New Britain suggest that a different form exists on this island.

TRIPTEROIDES (TRIPTEROIDES) BREVIPALPIS Brug.

Brue, 8S. L., 1934.—Bull. ent. Res., 25: 508.

Types: Described from four males, five females and two larval skins. Type male and
female in the British Museum.

Type Locality: North coast of Ceram (one of the following places—Warasiwa, Oewin or
Hoelong).

BY DAVID J. LEE. 241

DISTINCTIVE CHARACTERS.

The combination of yellow thoracic integument, with broad scaled pronotal lobes
and narrow scaled posterior pronota, the position of the fork cell bases (described as
level) together with the abdominal and femoral adornment should serve to distinguish
this species from its allies. There are apical silver bands on the abdominal tergites
which are only briefly interrupted medially. The femora of the fore legs are described
as having a narrow golden line on the apical half and two silvery patehes on the
basal half, those of the mid leg with two silvery patches apically and a silvery line
basally, and those of the hind leg with a preapical silvery patch and a broad silvery
stripe about the middle.

According to Brug (loc. cit.) all the dorsal head hairs of the larva are simple and
the clypeal spines are stout and bent. Hair 7 of both the mesothorax and metathorax
is modified as in JT. bimaculipes and other species. There are only 14 teeth in the
lateral comb of the eighth segment and seven pecten teeth are present on the siphon.
The ventral hair tufts are bifid and the dorsal ones simple or bifid. The saddle hair
is long and bifid and the ventral tuft five-branched. There is little in the description
to differentiate the larva of this species from that of JT. bimaculipes (as described
above). Perhaps the reduced number of lateral comb teeth may be significant.

Biology: The larvae were found breeding in bamboo stumps and cut bamboos.

Distribution: Only known from Ceram.

Note.—Two specimens from Oro Bay (W. V. King) agree in colouration, scaling of
the pronotal lobes and posterior pronota, and abdominal adornment with 7. brevipalpis
but differ in other details. The pleural integument is yellowish (instead of dark brown
as in T. brevipalpis) and the base of the lower fork cell is distinctly nearer the base
of the wing than that of the upper cell. Several other specimens from Biak (University
of Queensland Collection) agree with the Oro Bay material in all details and I feel
these must comprise a quite distinct species. However, since no differences can be
noted in the genitalia of the New Guinea specimens (both Biak and Oro Bay) and the
brief details and figure of the ninth tergite given by Brug, it would be wise to wait for
more adequate material before defining this probably new form. The ninth tergite of
a specimen from Oro Bay is illustrated in Text-figure 6.

TRIPTEROIDES (TRIPTEROIDES) DISTIGMA Hidwards.

EDWARDS, F. W., 1925.—Bull. ent. Res., 15: 257 (Rachionotomyia).
Type: Described from a single female in British Museum.
Type Locality: Tulagi on Florida I. in the Solomon Islands.
Synonymy: Rachionotomyia distigma, Edwards 1925. Loc. cit.

DISTINCTIVE CHARACTERS.

According to the description 7. distigma is allied to T. magnesiana and others with
pale scutal integument. The thoracic integument is mainly orange but the scutal
scaling consists of straight hair-like greenish scales with roundish dark brown spots
above each wing root clothed with flat deep black scales. The pronotal lobes bear flat
black scales and the posterior pronota a few narrow dark ones.

The mid femora have one small obscurely pale spot at the middle in front and the
hind femora have a dark dorsal line reaching the base, and on the outer side are pale
golden with the tip and a longish area beyond the middle, dark. There are no silvery
markings on the tergites and the venter is golden.

Note-—Edwards (1925) considered this species to answer the description of
T. ornata in many respects, but see synonymy of 7. bimaculipes (page 231).

Neither the male nor the larva has yet been recorded.
Biology: Nothing is known of the larval or adult habits of this species.
Distribution: So far only known from the type locality.

TRIPTEROIDES (TRIPTEROIDES) PURPURATA (Hdwards).

EDWARDS, F. W., 1921.—Bull. ent. Res., 12: 79 (Rachionotomyia).
, 1924.—Ibid., 14: 361 (Rachionotomyia).

242 NOTES ON AUSTRALIAN MOSQUITOES. VI,

Buxton, P. A., and HopkKINs, G. H., 1927.—Res. in Polynesia: 78 (Rachionotomyia). (Includes
description of larva.)
Types: Described from three females. In British Museum.
Type Locality: Suva, Fiji.
Synonymy: Rachionotomyia purpurata, Edwards 1921. Loc. cit.

DISTINCTIVE CHARACTERS.

From the description it appears that this species is closest to those with a light
coloured scutum. The scutal integument is dull orange, clothed with greenish scales
mingled with some black ones. There are broad flat black scales on both the pronotal
lobes and the posterior pronota.

The larva apparently closely resembles 7. bimaculipes but may be distinguished
from this and other species by the chitinous plates from which the lateral combs of
the eighth abdominal segment arise being united dorsally; the dorsal portion of the
plate is more heavily chitinized than that from which the comb arises. The clypeal
spines are not bent.

The male has not yet been described.
Biology: This species has been recorded breeding in an old kerosene tin and in tree-
holes.

Distribution: As far as I am aware, this species is only known from the type locality and
Moturiki, a small island nearby. Its larval habitat would suggest a reasonably wide distribution.

UNORNAMENTED SPECIES.

All those species lacking metallic silvery scaling on the thorax and abdomen and
generally of a drab appearance belong, within the Australasian Region, to either the
subgenus Rachisoura or Mimeteomyia. (There are two possible exceptions, T. subnudi-
pennis and T. concinna which are discussed elsewhere.) All are best subdivided on the
wing scaling which appears to be the most satisfactory character on which to base
groupings of species.

Subgenus RacHisourRA Theobald.
EDWARDs,. F. W., 1932.—Culicidae in Wystman’s Genera Insectorum, Fasc. 194: 76.

This subgenus is, with the exception of the monotypic Maorigoeldia, the most
restricted of all in its distribution, being confined to New Guinea, islands eastwards to
the Solomons and the north of Australia. It is also largely nepenthicolous in habitat
but some members of both Tripteroides and Mimeteomyia have also been found in
Nepenthes.

The adults all have veins R, to R, without any narrow outstanding scales although
the scales of these veins are broad and fairly large and dense. The larvae
characteristically possess enlarged, modified maxillae.

Two groups are recognized below, namely, the filipes-group in which there are no
narrow outstanding scales on the wing veins and the vanleewweni-group in which
narrow outstanding scales are present from vein R,,; to Cu,. No differentiation of the
larvae of the two groups is as yet obvious.

FILIPES-GROUP.

Apart from the wing scales being all broad there is always a posterior pronotal
bristle. The male palpi are variable in length, but no dorsocentral bristles are present.
The members of the group are T. filipes, T. papua, T. fuliginosa, T. latisquama, T. longi-
palpata and T. confusa. The male palpi of the first three are short and those of the last
two almost as long as the proboscis.

The male of 7. latisquama is not known.

TRIPTEROIDES (RACHISOURA) FILIPES (Walker).

WALKER, F., 1861.—Proc. Linn. Soc. Lond., 5: 229 (Culex).

Epwarps, F. W., 1924.—Bull. ent. Res., 14: 362 (Rachionotomyia).

——, 1927.—Nova Guinea (Zoologie), 15: 354 (Rachionotomyia).

Nec PAINE, R. W., and Epwarps, F. W., 1929.—Bull. ent. Res., 20: 305 (larval description, also
reproduced in part in Lee, 1944).

BY DAVID J. LEE. 243

Types: Described from a female. In British Museum? As the male in the original series
of T. sylvestris (which otherwise might be used as allotype for 7. filipes) is not conspecific with
the female (see below), I hereby designate an allohypotype male and four parahypotype males
which are lodged in the C.S.I.R. Museum. Four further parahypotype males are also designated
which are lodged two each in the British Museum and the National Museum, Washington. The
east larval and pupal skins of the allohypotype are also preserved in the C.S.I.R. Slide Collection
and these become morphohypotypes.

Type Locality: Dorey, New Guinea (Dore or Doreh, in Dutch New Guinea, immediately
west of Manokwari—0° 51’ S. by 134° 15’ E.). The type locality of the allohypotype and para-
hypotypes is Hollandia.

Synonymy: Culex filipes Walker 1861. Loc. cit. Rachionotomyia filipes Edwards 1924.
Loc. cit. Rachisoura sylvestris Theobald, F. V., 1910. Monogr. Cul., 5: 208. Stegomyia hilli
Taylor, F. H., 1914. Proc. Linn. Soc. N.S.W., 39: 456. Mimeteomyia hilli Taylor, F. H., 1916.
Ibid., 41: 566.

Note.—T. hilli has been included in the synonymy of T. filipes following Edwards (1924).
I have not been able to examine this type but I am informed that it is badly damaged. Some
doubt as to the correctness of this synonymy does seem justified as 7. filipes is a nepenthicolous
species, and as far as I have been able to discover Nepenthes are absent from the Northern
‘Territory.

I have taken as the basis of the present conception of this species a series of
specimens found at Hollandia, a locality rather similarly situated to, and less than 800
kilometres east of, the type locality.

DISTINCTIVE CHARACTERS.

This is a typical Rachisoura with all the wing scales broad, and short palpi in both
sexes. Although best characterized by the form of the ninth tergite of the male this
species belongs to a group which is characterized by the light pleural integument. It
is readily distinct from JT. latisquama by the reduced number of prescutellar bristles
(no more than two pairs as compared with about seven pairs) and from other species,
particularly J. confusa, the relatively short hind tibiae (only 75-78% of the mid
tibiae) serve to differentiate it. The larva appears adequately characterized by the
restricted development of the lateral comb and the enlarged apical tooth of the maxilla,
which is distinctly longer than the body of the maxilla.

DESCRIPTION.
Female.

Head: The head is clothed with broad greyish-black appressed scales. There are
some pale upright scales with fine pale narrow curved scales at the centre of the nape,
but laterally the upright scales are black. There is an area of flat pale scales low
down at each side of the head but no line of pale scales across the eye margins. The
proboscis is a little shorter than the abdcmen, equal to the fore femur, rather stout
and slightly recurved towards the tip. The palpi are black, 0-2 the length of the
proboscis and extending beyond the clypeus for three to four times its length. The
clypeus is round and dark brown; the pedicels are dark brown or black, pubescent,
with fine greyish hairs. The antennae are black and almost as long as the proboscis.

Thorax: The scutum is dark brown, the pleura yellowish-brown. The scales of the
scutum are dense, broad, somewhat curved and greyish-black. There are no dorso-
central bristles but one pair of prescutellars is present. The pleura are largely clothed
with pale yellowish scales, the pronotal lobes with pale scales but there are some
brownish ones above and behind; on the posterior pronota the scales are pale below
with light brownish ones on the upper part. One posterior pronotal bristle is usually
present; there are four or five black spiraculars and one upper sternopleural bristle;
the latter is not particularly strong and may be absent. The postnotum is dark brown
with a few scarcely noticeable hairs on the distal portion.

Legs: The coxae are yellow with pale scales laterally. The rest of the legs are
dark scaled except that the femora are pale beneath. The hind tibiae are only from
75% to 78% the length of the mid tibiae.

Wings (Plate xii, b): All the scales are broad and brown. The upper fork cell
is twice the length of its stem and the base of the lower fork cell is usually slightly
nearer the base of the wing than that of the upper cell, but they may be level or their
relative positions may be reversed.

244 NOTES ON AUSTRALIAN MOSQUITOES. VI,

Abdomen: This is black dorsally with strong apical lateral pale triangles (their
bases on the distal margins of the segments) on segments II to VII; the line of
demarcation between the colours is strongly serrate. The venter is pale and the apex
of the abdomen very hairy.

Male.

The palpi are one-third the length of the proboscis (not two-thirds as stated by
Edwards, 1924, p. 362). The upper fork cell is slightly shorter than in the female
(about 1:75 its stem). Two pairs of prescutellar bristles are occasionally present and
the abdomen is pale dorsally at the apex of segment VIII.

The lobes of the ninth tergite are long, broad and deeply emarginate and with
eight to ten strong spines on the distal margin. (See Text-fig. 24.)

Larva.

The clypeal spines are rather long and fairly slender. Head hairs A, B and C
are usually single, sometimes bifid, C is behind the level of A and longer than the rest;
B is well in front of C but in about the same longitudinal plane, d is a little in front
of C; e and f are single. The maxillae (Text-fig. 45) have the long apical tooth longer
than the body of the maxilla.

Most of the stellate tufts of the thorax and abdomen have two to four branches
with strong but sparse pectination and the termination of each spine is nearly always
a blunt point but occasionally there are several non-divergent points. There are no
modified thoracic spines.

The lateral comb comprises three to five teeth each with a long single or double
tooth and basal pectination (Text-fig. 51). The first pentad hair has a chitinous basal
plate and about seven strong branches (occasionally reduced to three), the second is
single, the third has three or four very small branches, the fourth is single and the
fifth has about three very small branches. The siphon is short and tapers rather
sharply apically. The ventral hairs of which there are twelve to fourteen have four
branches except the basal pair which are about seven-branched. The pecten comprises
two to four spines on the distal half of the siphon. Of these, one is situated at the
apex and is stronger, stout and slightly curved. The apicodorsal spine is rather long
but slender and the ventral siphonal valves have a very long four- or five-branched
hair. The dorsal siphonal hairs have three to five branches. The saddle covers the
dorsal half of the anal segment and there are only a few teeth (about six) on the
distal margin. The saddle hair is single and plumose, the dorsal subcaudal has six
finely plumose branches and the ventral tuft is long with six plumose branches.

Note.—Brug’s (1934) illustration of the ninth tergite of JT. sylvestris, presumably
made from a British Museum specimen (four females and one male comprised the type
series described by Theobald; doubtless the male of this series was used), does not
agree with the form of the ninth tergite found in the specimens examined by me,
which have been correlated with the females by larval characters (through cast skins),
by habitat association and by locality association in widely separated areas. It seems
most likely that the male figured by Brug actually represents some other species.
Furthermore, in Theobald’s original description, the length of the male palpi is given
as “not quite two-thirds” whereas the male described by me has the palpi only one-third
the length of the proboscis. This adds credence to the view that the male specimen in
Theobald’s type series is not conspecific with the female type and for this reason I
have designated a male allohypotype and a number of male parahypotypes. Unfortunately
I have not seen any species from Queensland with broad scaled wings in which the male
palpi are more than half as long as the proboscis, so cannot suggest what Theobald’s male
specimen might be, unless of course, it is not a Rachisoura at all.

Further, Paine and Edwards’ description of the larva of a species from the Solomon
Islands (called by them R. filipes), although close to T. jfilipes, differs firstly in the
larval habitat and in the following larval characters. The maxilla is smaller, the
largest tooth shorter and the average number of lateral comb teeth greater (six to nine).

BY DAVID J. LEE. 245

= YWly . a

GN

31
f ll -
INS,

32

33

35

39

-

41 40
42
Lee

Text-figs. 24-42—Lobe of ninth tergite of unornamented species of Tripteroides. All
figures x 190 approx.

24. T. filipes. 25. Specimen mentioned on page 246 as probably T. filipes. 26. T. filipes sensu
Paine and Edwards. 27. T. confusa. 28. T. fuscipleura. 29. T. fuliginosa. 30. T. pallida.
31. T. longipalpata. 32. Species near T. longipalpata, see page 250. 33. T. bisquamata. 34.
T. kingi (both lobes shown). 35. T. brevirhynchus (specimen from Jacky Jacky). 36. JT. sub-
obscura. 37. T. tasmaniensis. 38. T. collessi. 39. T. argenteiventris. 40. T. caledonica. 41.
T. atripes. 42. T. punctolateralis.

246 NOTES ON AUSTRALIAN MOSQUITOES. VI,

Biology: Adults have been taken biting in the vicinity of Hollandia and all larval
collections both from New Guinea and Cape York (north Queensland) have been from
pitcher plants, including some from climbing species. The species has been taken up
to an altitude of 460 metres but it has been most commonly associated with grassy
swamps.

Distribution: Specimens have been examined from Hollandia, 23.11.45; 4.ii1.45; 3.iv.45;
13.iv.45 (all 19th Med. Gen. Lab.); Hollandia, 220th M.S.U. (Malaria Survey Unit of the U.S.
Army); Dobodura, xii.43 (W. V. King) and Jacky Jacky (Cape York, north Queensland)
(Chippendale).

Note.—One specimen from Hollandia (23.11.45, 19th Med. Gen. Lab.—also labelled
718-1 with associated larval and pupal skins) in the C.S.I.R. Collection, closely resembles
T. filipes but the palpi are 0-4 the length of the proboscis and the hind tibia 94% the
length of the mid tibia. The genitalia are similar to those of T. filipes but the ninth
tergite has nine spines on one lobe (see Text-fig. 25) and twelve on the other. These
are somewhat irregularly disposed, particularly laterally. The larva has the stellate
tufts with only two to three branches and these are frayed rather than strongly pectinate
and the hairs of the ventral siphonal valves have only three branches. It is difficult to
say whether this is only an extreme variant of 7. filipes or a distinct form, but on present
evidence I am inclined to regard it as only a variant.

TRIPTEROIDES (RACHISOURA) FILIPES sensu Paine and Hdwards 1929.
PAINE, R. W., and Epwarps, F. W., 1929.—Bull. ent. Res., 20: 305 (Rachionotomyia).

As pointed out under T. filipes the larval description given by these authors does
not agree with T. filipes from New Guinea. Their specimens from Guadalcanal must
belong to an undescribed species. I have before me larvae from Guadalcanal completely
agreeing with their description (the maxilla is figured in Text-figure 44) and a male
specimen in poor condition from Tenaru (also in the Solomons) has the ninth tergite
(see Text-fig. 26) quite distinct from that of T. filipes but the palpi are short as in
T. filipes. Very likely this is the male of the species found by Paine on Guadalcanal
breeding in a large wild Aroid and in leaves on the ground. The larvae examined by
me were collected by L. J. Dumbleton breeding in Pandanus and coconut centres.

TRIPTEROIDES (? RACHISOURA) PAPUA Brug.

Brue, S. L., 1934.—Bull. ent. Res., 25: 505.
Types: Type male and female in British Museum.
Type Locality: Tanah Merah, Dutch New Guinea.

DISTINCTIVE CHARACTERS.

This species is obviously very clesely allied to 7. jfilipes in nearly all respects.
Indeed the resemblance is so close that one might be tempted to place it, tentatively
at least, in the synonymy of T. filipes. However the following points in the description
of 7. papua make it wise to withhold judgement until further evidence is available.
The lengths of the proboscis and palpi, though very close, are not quite as I have
found them in 7. filipes. The scutal scaling may be narrower in TJ. papua and there
are fewer spiracular bristles (two to three instead of four to five as in TJ. filipes).
Apart from these quite minor differences Brug suggests that the paraprocts may be
absent, or if not, are unusual, in the genitalia of this species. I have noticed that the
paraproct, in Tripteroides generally, has a tendency to fold backwards towards or
between the lobes of the ninth tergite. Very likely this is all that has happened to
Brug’s specimen but while the doubt remains it is better to retain specific status for
Brug’s name.

The larva of this species has not been described.

Biology: A nepenthicolous species (T. filipes is also nepenthicolous).

Distribution: Only known from the type locality.

TRIPTEROIDES (RACHISOURA) FUSCIPLEURA, Nn. Sp.

Types: Holotype male, allotype female, two male paratypes and a female paratype in the
C.S.I.R. Collection. One paratype of each sex in the National Museum, Washington, and one

BY DAVID J. LEE. 247

male paratype in the British Museum. All except the last one with associated larval and
pupal skins.

Type Locality: Doromena, near Hollandia, Dutch New Guinea. The female paratype in the
C.S.I.R. Collection and the paratypes in the National Museum, Washington, and the British
Museum are from Nakasawa, another village in the immediate vicinity of Hollandia.

DISTINCTIVE CHARACTERS.

Readily separable from 7. filipes, T. confusa and T. latisquama by the dark pleural
integument. The very short palpi distinguish it from T. fuliginosa (in addition to
genitalia characters). The siphon of the larva provides characters which are quite
distinct from those of T. filipes and its allies. The siphon is longer with the pecten
regularly disposed over the basal two-thirds or three-quarters, but it does not extend
to the apex as in 7’. longipalpata, nor is the ventral portion of the anal segment hirsute
as in the latter species. The enlarged and modified maxilla will of course immediately
place it as a member of the subgenus Rachisoura.

DESCRIPTION.
Male.

Head: This is black scaled with a very few pale flat and upright scales at the
middle of the nape at least, or even extending (flat scales) forwards towards the eyes.
Laterally the head is pale and there is a very narrow silvery border to the eyes. The
pedicels are black and the clypeus dark brown. The proboscis is black, fairly stout,
about equal in length to the fore femur and a little shorter than the abdomen. The
palpi are black, one-eighth the length of the proboscis (or up to one-sixth in other
specimens) and extending beyond the clypeus by somewhat more than twice its length.

Thorax: The integument (including the pleura) is dark brown with a rather
blackish tint. The scutum is clothed with dense, broad, slightly curved black scales.
There are no dorsocentral bristles and at most only one pair of prescutellars. A few
pale scales are found at the centre of the anterior promontory, the scutellum is clothed
with broad flat dark scales and the postnotum is dark brown and bare. The pleura
are densely clothed with white scales. The pronotal lobes are pale scaled below but
black scaled above and the posterior pronota are similarly covered, the upper half
being clothed with almost black scales. There is one posterior pronotal bristle and two
to three spiraculars, but no upper sternopleural.

Legs: The coxae are not as yellow as in 7. filipes, the fore coxae are largely clothed
with dark scales (some may be pale dorsally or in some specimens the majority may
appear silvery) and the mid and hind coxae are pale scaled. The femora are pale
beneath but the rest of the legs are dark. The hind tibiae are about 88% the length
of the mid.

Wings: All the scales are broad and dark. The upper fork cell is slightly greater
than twice its stem and the base of the upper fork cell is slightly nearer the base of
the wing than that of the lower cell.

Abdomen: This is dark scaled dorsally but the tergites are pale laterally with a
straight margin between the colours.

Genitalia: The lobes of the ninth tergite (Text-fig. 28) are short but fairly well
separated and carry four or five spines which are considerably longer than the lobe.
On the holotype four such spines are present on one side and five on the other. One
of the paratypes has five such spines on each side.

Female.

The female agrees closely with the male. The palpi are about one-sixth the length
of the proboscis and the latter is equal in length to the fore femur.
Larva.

The antennal shaft hair is single and situated at about three-quarters from the
base. The clypeal spines are rather long and slender, and the head hairs are all
simple and rather similar to T. filipes, but C is not as long and B is further behind the
level of d. The maxillae (Text-fig. 46) have the apical tooth shorter than the body

248 NOTES ON AUSTRALIAN MOSQUITOES. VI,

of the maxilla with three other shorter teeth of decreasing size and a number of
rudimentary teeth at the base. The stellate tufts of the thorax and abdomen have
three or four branches, each bluntly pointed, smooth, short and pale.

The lateral comb comprises four to six spines, each one having a median long
sharply pointed spine with shorter teeth near its base. The first pentad hair has
three or four very short pale branches, the second two branches, the third three, the
fourth is bifid and the fifth single. All of these are comparatively inconspicuous. The
siphon is pale, rather long, narrowing uniformly to the apex, and the pecten comprises
five to nine pointed but fringed spines distributed in a row over the basal portion of
the siphon. There are fourteen ventral hair tufts each with two to four branches
except the basal pair which have six or seven. The dorsal hairs are four- to five-
branched and the ventral siphonal valves each have a long bifid hair. The apicodorsal
spine is fairly long and slightly hooked apically. The saddle hair is short, single and
practically smooth and there are some four to seven small spines on the distal margin.
The ventral beard has five to seven plumose branches, the dorsal subcaudal six and
the ventral subcaudal one. The anal papillae are not adequately preserved on any of
the cast skins before me.

Biology: Apart from one specimen bred from a small stump hole all the rest of
those before me were bred from the leaf axils of taro.

Distribution: Doromena, 25.ii1.45; 7.ii1.45 (H. Hoogstraal) ; Nakasawa, 12.i1.45 (W. V. King
and D. Johnson).

TRIPTEROIDES (RACHISOURA) FULIGINOSA, N. Sp.

Type: Holotype male (19th Med. Gen. Lab.) in the C.S.I.R. Collection.
Type Locality: Hollandia, Dutch New Guinea.

DISTINCTIVE CHARACTERS.

The dark pleural integument distinguishes this species from 7. filipes and its close
allies. The palpi are longer than those of 7. fuscipleura, the head is entirely dark scaled
at the nape, the prescutellar bristles are better developed and the ninth tergite is
very distinct.

DESCRIPTION.
Male.

Head: This is entirely dark scaled except for a very narrow silvery rim to the
eyes. The pedicels are dark brown, the clypeus is small, brown, the palpi black and
exceeding the clypeus by four times its length and scarcely one-quarter the length of
the proboscis. The proboscis is black and slightly longer than the fore femur.

Thorax: The integument (including the pleura) is dark brown. The scutal scaling
is very dense, broad, curved and black. There are no dorsocentral bristles but three
pairs of prescutellars are present. The scutellum is clothed with broad black scales
and the postnotum is dark brown. The pronotal lobes are pale scaled below with some
black ones above and the posterior pronota are pale scaled on the basal third and black
above. There is one posterior pronotal bristle, four black spiraculars and one pale
upper sternopleural bristle.

Legs: The coxae are pale with pale scales, the fore coxae with a few dark scales
anteriorly as well. The rest of the legs are black except that the femora are pale
beneath basally.

Wings: The scales are all broad. The upper fork cell is three times the length of
its stem, and the base of the upper cell is slightly nearer the base of the wing than
that of the lower.

Abdomen: The tergites are black dorsally but pale laterally with a straight line
junction. The venter is pale.

Genitalia (Text-fig. 29): The lobes of the ninth tergite are well separated, of
medium length and with eight to nine bristles distally.

Neither the female nor the larva of this species is known.

BY DAVID J. LEE. 249

Biology: The specimen was taken in flight at the buttress of a large rain-forest
tree by H. Hoogstraal.
Distribution: Only known from the type locality.

TRIPTEROIDES (RACHISOURA) LATISQUAMA (Edwards).

Epwarps, F. W., 1927.—Nova Guinea (Zoologie), 15: 354 (Rachionotomyia).
Type : Holotype female in British Museum.
Type Locality: Nassau Mts., Dutch New Guinea (1,500 metres).
Synonymy: Rachionotomyia latisquama, Edwards 1927. Loe. cit.

DISTINCTIVE CHARACTERS.

T. latisquama is very similar to 7. filipes and 7. confusa but is readily distinguish-
able by the greater development of prescutellar bristles (six or seven pairs as contrasted
with only one pair in the other two species).

DESCRIPTION.
(Based on a female specimen from Mt. Dafonsero near Hollandia.)

Female.

Head: This is black scaled with a pale patch at the centre of the nape and a pale
rim to the eyes. The pedicels and clypeus are dark brown. The palpi and proboscis are
black, the former exceeding the clypeus by about five times its length, and almost
one-quarter the length of the proboscis which itself is longer than the fore femur.

Thorax: The scutum is dark brown, clothed with broad curved dark brown scales
laterally and somewhat narrower ones medially. The pleura are yellowish-brown (as
in T. filipes). The pronotal lobes are pale scaled below, dark scaled above and the
posterior pronota are pale below grading into darker above. There is one posterior
pronotal bristle, five to six spiraculars and two to three pale upper sternopleurals.
There are no dorsocentrals but six or seven pairs of prescutellars are present. The
scutellum is clothed with broad flat black scales and the postnotum is dark brown
and bare.

Legs: The coxae are pale yellowish-brown with pale scaling. The femora are pale
beneath basally only and the hind tibia is 88% the length of the mid.

Wings: All wing scales are broad. The upper fork cell is 1:5 times the length of
its stem and the base of the lower is a little nearer the base of the wing than that of
the upper cell.

Abdomen: The tergites are black dorsally with very prominent lateral triangles on
the apical half of segments II to VII. The venter is pale scaled.

Neither the male nor the larva of this species is yet known.
Biology: The specimen before me was collected biting at an altitude of about
1,400 metres.

Distribution: Apart from the type locality this species has been collected on Mt. Dafonsero,
near Hollandia, in Dutch New Guinea (3.iii.45, 19th Med. Gen. Lab.).

TRIPTEROIDES (RACHISOURA) LONGIPALPATA, Nl. Sp.

Types: Holotype male (19th Med. Gen. Lab., 18.iii.45) together with its cast larval and

pupal skin and a morphotype larva in the C.S.I.R. Collection. All were collected by J. P.
Toffaleti.

Type Locality: Hollandia, Dutch New Guinea.

DISTINCTIVE CHARACTERS.

The long male palpi relate this species to 7. confusa but the form of the ninth
tergite and the maxilla of the larva are striking characters of distinction.

DESCRIPTION.
Male.

Head: The head is black with a very few pale scales at the middle of the nape
and a very narrow silvery rim to the eyes can be seen. The proboscis is equal to the

fore femur and the palpi are almost equal in length to the proboscis. The pedicels and
the clypeus are brown.

250 NOTES ON AUSTRALIAN MOSQUITOES. VI,

Thorax: The scutum is dark brown and clothed with broad curved black scales.
There are no dorsocentrals and no prescutellars can be seen. The pronotal lobes are
pale below, dusky above and the posterior pronota are similar. There is one posterior
pronotal bristle, two to three spiraculars and no upper sternopleural. The pleural
integument is a medium brown. The postnotum is brown with a pair of small but
distinct fine bristles distally on the mid line.

Legs: The coxae are pale yellowish with silvery scales, but the fore coxae are
rather darker anteriorly and the scaling may appear dark in some lights. The femora
are indefinitely pale beneath to the apex and the hind tibia is about 88% the length
of the mid.

Wings: All the wing scales are broad but rather pale. The upper fork cell is about
2-5 times the length of its stem and its base is level with that of the lower cell.

Abdomen: This is dark scaled above with pale lateral triangles from segment III
onwards.

Genitalia: The ninth tergite (Text-fig. 31) has the lobes deeply emarginate but
not widely separated. They are of medium length with eight or nine distal spines
about equal in length to the lobe (see note. below).

The female is not yet known.

Larva.

The clypeal spines are fairly long and slender; head hair A is single and of
moderate length, C is single and very long and a little behind the level of A, B is
directly in front of C and close to the anterior margin and has six or seven branches,
and d is directly behind the clypeal spine and very slightly anterior to B; it also has
about six branches. The antennae are rather stout with the shaft hair single and
about two-thirds from the base. However the most conspicuous feature of the head is
the maxilla which carries an extraordinarily long, rather slender, apically rounded
tooth (Text-fig. 47).

Most of the stellate hairs of the abdomen have from two to four branches, each
being strongly pectinate. The terminations of each spine often comprise a number of
non-divergent points. The integument of the body is invested with minute pale spines.

The lateral comb comprises four to five spines. There appears to be wide variation
in the pentad hairs but the specimens before me are obscure in this region. The
siphon is of moderate length, evenly tapering with eleven to twelve pecten spines
extending in a row for the whole length of the siphon. The basal teeth are minutely
fringed. The ventral hairs are trifid except the basal pair (with six or more branches)
and in all there are about thirteen ventral tufts. The dorsal hairs are four- to five-
branched and the ventral siphonal valve has a long single hair. The saddle covers the
dorsal half of the anal segment, the saddle hair is single and plumose and the dorsal
subcaudal has five or more plumose branches. The ventral tuft on one side of the
cast skin of the holotype has three branches but on the other there are nine. A most
conspicuous feature of the anal segment is its ventral surface which is densely clothed
with short fine dark spines.

Biology: Larvae have been found in cut bamboos in rain forest both at Hollandia
and Mt. Dafonsero, and a larva which is indistinguishable from that of T. longipalpata
was taken from a pitcher plant in mossy forest at an altitude of 1,600 metres.

Distribution: The adult is only known from Hollandia but larvae have been identified from
Mt. Dafonsero, 19.iv.45, and Mt. Dafonsero at an altitude of 1,600 metres also 19.iv.45 (all
collected by 19th Med. Gen. Lab.).

Note.—Another specimen from the same collection as the type JT. longipalpata (date
and habitat and numbered 874-2) only differs in having the palpi a little shorter
(about three-quarters the proboscis) and the hind tibia longer (about equal to the
mid), and the lobes of the ninth tergites similar except that the outermost spine arises
well behind the distal margin (Text-fig. 32) and is associated with a larval skin which is
very similar in the head hairs, the siphon and even the hirsute area ventrally on the
anal segment. However, the extremely long apical spines of the maxillae cannot be

BY DAVID J. LEE. 251

made out, nor can it be decided whether they have been broken off or whether the
maxillae are intact. Further collections are required before it can be established
whether the specimen before me (now in the C.S.I.R. Collection) is a variant of
T. longipalpata or a distinct form. It should also be noted that no postnotal hairs can
be seen.

TRIPTEROIDES (RACHISOURA) CONFUSA, Nn. sp.

Types: Holotype male, allotype female and a morphotype larva in the C.S.I.R. Collection
together with cast larval and pupal skins correlated, but not individually, with the types.
Similar cast skins in British Museum and National Museum, Washington. All collected by
D. A. C. Cameron, 1943.

Type Locality: Milne May, Papua.

DISTINCTIVE CHARACTERS.

The female comes closest to T. filipes and T. latisquama but the relatively longer
hind tibia will distinguish it from the former and the fewer prescutellar bristles from
the latter. In most specimens I have seen, the postnotal bristles are rather characteristic,
being a pair of small but distinct dark hairs arising close together on the mid line
distally on the postnotum, the hairs themselves being widely divergent. The male comes
closest to T. longipalpata but the lobes of the ninth tergite are quite distinct. The
larva is close to TJ. filipes but the trifid ventral beard (instead of six-branched) and
the single hair on the ventral siphonal valves are easily discernible differences.

DESCRIPTION.
Male.

Head: The head is dark scaled with a few pale scales at the middle of the nape
and an obscure and narrow pale border to the eyes. The palpi are two-thirds the
length of the proboscis which itself is equal in length to the fore femur and not
noticeably thickened. The pedicels and the clypeus are yellowish-brown to brown.

Thorax: The integument of the scutum is dark brown and that of the pleura
yellow-brown. The scales of the scutum are broad, curved and dark brown and appear
to be somewhat narrower down the middle. There are no dorsocentrals and only one
prescutellar pair appears to be present. The pronotal lobes and the posterior pronota
are pale below and somewhat darker above. There is one posterior pronotal bristle (a
small second one may also be present), four spiraculars and one strong upper sterno-
pleural. The postnotum is brown with one or two quite strong hairs at the middle
towards the distal margin.

Legs: The femora are pale beneath to about two-thirds from the base, otherwise the
legs are dark. The hind tibia are from 94% to almost equal the mid.

Wings: All the wing scales are broad and rather dark. The upper fork cell is a
little more than twice the length of its stem and its base is slightly nearer the base
of the wing than that of the lower cell.

Abdomen: The tergites are black with distinct lateral pale apical triangles.

Genitalia: The lobes of the ninth tergite are long, rather narrow, deeply separated
and surmounted by seven or eight spines which increase considerably in length from
the inner to the outer margin (see Text-fig. 27).

Female.

The palpi are barely one-quarter the length of the proboscis, the clypeus is dark
brown and the pedicels light brown. The proboscis is equal in length to the fore femur.
One pair of prescutellar bristles is present. The femora are pale beneath to the tip
and the upper fork cell is twice the length of its stem, the bases of both cells being
about level. The abdominal markings are distinctly serrate.

Larva.

The clypeal spines are fairly long and slender. Head hair A is single or bifid,
B is trifid, directly in front of C but near the anterior margin; C is long, single and
well behind the level of. A; d is bifid and immediately behind the clypeal spines.

Z

252 NOTES ON AUSTRALIAN MOSQUITOES. VI,

The antennal shaft hair is single and short and about two-thirds from the base. The
maxillae are as figured in Text-figure 43. There are no modified hairs on the thorax,
the stellate tufts have three to four branches and each branch is finely spinose and
simple ended. The lateral comb comprises four to five pointed spines (Text-fig. 55)-
The first pentad hair has three or four finely and sparsely spinose branches, the second
is single and of moderate length, the third has about five very short branches, the
fourth is single and the fifth has about four branches, again very short. The siphon is
short with three or four pecten teeth (Text-fig. 58), one at the apex and the others
also on the distal half. The ventral tufts are nearly all four-branched, but the basal
pair has about seven branches. The dorsal hairs are two- to four-branched and the
ventral siphonal valves bear a single hair. The anal segment has a small saddle with
the saddle hair single and plumose and there are a few spines on the distal margin.
The anal papillae are very large and pointed. The terminal segments are figured in
Text-figure 56.

Biology: Larvae were found in tins and coconut shells in association with
T. bimaculipes.

Distribution: Known with certainty only from the type locality but probably quite widely
distributed. This is very likely the species which has usually been called T. atra, owing to the
paratype of that name seen by Edwards not being conspecific with the type.

Note.—A female specimen from Hollandia (10.xi.44, 19th Med. Gen. Lab. No. 270,
bred from a treehole in rain forest) closely resembles 7. confusa even to the rather

characteristic postnotal bristles and seems likely to be that species.

VANLEEUWENI-GROUP.

Two characters distinguish members of this group from those of the filipes-group,
namely, the presence of narrow outstanding scales on the posterior wing veins (from
R,,, to Cu,) and the lack of a posterior pronotal bristle. Several of the species have a
few broad pale scales on the clypeus (7. pallida, T. kingi and. T. brevirhynchus), the
rest have no such scales on the clypeus (T. bisquamata, T. vanleeuweni, T. pilosa and
T. simplex). Two of the species, 7. brevirhynchus and T. simplex, are to be grouped
together because the lobes of the ninth tergite are fused and not emarginate.

TRIPTEROIDES (RACHISOURA) BISQUAMATA, 0. Sp.

Types: Holotype male, allotype female, three male and one female paratypes in the C.S.IR.
Collection, two male paratypes in the British Museum and one male and one female paratypes
in the National Museum, Washington. No cast skins were associated with any of the type
series but larval correlations were established through a number of males not used in the type ©
series. Morphotype larvae are lodged in each of the Museums mentioned. The type series was
collected by C. J. Steinhauer. :

Type Locality: Hollandia, Dutch New Guinea.

DISTINCTIVE CHARACTERS.

T. bisquamata comes very close to 7. vanleeuweni but is to be distinguished from the
latter by the somewhat shorter palpi, fewer spiracular bristles and the lack of prescu-
tellars. The single hind tarsal claw of T. vanleewweni is also an unusual feature and
possibly the scutal scales, described as “narrowly-spindle-shaped” may be rather different
to those of T. bisquamata. The broad scales of the upper part of the posterior pronota are
in marked contrast to 7. pilosa, and the lack of clypeal scales distinguishes it from
T. kingi and T. brevirhynchus, and the dark scaling on the upper part of the posterior
pronota from both the last-named and 7’. pallida. The larva has the ventral hair tufts
of the siphon much more strongly developed than in the last three species and the form
of the lateral comb will distinguish it from T. filipes and its allies.

DESCRIPTION.
Male.

Head: Dorsally the head is dark scaled with a few pale scales at the centre of the
nape, and lateral pale areas. There is a very narrow pale rim to the eyes which is
usually obscured by the overhanging broad dark scales. The pedicels are dark brown
with fine greyish hairs and scales; the clypeus is dark brown, round and bare. The

BY DAVID J. LEE. 253

palpi and proboscis are black, the former 0-75 the length of the proboscis. The latter
is shorter than the abdomen and 0:9 the length of the fore femur.

Thorax: The integument of the thorax is dark brown. The scales of the scutum
are dense, dark, broad and irregularly spindle-shaped, with some fine pale scales at the
anterior margin. There are no dorsocentrals and no prescutellars. The pronotal lobes
and the posterior pronota are pale scaled below and dark scaled above. There is no
posterior pronotal bristle, two to three spiraculars (occasional specimens have up to
five on one side), and no upper sternopleural.

Legs: The coxae are yellowish-brown with pale scales on the outer lateral surfaces.
The femora are pale scaled beneath at least half-way and even to the tips in some lights,
otherwise the legs are dark scaled. The hind tarsal claw is double and the hind tibia
is 87% the length of the mid.

Wings (Plate xii, c): R,, R. and R, are broad scaled but the rest of the veins have
narrow outstanding scales, particularly distally. The base of the lower fork cell is very
slightly nearer the base of the wing than that of the upper cell, the latter being a little
more than twice the length of its stem. \

Abdomen: The dorsum is dark scaled with lateral pale markings; the line of
demarcation between the colours is irregular but not markedly serrate. Tergite VIII
is pale laterally (visible dorsally) and may be pale dorsally as well. The venter is
pale.

Genitalia: The lobes of the ninth tergite are broad, not deeply emarginate, and
carry six to eight spines (Text-fig. 33).

Female.

This is similar in most respects to the male but the palpi are only 0-25 the length
of the proboscis and exceeding the clypeus by twice its length. The upper fork cell is
about three times the length of its stem and the lateral abdominal markings are only
slightly irregular.

Larva.

The clypeal spines are slender, head hair A is long and single, C is similar and well
behind the level of A, B is bifid or trifid and near the anterior margin of the head, and
d is single or bifid and fairly long. The maxilla is as illustrated in Text-figure 48. The
antenna is moderately long with the shaft hair single and situated at about three-
quarters from the base.

The stellate hairs of the thorax and abdomen have three to seven pectinate branches
and each ends in two or more small non-divergent points. The thoracic: chaetotaxy is
illustrated in Text-figure 52.

The lateral comb of the eighth abdominal segment comprises two to five spines
(Text-flg. 54); one or two of these are long pointed teeth, the rest short fringed spines.
The first pentad hair is well developed with about eleven pectinate branches, the second
is long and single, the third is very short with about ten branches, the fourth is long
and single and the fifth similar to the third.

The siphon is moderately long, narrowing apically, the index about three, and the
pecten comprises some five to six teeth disposed in a row from near the base to the
apex. The sixteen to nineteen ventral hairs are long, bifid or trifid except the basal
pair which have some seven branches; all are plumose. The dorsal hairs are well
developed with three to four plumose branches. There is a long four-branched hair on
each of the ventral siphonal valves. The saddle has distinct spines on its distal margin,
the saddle hair is long, single and plumose, the dorsal subcaudal has four or five long
plumose branches and the ventral subcaudal is single and plumose. The ventral beard
comprises a long six-branched plumose tuft. The anal papillae are long, slender and
pointed (see Text-fig. 53 for illustration of terminal segments).

Biology: -The larvae are found in pitcher plants (including climbing Nepenthes),
often growing in swampy grassland. Specimens of Nepenthes in which this species was
found appear to be either N. mirabilis or N. papuana according to Danser’s (1928)
revision of this genus.

Distribution: A common species in the vicinity of Hollandia but it has not been identified
with certainty (due to lack of male specimens) from any other locality.

254 NOTES ON AUSTRALIAN MOSQUITOES. VI,

Lee

Text-figs. 43-50.—Larval maxillae of species of Tripteroides (Rachisoura). All figures x
85 approx.

43. T. confusa. 44. T. filipes sensu Paine and Hdwards. 45. T. filipes. 46. T. fusciplewra.
47. T. longipalpata. 48. T. bisquamata. 49. T. brevirhynchus. 50. T. pallida.

BY DAVID J. LER. 255

TRIPTEROIDES (RACHISOURA) VANLEEUWENI (Edwards).

Epwarps, F. W., 1927.—Nova Guinea (Zoologie), 15: 355 (Rachionotomyia).

Types: Described from four females in the British Museum.

Type Locality: Rouffaer R. (tributary of Mamberano R.), Dutch New Guinea.

Synonymy: Rachionotomyia vanleeuweni, Edwards 1927. Loc. cit.

I have not seen this species but apparently it comes close to T. bisquamata and may
be distinguished in the manner indicated under the description of that species.

To suggest that it is very similar to 7. atripes is misleading. The lack of dorso-
central bristles and the entirely different wing scaling make it immediately distinct
from that species.

Neither the male nor the larva has yet been described.

Biology: No information is available.
Distribution: Only known from the type locality.

TRIPTEROIDES (RACHISOURA) PILOSA, N. Sp.

Type: Holotype female in C.S.1.R. Collection. In case the larval skin is located at any time
the reference number is 1010-9 (19th Med. Gen. Lab., 20.iv.45).

Type Locality: Mt. Dafonsero, near Hollandia, Dutch New Guinea, at an altitude of about
1,250 metres.

DISTINCTIVE CHARACTERS.

The form of the scaling of the posterior pronota (hair-like above) is a most tinusual
feature and should prove to be specifically distinctive.

DESCRIPTION.
Female.

Head: The head is black scaled with a few pale scales at the centre of the nape.
No pale rim around the eyes can be seen. The clypeus is round, bare and dark brown
and the pedicels are also dark brown with fine dark hairs on the inner surface. The
palpi and proboscis are black, the former exceeding the clypeus by twice its length and
being 0-225 the length of the proboscis, which itself is 0-9 the length of the fore femur.

Thorax: The integument is dark brown and the scutum is clothed with broad, some-
what curved spindle-shaped dark scales; these are rather narrow laterally and in the
middle. There are no dorsocentral bristles, three pairs of prescutellars, no posterior
pronotal, seven spiraculars and no upper sternopleural. The pronotal lobes are clothed
with flat white scales, there are similar scales on the lower part of the posterior pronota,
but above, the scales are brownish, curved and hair-like.

Legs: The coxae are light brown, with white scales laterally. The femora are pale
beneath to the tip but otherwise the legs are dark. The hind tibia is 80% the length
of the mid.

Wings: The scaling is as in 7. bisquamata, the base of the upper fork cell is slightly
nearer the base of the wing than that of the lower cell and the former is three times the
length of its stem.

Abdomen: The margin of the lateral pale markings is not distinct but is probably
irregular.

The male and the larva are not yet known.

Biology: The type was bred from a larva found in a pitcher plant (W. E. Brewer).

Distribution: Only Known from the type locality.

TRIPTEROIDES (RACHISOURA) PALLIDA, N. Sp.

Types: Holotype male, allotype female and one paratype of each sex, together with cast
larval and pupal skins of the allotype and male paratype, in the C.S.I.R. Collection. All 19th
Med. Gen. Lab., 23.11.45.

Type Locality: Hollandia, Dutch New Guinea.

DISTINCTIVE CHARACTERS.

Although very similar to 7. brevirhynchus and T. kingi, both the male and the
larva are quite distinct. The female palpi are rather longer than in 7. brevirhynchus
and the general colouration not such a pronounced rusty brown, nor again are the
abdominal markings so obviously serrate. TJ. kingi has the upper border of the posterior

256 NOTES ON AUSTRALIAN MOSQUITOES. VI,
pronota distinctly brown scaled. The differentiation of the larva is discussed under
T. king.

DESCRIPTION.
Male.

Head: Dorsally the head is dark scaled with no pale scaling on the nape but a pale
vim to the eyes is present. The pedicels and clypeus are dark brown, the palpi and
proboscis are black. The palpi are 0-22 the length of the proboscis which itself is 0-9
the length of the fore femur. Several large pale scales are present on the clypeus.

Thorax: The integument is dark brown, the scutal scales are broad and curved
though somewhat narrower along the midline. There are no dorsocentral bristles and
three pairs of prescutellars. The pronotal lobes and the posterior pronota are pale
scaled, there is neither posterior pronotal nor upper sternopleural bristle, but there are
five to six spiraculars.

Legs: The coxae are light brown with pale scales laterally. The femora are pale
beneath to the tip but otherwise the legs are dark scaled. The hind tibia is 80% the
length of the mid.

Wings: The scaling is as in 7. bisquamata. The base of the upper fork cell is
slightly nearer the base of the wing than that of the lower cell and the former is about
2-3 times the length of its stem.

Abdomen: Dorsally the abdomen is dark scaled with the lateral pale areas having
an indefinitely serrate margin.

Genitalia: The lobes of the ninth tergite are short, well divided, with five long spines
distally (Text-fig. 30).

Female.

The female is similar to the male in most respects. However, no scales could be
observed on the clypeus on most specimens examined, and it is thought that they are
seldom present since no pits at which scales might have been attached could be seen.
The palpi barely exceed the clypeus by twice its length. The proboscis is 0:8 to 0:9
times the length of the fore femur. The lateral markings of the abdomen are straight
to segment IV then indefinitely serrate to the apex. The upper fork cell is about three
times the length of its stem.

Larva.

The antennal shaft hair is single and arises at three-quarters from the base; head
hair A is single, C is similar and behind the level of A, and B and d are bifurcate at
about half their length and situated near the anterior margin. The maxilla is as
figured in Text-figure 50.

The stellate hairs have from three to seven branches but are only minutely frayed.
The lateral comb is reduced to a single long smooth spine. The first pentad hair has
about five strong branches, the second is fairly long and single, the third very short
with about five branches, the fourth is single and the fifth bifid and fairly long.

_ The siphon is very short with the siphonal index barely more than one. There are
four or five pecten spines and six ventral hair tufts of which all are bifid or the basal
two may be trifid. The dorsal hairs (of which there are two pairs) are bifid. There
are minute spines distally on the saddle and the saddle hair has three long plumose
branches. The dorsal subcaudal tuft has three finely plumose branches and the ventral
subcaudal one. The ventral beard comprises a long six-branched hair.

Biology: The type series was bred from larvae collected in climbing pitcher plants
by W. V. King. Other adult females were taken biting in sago swamps and rain forest.

Distribution: Hollandia, 23.i1.45 (10 specimens); Nakasawa (2 specimens); all collected
by 19th Med. Gen. Lab. The species has not yet been recorded elsewhere.

TRIPTEROIDES (RACHISOURA) KINGI, D1. Sp.

Types: Holotype male, allotype female, two female paratypes together with the cast larval

and pupal skins of the holotype in the C.S.I.R. Collection. All collected by 19th Med. Gen. Lab.
{W. E. Brewer). :

Type Locqlity: Mt. Dafonsero, Dutch New Guinea (1,230 to 1,450 metres).

BY DAVID J. LEE. 257

DISTINCTIVE CHARACTERS.

A very dark species closely related to T. pallida and T. brevirhynchus. The male
genitalia are distinct from both but the larva is very similar. In 7. pallida there are
only two pairs of dorsal hairs on the siphon (instead of four to five), and no more than
six ventral hair tufts (instead of six to nine). Again, the siphonal index of 7’. pallida is
distinctly less than that of TJ. brevirhynchus. T. kingi probably comes closest to
T. brevirhynchus but its siphonal index tends to be less and the saddle hair has two
(rarely three on one side only) branches as compared with three to four in TJ. brevi-
rhynchus. Further material of the latter is required before more exact means of differen-
tiation can be proposed. The female is darker than both 7. pallida and T. brevirhynchus
but particularly darker than the latter, and in contrast to both, it has the scales on the
upper part of the posterior pronota distinctly brown.

DESCRIPTION.
Male.

Head: The head is black with a silver rim and there is a small pale patch at the
middle of the nape best visible from in front. The pedicels are black with fine dark
hairs on the inner stirface and the clypeus is very dark brown with some three to five
large pale scales. The palpi and proboscis are black; the former are a little more than
one-quarter the length of the latter, which is distinctly longer than the fore femur
(9:8).

Thorax: The integument is dark brown, rather blackish, particularly on the exposed
parts of the pleura. The scutum is clothed with rather broad spindle-shaped scales, the
pronotal lobes with pale scales, and the posterior pronota are pale scaled below and
brown scaled above. There are no dorsocentrals but about three pairs of prescutellar
bristles are present. There is no posterior pronotal nor upper sternopleural bristle but
there are at least four spiraculars. i

Legs: The coxae are brown with pale scaling laterally. The femora are pale beneath
to about half-way and the hind tibia is about 82% the length of the mid.

Wings: The scaling is as in T. bisquamata, the base of the upper fork cell ig nearer
the base of the wing than that of the lower cell and the former is three times the
length of its stem.

Abdomen: This is dark dorsally with lateral pale markings which are serrate on
the distal segments (IV onwards).

Genitalia: The lobes of the ninth tergite are broad, emarginate, with six to eight
strong spines distally (see Text-fig. 34; both lobes are figured since there is some
variation on either side).

Female.

This resembles the male in most respects except that the palpi are only one-sixth
the length of the proboscis and extend beyond the clypeus a little more than twice its
length and the proboscis is only 0-9 the length of the fore femur. There are five or
six spiracular bristles on the allotype but one of the paratypes has up to eleven.

Larva.

Head hair A is long and single, C is behind the level of A, single but not very
long, B and d are near the anterior margin of the head and both bifurcate at some
distance from the base. The antennal shaft hair is rather long, usually bifid and
situated beyond three-quarters from the base.

The stellate hairs have from two to five branches (usually four) with very fine
fraying and simple points.

The lateral comb comprises a single large pointed spine. The first pentad hair
has about seven strong branches, the second is fairly long and bifid, the third very
short with five or more branches, the fourth is similar to the second and the fifth to
the third. The siphon is short with index about 1:5, the pecten comprises two to four
pointed teeth situated on the distal two-thirds, with one or two spines very close to
the apex. The ventral hairs (of which about nine are present) are bifid and the dorsal
hairs (of which there are eight or nine) are also bifid. The saddle is only finely
spinose distally and the saddle hair has two long plumose branches. The dorsal

258 NOTES ON AUSTRALIAN MOSQUITOES. VI,

subeaudal has three plumose branches, the ventral subcaudal one and the ventral beard
four. The anal papillae are long, narrow and pointed. The terminal segments are
illustrated in Text-figure 57.

Biology: Larvae have been taken in pitcher plants and adult females biting during
the day but all collections have been from considerable altitudes.

Distribution: All specimens examined have been from Mt. Dafonsero at altitudes ranging
from 925 to 1,450 metres.

NN

\

Lee \

Text-figs. 51-58.—Larval structures of unornamented species of Tripteroides (Rachisoura)-
51, 54, 55 and 58 x 100 approx.; 52, 53, 56 and 57 x 40 approx.

51. Lateral comb scales of TJ. filipes. 52. Thoracic chaetotaxy (dorsal) of JT. bisquamata.
53. Terminal segments of T. bisquamata. 54. Lateral comb scales of J. bisquamata. 55. Lateral
comb scales of T. confusa. 56. Terminal segments of JT. confusa. 57. Terminal segments of
T. kingi. 58. Pecten spines of JT. confusa.

BY DAVID J. LEE. 259

TRIPTEROIDES (RACHISOURA) BREVIRHYNCHUS Brug.

Brue, S. L., 1934.—Bull. ent. Res., 25: 508.

Types: Type male and female in British Museum. Cotype male and female were deposited
in Institute voor Tropische Hygiene, Amsterdam.

Type Locality: Tanah Merah, Dutch New Guinea.

DISTINCTIVE CHARACTERS.
The wing scaling immediately relates this species to the vanleeuweni-group. It is
closest to 7. pallida but it is generally a rather brighter brown than that species and

the abdominal markings are distinctly serrate. The larva is very close to that of
T. kingi but may be distinguished as indicated under the description of that species.

DESCRIPTION.

(Based on northern Queensland specimens. )
Male.
Head: The head is brown with a distinct pale patch at the centre or generally pale
greyish-brown and there is a distinct pale scaled margin to the eyes. The pedicels and
clypeus are rather light brown, the latter with a very few large pale scales. The palpi
are 0-2 the length of the proboscis which itself is rather stout and about 0-8 the length
of the fore femur.

Thorax: The integument is brown and the scutal scaling mixed broad and narrow
brownish spindle-shaped scales. The pronotal lobes and the posterior pronota are clothed
with flat whitish scales and some rather greyish ones dorsally on the latter. There are no
dorsocentral bristles, two strong prescutellars, no posterior pronotal or upper sterno-
pleural and three or four spiraculars.

Legs: The coxae are light brown with pale scaling; the femora are pale beneath
to the tip.

Wings: The wing scaling is as in 7. bisquamata. The upper fork cell is distinctly
nearer the base of the wing than that of the lower cell and the former is three times
the length of its stem.

Abdomen: This is brown scaled dorsally with pale apical lateral triangles.

Genitalia: The ninth tergite (Text-fig. 35) is undivided, with ten comparatively
long spines distally (eight to twelve are mentioned in the original description).

Female.

This is similar to the male in most details. The pedicels and clypeus are somewhat
darker brown and the palpi exceed the clypeus by twice its length. The hind tibia is
78% the length of the mid.

Larva.

Only fragmentary material from northern Queensland is at present available but
there is general agreement with the description and figures published by Brug. The
maxilla is rather small (see Text-fig. 49).

Biology: This is a nepenthicolous species. Those from northern Queensland were
collected in N. Kennedyi(ana) (? synonym of N. mirabilis).

Distribution: Known from the type locality (Tanah Merah) and Endeavour Strait, north
Queensland (F. Chippendale). It is interesting to note that this adds a further species to the
list of those common to the southern part of Dutch New Guinea and north Australia.

TRIPTEROIDES (? RACHISOURA) SIMPLEX Brug.

Brue, 8. L., 1934.—Bull. ent. Res., 25: 507.
Types: Described from a single male. In British Museum.
Type Locality: Tanah Merah, Upper Digoel R., Dutch New Guinea.

DISTINCTIVE CHARACTERS.

The description of this species is scarcely adequate for it to be placed in its proper
group. Brug indicates that it is very similar to 7. brevirhynchus, and as in that species,
the ninth tergite is undivided. An examination of the wing is necessary before it can
definitely be placed in Rachisoura but this supposition is not in conflict with the details

260 NOTES ON AUSTRALIAN MOSQUITOES. VI,

given in the description. (It will be seen later that species in which the wing scales
are all narrow rarely lack a posterior pronotal bristle). Hxamination of the terminalia
would be necessary to recognize this species although there is still the possibility of
confusion with 7. brevirhynchus. The palpi of the latter are somewhat longer, extending
beyond the clypeus for twice its length instead of only a little more than the length of
the clypeus as in TJ. simplex.

Neither the female nor the larva of this species has yet been described.

Biology: A nepenthicolous species.
Distribution: Only known from the type locality.

SPECIES NOT PLACED IN ANY SUBGENUS.

The following two species must at present be treated with the unornamented species
but their subgeneric position is still in doubt, since neither the male nor the larva of
either is yet known. Nevertheless the two are certainly closely related since the scaling
ot the wings is distinctive. The wing scales generally are rather small, although broad,
and there are sparse narrow outstanding scales at the terminations of the posterior
veins.

TRIPTEROIDES CONCINNA, 0. Sp.

Types: Holotype female and one paratype female in the C.S.I.R. Collection. Both 19th
Med. Gen. Lab., 2.iii.45 (W. R. Fullem and H. W. Cook).

Type locality: Mt. Dafonsero, near Hollandia, Dutch New Guinea, at an altitude of 1,400
metres.

DISTINCTIVE CHARACTERS.

The small broad scales on the upper surface of the wings and narrow scales below
on veins R,,; onwards will separate this species from all other unornamented ones
except T. subnudipennis. From the latter the entirely dark upright forked scales will
be distinctive.

DESCRIPTION.
Female.

Head: The scales of the head are entirely dark dorsally, silvery white laterally
and with a very narrow silvery rim to the eyes. The pedicels and clypeus are brown,
the latter bare. The palpi exceed the clypeus by at least three times the length of the
latter and are 0-2 the length of the proboscis. The proboscis is rather long, equal to the
fore femur and at least as long as the abdomen.

Thorax: The scutal integument is dark brown, the pleural integument brown. The
scutal scaling is dense, dark, rather narrow medially but with broad curved spindle-
shaped scales laterally and distally. There are no dorsocentrals but four pairs of
prescutellars are present. The pronotal lobes and the posterior pronota are pale scaled
below and light brownish above but there is no marked contrast in the colours. There
is a single posterior pronotal bristle, three to four spiraculars but no upper sternopleural.
The pleural scales cover practically all but the anterior half of the sternopleura and are
silvery white. The scutellum is clothed with dark flat scales and the postnotum is
dark brown.

Legs: The coxae are pale with silvery white scales laterally. The rest of the legs
are dark scaled but the femora are pale beneath to the tip. The hind tibia is about
95% the length of the mid. :

Wings (Plate xii, a): The scaling is small but broad on the upper surface, but
there are narrow outstanding scales below distally on veins R,,;, M,, M., Ms., and Cu,.
The base of the upper fork cell is decidedly nearer the base of the wing than that of
the lower cell. The upper cell is almost three times the length of its stem.

Abdomen: This is dark scaled dorsally and yellowish ventrally. The lateral
yellowish scaled areas of the tergites have an indefinitely straight margin.

Both the male and the larva are as yet unknown.

Biology: All specimens examined were caught biting at an elevation of 1,400 metres.

Distribution : Only known from the type locality.

BY DAVID J. LEE. 261

TRIPTEROIDES SUBNUDIPENNIS (HMdwards).

Epwarps, F. W., 1927.—Nova Guinea (Zoologie), 15: 355 (Rachionotomyia).
Type: A single female in the British Museum.
Type Locality: Nassau Mts., Dutch New Guinea (2,400 metres?—stated as 2,400 feet but
as all other altitudes cited in this paper are in metres it seems likely that metres was intended).
Synonymy: Rachionotomyia subnudipennis, Edwards 1927. Loc. cit.

DISTINCTIVE CHARACTERS.

From the description it is difficult to place this species. Edwards relates it to
T. vanleeuweni but states that the wings are much more scantily scaled, the scales
smaller and narrower. To my mind this relates it to T. concinna and it seems likely
that the two comprise a group of their own. The presence of a posterior pronotal
bristle is a further indication that it is not really closely related to 7. vanleeuweni.
However, it should not be difficult to recognize this species because all the upright
scales of the head are yellow. Other details are that the proboscis is as long as the
abdomen and the palpi are less than one-quarter the length of the proboscis. There is
one small yellow posterior pronotal bristle, about three yellow spiracular bristles and
two or three pairs of prescutellars. The scutal scales are moderately broad and
slightly curved and the sides of the abdominal tergites are broadly whitish, the
separation of the colours being straight. The base of the upper fork cell is very
distinctly nearer the base of the wing than that of the lower cell.

Neither the male nor the larva of this species has been described.

Biology: No information is available.
Distribution : Only known from the type locality.

Subgenus MIMETEOMy1IA Theobald.
EDWARDS, F. W., 1932.—Culicidae in Wystman’s Genera Insectorum, Fasc. 194: 76.

The members of this subgenus are all unornamented species which may be
distinguished from all the preceding species on the scaling of the wings, there being
fairly dense, almost linear outstanding scales on all the wing veins.

ATRIPES-GROUP.

,

The members of this group have the proboscis shorter than the abdomen, the male
palpi almost as long as the proboscis and the female palpi about one-sixth the length
of the proboscis. There are modified spines on both the mesothorax and metathorax
of the larva and the lateral comb arises from a lateral chitinous plate. All the species
are closely similar and require care in their differentiation. One species, 7. digoelensis,
is tentatively included in this group but its description is not adequate for it to be
placed with any certainty.

TRIPTEROIDES (MIMETEOMYIA) ATRIPES (Skuse).

SKUSE, F. A., 1899.—Proc. LINN. Soc. N.S.W., 13: 1750 (Culex).
THEOBOLD, F. V., 1901.—Monogr. Cul., 2: 58 (Culex).
TAYLorR, F.. H., 1914.—Proc. LINN. Soc. N.S.W., 38: 750 (Scutomyia).
, 1919.—Ibid., 43: 829 (Mimeteomyia).
EDWARDS, F. W., 1924.—Bull. ent. Res., 14: 362 (Rachionotomyia).
Coouine, L., 1924.—Commonw. Aust. Dept. Hlth. Serv. Publ. (Trop. Div.), 8: 15 (Rachiono-
tomyia—includes larval description).
EDWARDS, F. W., 1927.—Nova Guinea (Zoologie), 15: 352 (Rachionotomyia).
Lee, D. J., 1944.—Atlas of Mosquito Larvae of the Australasian Region. Australian Military

Forces (Restricted) : 19 (larval description).

Type: Type female in Macleay Museum, University of Sydney.

Type Locality: Knapsack Gully, Blue Mountains, N.S.W. (Homebush is the locality listed
first by Skuse but this is not the type specimen.) It should also be noted that the specimen
mentioned by Skuse from Mossman’s Bay-——that is, Mosman—is not 7. atripes but is actually a
specimen of 7. tasmaniensis.

Synonymy: Culex atripes, Skuse 1899. Loc. cit. Culex atripes, Theobald 1901. Loc. cit.
Scutomyia atripes, Taylor 1914. Loc. cit. Mimeteomyia atripes, Taylor 1919. Loc. cit. Rachio-
notomyia atripes, Edwards 1924. Loc. cit. Mimeteomyia apicotriangulata Theobald, F. V., 1910.
Monogr. Cul., 5: 211—-Single female type in British Museum from Kuranda, north Queensland.

262 NOTES ON AUSTRALIAN MOSQUITOES. VI,

DISTINCTIVE CHARACTERS.
The outstanding scales of the wing are all narrow and the prcboscis short (it is
shorter than the abdomen but usually a little longer than the fore femur; in spite
of this, the proboscis is not likely to be considered long since the femur itself is also
short). The black scaled head will distinguish it from the species most resembling it,
namely JT. punctolateralis and the bronzy scaled scutum from J. solomonis. In the
larva the presence of both mesothoracic and metathoracic spines together with the
lateral comb arising partly from a chitinous plate and the stout divided antennal shaft
hair will serve to distinguish it from all but T. solomonis (but see below under
T. solomonis for further remarks concerning that form). The bluntly rounded points
of the lateral comb spines arising from the lateral plate are also important.

DESCRIPTION.
Female. °

Head: The head is black sealed with a narrow but conspicuous silvery border to
the eyes. The proboscis is shorter than the abdomen and in the holotype it is equal
in length to the fore femur but in most other specimens I have examined it is slightly
longer. The female palpi are one-sixth the length of the proboscis and those of the
male are equal in length to the proboscis.

Thorax: The thoracic integument is dark brown and the scutum is clothed with
moderately large light bronzy scales with a patch of white scales above the wing root
and in front of the scutellum. Dorsocentral bristles are present at least posteriorly
with three to five pairs of prescutellars. A single posterior pronotal bristle is present
in some specimens but in others this cannot be made out; there are usually two
spiracular bristles. The pleura are clothed with flat white scales and the posterior
pronota are pale scaled below and bronzy or grey above.

Legs: The coxae are pale scaled laterally, the femora and tibiae are pale beneath
but otherwise the legs are dark. The hind tibia is about 90% the length of the mid.

Wings (Plate xii, e): The scaling is dense, the outstanding scales on all veins
long and narrow. The upper fork cell is 2:5 the length of its stem and its base is
considerably proximal to that of the lower cell.

Abdomen: The tergites have large apical lateral patches of white in the form of
distinct triangles giving a markedly serrate junction to the colours. The venter is
pale.

Male.

This sex closely resembles the female except that the palpi are as long as the
proboscis. The lobes of the ninth tergite (Text-fig. 41) are not widely separated, longer
at the outer margin and bearing about five strong spines. The venter is banded at least
on the posterior segments.

Larva.

The antennal shaft hair arises at about two-thirds from the base and is a stout
divided spine; beyond this the antenna is distinctly narrower than below. The clypeal
spines are moderately stout and curved with a tuft of very fine hairs at the tip. Head
hair A is long and trifid, B is long with four to six branches and C is four- to five-
branched. Both the latter are well in front of the level of A. Hair d is further in
front still and bifid.

Both mesothoracic and metathoracic spines are present (Text-fig. 68). The stellate
tufts of the thorax and abdomen are strongly developed, each branch being plumose
with a minutely divided tip.

The lateral comb arises from a chitinous plate, the teeth are long, strong, with
rounded tips and below those arising from the plate are a further series of finer spines
often extending to the ventral surface. The first pentad hair is a strongly developed
stellate tuft, the second single, the third with about four branches, the fourth long and
single and the fifth similar to the third. The siphon is short with twelve to thirteen
ventral tufts most of which are bifid but the basal pair may be trifid. The dorsal hairs

BY DAVID J. LEE. 263

are mostly bifid, the lower ones being fairly stout. The saddle has a strong fringe of
spines distally and the saddle hair has three or four branches. The dorsal subcaudal
is five-branched, the ventral subcaudal is single and the ventral tuft has also about
five short branches. The anal papillae are short and pointed. The terminal segments
are illustrated in Text-figure 69.

Biology: The larvae are usually found in treeholes, tanks and water barrels.

Distribution: I have seen specimens from the vicinity of Sydney, Queanbeyan, 20.11.43
(D. F. Waterhouse), and Mt. Victoria, 13.1.44 (R. H. Wharton).

TRIPTEROIDES (MIMETEOMYIA) PUNCTOLATERALIS (Theobald).
THEOBOLD, F. V., 1903.—Entom., 36: 156 (Stegomyia). :

EDWARDS, F. W., 1927.—Nova Guinea (Zoologie), 15: 352 (Rachionotomyia).

Types: Types of both sexes presumably in British Museum.

Type Locality : Southern Queensland.

Synonymy: Stegomyia punctolateralis, Theobald 1903. Loc. cit. Rachionotomyia atripes
var. punctolateralis, Hdwards 1927. Loc. cit. Tripteroides atripes var. occidentalis Brug, S. L.,
1934. Bull. ent. Res., 25: 506.

DISTINCTIVE CHARACTERS.

T. punctolateralis appears to be a distinct species, closely allied to 7. atripes, but
differing in its generally paler colouration. The head is fawn above, instead of black,
the scales of the posterior pronota are distinctly white all over, and the scutal scaling
is generally a paler brown. The proboscis is shorter than the abdomen but longer (7:6)
than the fore femur. The scales of the scutellum are very light, appearing whitish in
some lights, fewer dorsocentral bristles are present, and in the male, at least the basal
third of the palpi are whitish scaled and there are pale reflections at the base of the
proboscis as well. Some pale scaling is present at the base of the proboscis of the
female also but it may be difficult to see. The pale abdominal bands are visible dorsally
and even unite above on segment VIII.

I have taken as a basis for this species a series of specimens in the Macleay
Museum, which although unlabelled as to collector or locality, were separated as
T. atripes var. punctolateralis and could be recognized as specimens collected and
mounted by Bancroft. A similar series in the Brisbane Museum, examined by Miss
E. N. Marks, is almost certainly the same and neither series agrees with Theobald’s
description in certain characters. Nevertheless, it seems likely that Theobald overlooked
the pale scaling at the base of the proboscis and palpi when he described this species
and this has been the cause of subsequent confusion. It is significant that Edwards
(1927) considered specimens from Atapoepoe, Timor, as the same as T. punctolateralis
even though Brug later described these same specimens as a new variety (7. atripes
var. occidentalis). Other material that I have examined from northern Queensland
and the Northern Territory cannot be differentiated from the supposed series of
T. punctolateralis and also fits Brug’s description of 7. atripes var. occidentalis perfectly.
Hence I feel that the latter must now go into the synonymy of 7. punctolateralis.

Text-figure 42 illustrates the ninth tergite of a male specimen from the T.
punctolateralis series in the Macleay Museum. Although the separation of the lobes
appeared distinctly angular in this specimen, in contrast to the rounded indentation
usually found, other specimens examined from the Northern Territory had the usual
rounded indentation and so it does not appear possible to differentiate the genitalia of
this species from those of T. atripes.

The only series of larvae correlated with specimens identified by me as
T. punctolateralis came from Adelaide River, Northern Territory, and although they
are almost identical with those of 7. atripes it does appear possible to distinguish them
on the branching of the dorsal head hairs. Hair A is usually bifid (instead of four-
branched), B has three branches (instead of six), C has three to five (instead of usually
five), and in both, d is bifid. All other characters appear to be as detailed for 7. atripes.
A larval specimen from Karumba was in agreement with those from Adelaide River.

264 NOTES ON AUSTRALIAN MOSQUITOES. VI,

Biology: Bancroft (1908) records this as a biting mosquito and says that the larvae
are found in water butts and tanks. The Adelaide R. material was bred from larvae
found in rot holes in fallen logs.

Distribution: South Queensland (Bancroft); Karumba, north Queensland (F./Sgt. S. J.
Stewart, R.A.A.F., 20.xi.44); Adelaide R., Northern Territory (A. R. Woodhill, 1943) and
Atapoepoe, Timor.

TRIPTEROIDES (MIMETEOMYIA) SOLOMONIS (Hdwards).

Epwarps, F. W., 1924.—Bull. ent. Res., 14: 363 (Rachionotomyia).

PAINE, R. W., and Epwaprps, F. W., 1929.—Ibid., 20: 308 (Rachionotomyia—description of larva).
Type: Type female in British Museum.
Type Locality: Tulagi, Guadalcanal, Solomon Islands.
Synonymy: Rachionotomyia solomonis, Edwards 1924. Loc. cit.

DISTINCTIVE CHARACTERS.

Edwards differentiated this species from 7. atripes as follows: “general colouration
much darker; scutal scales black, but with a conspicuous margin of pure white scales
around the bare space in front of the scutellum, alsc more numerous white scales in
front of the mesonotum; dorsum of abdomen with a strong bluish gloss; front and
middle femora much less extensively white beneath; upper fork ceil relatively somewhat
longer’. On this information and on the basis of a single female specimen from Tenaru
(L. J. Dumbleton), I have been able to differentiate 7. solomonis from T. atripes. The:
characters of most importance are the black scutal scales and the restriction of dorso-
central bristles to the posterior third of the scutum; also there is no white scaling at
the base of the palpi or the proboscis (cf. T. punctolateralis) and the lateral abdominal
markings are not visible dorsally.

Again, in the description of the larva of JT. solomonis (Paine and Edwards, 1929),
there is little to distinguish this species from TJ. atripes, the only obvious characters
of value being the lack of a series of finer lateral comb spines arising below those
attached to the lateral chitinous plate and the sharply pointed nature of those arising
from the chitinous plate, these being quite distinct from the blunt spines of T. atripes.

The male has not been described.

Biology: The larvae were found in coconut husks.
Distribution: Tulagi, Ufa, Fai-ami and Tenaru, all localities in the Solomon Islands.

TRIPTEROIDES (MIMETEOMYIA) DIGOELENSIS Brug.

Brue, 8S. L., 1934.—Buwll. ent. Res., 25: 506.
Type: Single male in British Museum.
Type Locality: Tanah Merah, Upper Digoel R., Dutch New Guinea.

DISTINCTIVE CHARACTERS.

This species is scarcely adequately described but it apparently does not come into
the group of species with long proboscis (proboscis is equal to fore femur). The three
spiracular bristles are yellow, there is a posterior pronotal bristle but no upper sterno-
pleural. The base of the upper fork cell is nearer to the base of the wing than that
of the lower cell and the former is 3-5 times the length of its stem. The separation of
the abdominal colours is straight (this is quite distinct from the other species with
short proboscis and long male palpi) and the lobes of the ninth tergite are deeply
separated, each lobe bearing six bristles a little longer than the lobes.

Neither the female nor the larva has yet been described.

Biology: A nepenthicolous species.
Distribution: Only known from the type locality.

CALEDONICA-GROUP.

The members of this group are distinguished by the proboscis which is longer than
the abdomen and slender. Three species, 7. caledonica, T. rotumana and T. tasmaniensis,
are readily recognized by the confinement of the pleural scaling to longitudinal bands.
The rest of the species have the pleura normally covered, with pale scales over almost

BY DAVID J. LEE. 265

the whole surface. Of these 7. argenteiventris and T. atra are very closely related, with
no striking characters of distinction, 7. microlepis has the scutal scaling very small
and 7’. collessi is distinguished by its banded venter in both sexes.

TRIPTEROIDES (MIMETEOMYIA) CALEDONICA (Edwards).

EpWaArps, F. W., 1922.—Bull. ent. Res., 13: 100 (Rachionotomyia).

Buxton, P. A., and HopKINs, G. H., 1927.—Res. in Polynesia: 74 (Rachionotomyia—description
of larva).

Len, D. J., 1944.—Atlas of Mosquito Larvae of the Australasian Region. Australian Military
Forces (Restricted) : 22 (larval description).
Types: One male and one female cotypes in British Museum.
Type Locality: Houailou, New Caledonia.
Synonymy: Rachionotomyia caledonica, Edwards 1922. Loc. cit.

DISTINCTIVE CHARACTERS.

This species is only likely to be confused with T. rotumana from which it differs
in having the pale abdominal bands situated apically and continuous across the dorsum
instead of basally and laterally as in 7. rotumana. The larva, although superficially
resembling 7. bimaculipes, is quite distinct since hair 7 of the mesothorax is not
noticeably modified. 7. alboscutellata is similar in this feature but has the lateral
comb of the eighth abdominal segment arising from a distinct chitinous lateral plate.
The differentiation of 7. rotumana is detailed under that species.

The proboscis is very long and slender, the female palpi extend beyond the clypeus
almost twice its length and the male palpi are about two-thirds the length of the
proboscis. The scutal scaling is very small, narrow and curved, and the pale pleural
sealing is confined to a median diagonal band beneath which the integument is pale,
in strong contrast to the dark brown integument above and below. Dorsocentral
bristles are present, together with about five pairs of prescutellars, a single posterior
pronotal, one to three spiraculars and an upper sternopleural. The abdomen has the
tergites pale banded apically, the bands being of even width and continuous across the
dorsum. The venter is entirely pale. The lobes of the ninth tergite are small, distinctly
separated, with from three to seven long spines distally (see Text-fig. 40).

Larva.

The clypeal spines are only slightly curved, head hair A is bifid and B, C and d are
single. Their disposition is apparent in Text-figure 64. The antennal shaft hair
is bifid.

The mesothoracic hairs (2-7) are shown in Text-figure 65. Hair 7 of the metathorax
is a strongly developed spine with one very long and one short branch (Text-fig. 66).

The stellate tufts of the thorax and abdomen have from nine to fourteen branches
each of which is finely spinose and terminates in a minutely divided tip and the points
are non-divergent. ;

The lateral comb of the eighth abdominal segment is very regular, comprising some
thirty or more spines, the subdorsal ones being the largest. The first pentad hair is
a well-developed stellate tuft, the second is single, the third is trifid and moderately
long, the fourth long and single and the fifth trifid and also moderately long. The siphon
is rather long with thirteen to fifteen bifid or trifid ventral hair tufts and a distinct
pecten of about eight rather short spines. The dorsal hairs are bifid. The saddle
terminates distally in a very strong fringe of spines the size of which decreases towards
the saddle hair which is bifid. The ventral tuft is a three-branched plumose hair, the
dorsal subcaudal has four such branches and the ventral subcaudal one. The anal
papillae are rather longer than the saddle. The terminal segments are illustrated in
Text-figure 67.

Biology: The larvae were originally recorded from Nepenthes but have since been
found commonly in treeholes.

Distribution: I have examined specimens from New Caledonia, Efate (New Hebrides) and
Havannah (New Hebrides). All were collected by L. J. Dumbleton.

266 NOTES ON AUSTRALIAN MOSQUITOES. VI,

WI

RA

BY DAVID J. LEE. 267

TRIPTEROIDES (MIMETEOMYIA) ROTUMANA (Hdwards).

EDWARDS, F. W., 1929.—Bull. ent. Res., 20: 337 (Rachionotomyia).
Types: Type male, one other male and five females in the British Museum.
Type Locality: Rotuma Island, north of Fiji.
Synonymy: Rachionotomyia rotumana, Edwards 1929. Loe. cit.

DISTINCTIVE CHARACTERS.

This species is only to be confused with 7. caledonica from which it is distinct in
having lateral basal creamy spots on the tergites which are larger and just visible
dorsally on the last few segments, in contrast to the continuous apical bands of
T. caledonica.

According to Edwards (1929) the larva is similar in many respects to that of
T. caledonica but “the metathoracic spines are very inconspicuous and bristle-like, fully
half as long and very little thicker than the hairs in the large thoracic tufts; the two
‘spines’ subequal in length (in TJ. caledonica one is much shorter than the other).
Siphon shorter (index about 2), with rather fewer postero-ventral hair tufts (about
10 instead of 13-16)”.

Biology: The larval habitat has not been recorded.

Distribution: The species is still only known from Rotuma I.

TRIPTEROIDES (MIMETEOMYIA) TASMANIENSIS (Strickland).

STRICKLAND, C., 1911.—Entom., 44: 249 (Stegomyia).
TAY Lor, F. H., 1916.—Proc. LINN. Soc. N.S.W., 41: 564 (Stegomyia).
EXpwaArbs, F. W., 1924.—Bull. ent. Res., 14: 362 (Rachionotomyia).
Les, D. J., 1944.—Atlas of Mosquito Larvae of the Australasian Region. Australian Military
Forces (Restricted) : 21 (larval description).
Types: Described from four females. Type in British Museum.
Type Locality : Tasmania.
Synonymy: Stegomyia tasmaniensis, Strickland 1911. Loc. cit., Rachionotomyia cephasi
Edwards, F. W., 1923. Bull. ent. Res., 14: 8 (Hole Creek, Tasmania—type and one other female
in British Museum). Rachionotomyia tasmaniensis, Edwards 1924. Loe. cit.

DISTINCTIVE CHARACTERS.

This species is closest to 7. caledonica in having rather small scutal scales, dorso-
central bristles and incompletely scaled pleura although the bare area of 7. tasmaniensis
corresponds with the scaled area of T. caledonica. From T. caledonica and T. rotumana,
and indeed all other unornamented species it is distinct in having white tipped femora
and tibiae and the apical tarsi white, if not entirely, then all but a dark dorsal line.

The larvae should not readily be confused with those of other species and the
absence of modified hairs on the thorax, the blunt lateral comb scales and the greatly
enlarged anal papillae should prove distinctive.

Male.

Head: The head is entirely dark dorsally, pale laterally and with an incomplete
narrow pale border to the eyes. The palpi are 0-8 the length of the proboscis which in
its turn is 1:25 the length of the fore femur.

Thorax: The integument is brown to dark brown, the scutal scales are small (but
not as small as in 7. caledonica), narrow and curved. Dorsocentral bristles are present
and there are two to three pairs of prescutellars. There is a single posterior pronotal
bristle, two spiraculars and a strong upper sternopleural. The pronotal lobes are pale
scaled, the posterior pronota pale below and brown scaled above. There is a band of
pale scales on the pleura from the lower part of the posterior pronota distally to the
upper third of the mesepimeron. Below this is a distinct bare stripe, broadest across
the middle of the mesepimeron, and below this again the pleura are pale scaled.

Text-figs. 59-69.—-Larval structures of unornamented species of Tripteroides (Mimeteomyia).
59, 61, 62, 638, 64, 65, 66, 67, 68 and 69 x 40 approx.; 60 x 100 approx.

59-60. T. argenteiventris. 59. Terminal segments. 60. Lateral comb scales. (Hale Yee
tasmaniensis, terminal segments. 62-63. T. collessi. 62. Head (dorsal). 63. Terminal segments.
64-67. TT. caledonica. 64. Head (dorsal). 65. Mesothoracic hairs 2-7. 66. Metathoracic spine.

67. Terminal segments. 68-69. JT. atripes 68. Mesothoracic and metathoracic spines. 69.
Terminal segments.

AA

268 NOTES ON AUSTRALIAN MOSQUITOES. VI,

Legs: The legs are conspicuous for their white tipped femora and tibiae and the
pale scaling of hind tarsi III-V, at least ventrally and laterally.

Wings: The outstanding scales are long and narrow and the base of the upper fork
cell is considerably nearer the base of the wing than that of the lower cell. The upper
cell is also about five times the length of its stem.

Abdomen: This is brown dorsally with pale lateral markings in the form of small
apical triangles interrupted by dark areas at the bases of each segment. The venter is
largely but not entirely pale scaled, brown scales being irregularly present.

Genitalia: The coxites are rather long and narrow, and the lobes of the ninth
tergite are of only moderate size, separated, and carrying some nine distal spines (see
Text-fig. 37).

Larva.

Head hair A is four-branched, B is fairly long and single and behind the level of
A, C is bifid or trifid and close to the anterior margin at the level of hair d which
is bifid. The clypeal spines are fairly long and curved and the antennal shaft hair
is single.

There are no modified hairs on the thorax and the stellate tufts of both the thorax
and abdomen have from five to twelve branches (usually eight to ten on the abdomen)
and each branch is very inconspicuously frayed with a divided but non-divergent point.

The lateral comb comprises a row of some twenty fringed spines with bluntly
rounded ends. The first pentad hair is a strongly developed stellate tuft, the second
is simple, the third with three or four branches, the fourth is exceptionally long and
single and the fifth has some six branches. The siphon is rather stout, its index a
little more than two, with a pecten of about eight multidentate teeth extending from
the base to almost halfway. The two basal hairs of the ventral tufts have four to five
branches and are followed by an irregular series of eleven bifid hairs. All are long and
finely plumose. The dorsal hairs are short and single or bifid. There is a row of
dentate spines at the distal margin of the saddle, the saddle hair is long and single,
there are six branches in the dorsal subcaudal, one in the ventral subcaudal, and the
ventral tuft is trifid; all these hairs are plumose. The anal papillae are very long
and sausage-shaped. The terminal segments are illustrated in Text-figure 61.

Biology: All larval records have been from small rock pools in mountain streams
or treeholes. :

Distribution: The species is known to occur in Tasmania and the eastern portion of New
South Wales. Tasmania: Waratah, 13.iii.16 ; Ferntree Gully, ii.28 (Irwin Smith) ; Mt. Wellington,
1.xii.22 (A. Tonnoir); Harz Mt., 9.xii.22 (A. Tonnoir) ; King R., 4.ii.23 (A. Tonnoir) ; Eagle-
hawk Neck, 23.xi.22 (A. Tonnoir). New South Wales: Barrington, i.25 (S.U. Zoo. Exp.) ;
National Park, 12.iv.25, 1.1.26 (Mackerras) ; Mosman Bay; Mt. Victoria, 3.1.44 (R. H. Wharton) ;
Northwood, 19.xi.40 (A. R. Woodhill).

TRIPTEROIDES (MIMETEOMYIA) ARGENTEIVENTRIS (Theobald).

THEOBALD, F. V., 1905.—Ann. Nat. Mus. Hung., 3: 118 (Polylepidomyia).
EDWARDS, F. W., 1924.—Bull. ent. Res., 14: 361 (Rachionotomyia).
Brue, 8S. L., 1934.—Ibid., 25: 510 (includes larval description).

Types: Described from five females. Type in National Museum of Hungary, one paratype
in British Museum.

Type Locality: Paumomu R., Papua (almost certainly Paimumu R., opposite Yule Island—
146° 45’ BE. by 8° 50’ S.).

Synonymy: Polylepidomyia argenteiventris, Theobald 1905. Loe. cit. ~Rachionotomyia
argentieventris, Edwards 1924. Loe. cit.

DISTINCTIVE CHARACTERS.

T. argenteiventris will not be confused with TJ. caledonica, T. rotumana or
T. tasmaniensis since the pleura are wholly covered with pale scales instead of having
the scaling confined to longitudinal stripes. The large scutal scales immediately
differentiate it from 7. microlepis, and the entirely white venter from T. collessi. As
only the female of 7. atra is known it is only possible to distinguish the female from
the former on the length of the palpi (one-sixth of the proboscis in 7. atra, one-tenth
in T. argenteiventris). The larva of T. argenteiventris will only be confused with that

BY DAVID J. LEE. 269

of T. tasmaniensis, these being the only two species without modified maxillae in which
both mesothoracic and metathoracic spines are absent, but the pointed lateral comb
spines of the former are in contrast to the blunt fringed spines of the latter.

DESCRIPTION.
(Based on Milne Bay material.)
Male.

Head: This is dark, almost black scaled above with the upright forked scales
yellowish at the centre of the nape and a very narrow pale border to the eyes which is
often obscured by the overhanging dark scales. The clypeus is light brown and the
proboscis black, long and slender and 1:3 the length of the fore femur. The palpi are
almost as long as the proboscis (85:95).

Thorax: The scutal scaling is broad, dense and dark brown. The thoracic
integument, including the pleura, is dark brown; there are no dorsocentral bristles
but three to four pairs of prescutellars are present. The pleura are clothed with pale
scales but the upper part of the posterior pronota are dark brown. There is a single
posterior pronotal bristle, two to three spiraculars and one upper sternopleural.

Legs: The femora are pale beneath to the tip, otherwise the legs are dark scaled.
The hind tibia is 88% the length of the mid.

Wings (Plate xiii, e): These are clothed with narrow outstanding scales on all
veins. The base of the upper fork cell is nearer the base of the wing than that of the
lower cell and the former is about 2-7 the length of its stem.

Abdomen: The abdomen is blackish dorsally with lateral pale markings separated
by a straight line. The venter is pale scaled.

Genitalia: The lobes of the ninth tergite (Text-fig. 39) are separated but small,
with some six to ten spines arranged around the margin.

Female.

Generally the female is very similar to the male but there are a few pale flat
scales at the centre of the nape; the palpi are 0:1 the length of the proboscis and exceed
the clypeus by less than 1:5 the length of the latter. The hind tibia is about 92%
the length of the mid and the upper fork cell is 3-0 the length of its stem.

Larva.

The antennae are slightly swollen at the middle and the shaft hair is long, single
and arising at three-quarters from the base. The clypeal spines are moderately stout,
curved and divided into two close-set branches at half-way (in occasional specimens
from both Milne Bay and Hollandia they are not divided). Head hair A has some
four to eight fine, rather long branches; B is similar but usually with fewer branches
and is situated in front of the level of A; C is long with two branehes and arises
behind the level of A and d is single or bifid, of moderate length and near the anterior
margin of the head; e is single and f is bifid. All the hairs are very fine and limp.

There are no modified hairs on the thorax. The stellate tufts of the thorax and the
abdomen have rather fine though numerous branches, finely frayed, with their apices
divided into two or more minute points. ,

The lateral comb (individual spines are shown in Text-figure 60) comprises a row
of six to fifteen teeth of which the upper ones are sharply pointed but the more ventral
ones may be rounded and fringed (the latter are not always present). The first pentad
hair is a stellate tuft, similar to the rest, the second is single and rather short, the
third is of similar length with about three or four branches, the fourth is long and
single and the fifth is either similar to the second or bifid. The siphonal index is
about 2-0; the pecten comprises two to six small fringed pointed spines. There are
eleven to twelve ventral tufts mostly bifid or trifid but the basal pair may have four
branches; the dorsal hairs have three to four branches. The saddle has a row of
coarse spines distally and the saddle hair is single or occasionally bifid. The dorsal
subcaudal has about eight branches, the ventral subcaudal is single and the ventral tuft
is usually six-branched. The anal papillae are considerably longer than the saddle and
sausage-shaped. The terminal segments are illustrated in Text-figure 59.

270 NOTES ON AUSTRALIAN MOSQUITOES. VI,

Biology: The larvae have been found in relatively fresh water in tins and old
coconuts at Milne Bay by D. A. C. Cameron and in treeholes at Hollandia by W. V. King.
Some specimens have been taken biting.

Distribution: I have examined specimens from the following localities: Milne Bay, 1943
(D. Cameron) ; ix.43 (Ratcliffe) ; x.43 (W. V. King); Tsili Tsili, ix.43 (W. V. King); Nadzab,
ix.43 (W. V. King); Dobodura, ix.43 (Ratcliffe); x.43 (W. V. King); Lae, xi.43 (Ratcliffe) ;
Hollandia, 28.iii.45; 10.iv.45; 16.ii1.45; 2.iv.45 (all collected by 19th Med. Gen. Lab.) ; undated,
220th Mal. Surv. Unit, U.S. Army.

TRIPTEROIDES (MIMETEOMYIA) ATRA (Taylor).

Taytor, F. H., 1914.—Trans. ent. Soc. Lond., 1914: 190 (Stegomyia).
Nec Epwarps, F. W., 1924.—Bull. ent. Res., 14: 362 (Rachionotomyia).
Nec ————,, 1927.—Nova Guinea (Zoologie), 15: 353 (Rachionotomyia).

Types: Described from three females, of which at least one was not conspecific with the
holotype, which is in the School of Public Health and Tropical Medicine, University of Sydney.

Type Locality: Lakekamu Goldfield, Papua.

Synonymy: Stegomyia atra, Taylor 1914. Loc. cit. Nec Rachionotomyia atra, Edwards
1924. Loe. cit. Rachionotomyia brugi Edwards, F. W., 1927. Nova Guinea (Zoologie), 15: 356
(deseribed from ten females, in the British Museum; type locality, Nassau Mts., Dutch New
Guinea (1,500 metres).

Note.—Due to the fact that Edwards viewed a paratype of 7. atra which was not conspecific
with the type it has usually been assumed that 7. atra was a species with all the outstanding
wing scales broad (hence a Rachisoura). The type, however, has only narrow outstanding
wing scales and appears to agree in all details with JT. brugi, hence I have placed the latter
in its synonymy.

DISTINCTIVE CHARACTERS.

I have identified a series of fifteen female specimens from various elevations on
Mt. Dafonsero as this species. They are very similar to 7. argenteiventris but may
best be distinguished on the length of the palpi which are one-sixth the length of the
proboscis and extend beyond the clypeus for twice the length of the latter. Amongst
the series are two specimens in which the palpi are shorter and come within the
limits of 7. argenteiventris but it is unlikely that they are really this species. The
length of the upper fork cell may also be useful, it varies from 2:5 to almost 3-0 times
the length of its stem.

Edwards, in his description of 7. brugi, does not mention the lack of dorsocentral
bristles and in his comparative table indicates that only one pair of prescutellar
bristles is present. In the specimens before me the alveoli of three or four pairs of
prescutellars may be seen and in a specimen from the Upper Rouffaer R., identified by
Brug as T. brugi, two pairs of prescutellars are quite distinct.

Neither the male nor the larva is yet known.

Biology: The adult female is a diurnal biting species, most specimens from Mt.
Dafonsero having been taken biting.

Distribution: I have examined specimens from the following localities: Upper Rouffaer R.,
i.27 (V. Leeuwin); Mt. Dafonsero, biting between noon and 4 p.m. in mossy forest, 18.iv.45;
biting at altitudes from 950 to 1,385 metres, 1.iii.45; biting in rain forest at 770 metres,
21.iv.45; biting at creek at noon, 770 metres; mountains above Doromena, 21.iv.45; biting in
rain forest, 400 metres (all the above collected by 19th Med. Gen. Lab.).

TRIPTEROIDES (MIMETEOMYIA) MICROLEPIS (Edwards).

EDWARDS, F. W., 1927.—Nova Guinea (Zoologie), 15: 353 (Rachionotomyia).
Type: Described from a single female in the British Museum.
Type Locality: Nassau Mts., Dutch New Guinea (1,500 metres).
Synonymy: Rachionotomyia microlepis, Edwards 1927. Loc. cit.

DISTINCTIVE CHARACTERS.

I have identified some twenty-seven specimens taken on Mt. Dafonsero as
T. microlepis since they agree very closely with the description of that species. Although
Edwards describes the species (from a single specimen) as lacking a posterior pronotal
bristle I have found that this is usually but not always present. It is often strong,
sometimes weak and in other specimens not even a pit from which such a bristle may

BY DAVID J. LEE. 271

‘have arisen can be seen. According to Edwards the spiracular bristles are golden but
in most of those before me they are black but in occasional specimens they are pale
brown. The only other character at variance from the description is the abdominal
markings which are usually indefinite but occasionally they may appear triangular on
the more posterior tergites. Since the specimens before me almost certainly comprise
a single species I feel that, in view of their variability, it would not be sound to
consider them distinct from T. microlepis.

The species may be recognized by the long slender proboscis in association with
the very small scutal scaling. In general characters there is considerable resemblance
to T. caledonica and T. rotumana but the evenly dark pleural integument of 7. microlepis,
together with the almost complete covering of the pleura by pale scales, will at once
be distinctive.

DESCRIPTION.

(Mt. Dafonsero material.)
Female.

Head: The head is black scaled above, pale laterally and with a silvery rim to the
eyes. The pedicels and clypeus are dark brown to black and the latter is bare. The
palpi are short, exceeding the clypeus by about 1-5 to 2-0 times the length of the latter
and the proboscis is long and slender and 1-25 the length of the fore femur.

Thorax: The thoracic integument is usually very dark, almost black. The scutal
scaling is dark bronzy, the individual scales being very small, narrow and curved.
There is a narrow area of pale scales above the wing root and at the anterior margin
of the scutum. Dorsocentral bristles are present and there are two to three pairs of
prescutellars. A dark posterior pronotal bristle is usually present but does appear to
be absent in some specimens, there are two to four spiraculars and one or two pale
upper sternopleurals. The pronotal lobes are pale scaled and the posterior pronota are
pale scaled below and dark on the upper half.

Legs: The femora are pale beneath to the tip and there are also scattered pale
areas beneath the tibiae. The hind tibiae are about 86% the length of the mid.

Wings (Plate xiii, a): The outstanding scales are long and narrow. The upper
fork cell is about 2-3 times the length of its stem and its base is distinctly nearer the
base of the wing than that of the lower cell.

Abdomen: This is dark above and the pale lateral areas of the tergites are not
pronounced, with an indefinite margin. The venter is entirely pale.

Neither the male nor the larva has yet been discovered.

Biology: All the specimens I have examined were taken biting during the day at
high altitudes.

Distribution: Apart from the type locality this species is only known from Mt. Dafonsero,
Dutch New Guinea. Mt. Dafonsero (1,385 metres), 2.ii1.45; 4.11145; (1,445 metres), 19.iv.45;
(950-1,385 metres), no date (all collected by 19th Med. Gen. Lab.).

TRIPTEROIDES (MIMETEOMYIA) COLLESSI, nh. sp.

Types: Holotype male, allotype female, four male and four female paratypes, all with
associated larval skins in the C.S.I.R. Collection; one male and two female paratypes with
cast skins of the latter two in the British Museum; one paratype of each sex in the Macleay
Museum; one male and two female paratypes with cast skins in the National Museum,
Washington, and one paratype of each sex in the Brisbane Museum. All collected by D. H.
Colless, 10.i.45.

Type Locality: Upper Barron, north Queensland.

DISTINCTIVE CHARACTERS.

This species is quite a distinct one. The presence of dorsocentral bristles, the long
proboscis, pale base to the palpi, distinct apical lateral triangles to the abdominal
tergites together with the banded venter should distinguish it from all related species.
The multi-branched head hairs of the larva (including the antennal shaft hair) together
with the spinose siphon are distinctive characters.

272 NOTES ON AUSTRALIAN MOSQUITOES. VI,

DESCRIPTION.
Male.

Head: Above, the head appears almost white in most lights, pale brownish in
some. The pedicels and clypeus are brown, the palpi and proboscis are dark brown,
the latter are pale scaled at the very base and both have pale reflections basally. The
palpi are 0-75 the length of the proboscis which itself is slightly longer than the
abdomen and 1-2 the length of the fore femur.

Thorax: The thoracic integument is dark brown, the scutal scales are dense, dark
bronzy brown, moderately narrow and curved. Dorsocentral bristles are present
together with three to five pairs of prescutellars. There is flat pale scaling over most
of the pleura except the anterior half of the sternopleuron, the pronotal lobes are pale
sealed and the posterior pronota pale below and bronzy brown above. There is a single
posterior pronotal bristle, two spiraculars but no upper sternopleural.

Legs: The coxae are very pale with pale scaling laterally, the femora are pale
beneath and at least the hind tibiae may appear so in some lights. Otherwise the legs
are dark brown with the hind tibiae about 87% the length of the mid.

Wings (Plate xiii, d): The outstanding scales of all veins are long and narrow.
The base of the upper fork cell is nearer the wing base than that of the lower cell
and the length of the upper cell is 3:5 that of its stem.

Abdomen: This is dark brown above with distinct apical lateral pale triangles
which do not extend quite to the bases of the segments. The venter is pale with very
obvious dark basal bands on segments III—VII.

Genitalia: The lobes of the ninth tergite (Text-fig. 38) are well separated, rounded
apically with from nine to fifteen spines.

Female.

This sex resembles the male closely but the palpi are very short, barely 0-1 the
length of the proboscis and exceeding the clypeus by scarcely more than its length and
with a distinct pale scaled basal area. The proboscis is 1:25 the length of the fore femur.
Larva.

The antennal shaft hair arises not much beyond half way and has twelve or more
branches. The clypeal spines are long, stout, divided into two near the base, each arm
tapering to a fine curved hair. The head hairs are most striking. A, B and C arise
more or less in a line behind the level of the antennae, hair d is situated at somewhat
less than half way to the anterior margin and all these hairs have from ten to twenty
radiating plumose branches. Hairs e and f are bifid or trifid (see Text-fig. 62 for
illustration of head). .

The mesothoracic hairs are not obviously modified but there is a strongly developed
spine on the metathorax with three (occasionally more) very uneven branches. The
stellate hairs of both the thorax and abdomen are very strong with at least twenty
dark plumose branches.

The lateral comb comprises a close set row of very many long fine spines of which
those towards the ventral surface are the longest. The first pentad hair is a strongly
developed stellate tuft, the second is long and single, the third is strong and multi-
branched, the fourth long and single and the fifth stronger than the third and also
multi-branched. The siphon tapers to the tip and its index is almost 3:0. There are
some fifteen three- to four-branched plumose hair tufts ventrally but the pecten cannot
be differentiated from the complete covering of pecten-like spines over the whole
siphon. The dorsal hairs have about fifteen branches in the basal hairs decreasing to
four or even two in the apical ones. The saddle has a strong fringe of long fine spines
distally and the saddle hair has six plumose branches. The dorsal subcaudal is
eight-branched, the ventral subcaudal is single and the ventral tuft has eight or nine
branches. The anal papillae are scarcely as long as the saddle. The terminal segments
are illustrated in Text-figure 63.

Biology: The type series was bred from larvae found in treeholes.

Distribution : As yet this species is only known from the type locality.

BY DAVID J. LEE. 273

OBSCURA-GROUP.

Only two species belong to this group, namely, 7. obscura and T. subobscura. They
are distinct from all other species of Mimeteomyia because of the palpi being about
one-third the length of the proboscis in both sexes.

TRIPTEROIDES (MIMETEOMYIA) SUBOBSCURA, Nl. Sp.

Types: Holotype male and allotype female in the C.S.I.R. Collection and a paratype of
each sex in the British Museum. All collected November, 1942, by I. M. Mackerras.
Type Locality: Jacky Jacky, Cape York, north Queensland.

DISTINCTIVE CHARACTERS.

Both sexes are distinguishable from all but 7. obscura on the length of the palpi.
From the latter the lack of a posterior pronotal bristle and the strongly serrate
abdominal markings are characters of distinction.

Note.—This species closely resembles 7. obscura Brug even in the form of the ninth
tergite, but owing to the lack of a posterior pronotal bristle on all four specimens,
together with the very markedly serrate abdominal markings, I have deemed it wise to
consider the present form a distinct species. Further study of material from Tanah
Merah, or a comparison of larvae from the two localities, is necessary before the exact
relationships of the two can be determined.

DESCRIPTION.
Male. :

Head: The scaling is black dorsally. The pedicels and clypeus are very dark and
the latter is bare. The palpi and proboscis are black, the former being one-third the
length of the proboscis and tie latter is short, stout and 0:8 the length of the fore femur.

Thorax: The scutal integument is almost black, the scaling is black, dense, rather
narrow medially but broad laterally. There are no dorsocentral bristles but two pairs
of prescutellars are present. The pleural integument is brown, the scales on the pronotal
lobes are pale with some darker ones dorsally and those on the posterior pronota are
pale below and very dark on the upper half. There is no posterior pronotal bristle,

-four or five spiraculars and no upper sternopleural. The scutellum is black scaled and
the postnotum is bare and dark brown.

Legs: The coxae are yellowish-brown with pale scaling laterally. The rest of the
legs are black although the femora are pale beneath for about the basal two-thirds.
The hind tibia is only 75% the length of the mid.

Wings: All the outstanding scales are long and narrow. The upper fork cell is
three times the length of its stem and its base is distinctly nearer the base of the
wing than that of the lower cell.

Abdomen: This is dark dorsally with distinct lateral pale triangles on the tergites,
the junction of the colours being decidedly serrate.

Genitalia: The lobes of the ninth tergite are long, deeply separated with seven
strong spines as shown in Text-figure 36.

Female.

This sex is very similar to the male. The palpi are not quite one-third the length
of the proboscis and extend beyond the clypeus for four times its length. There are
up to six spiracular bristles.

No larval material is available for description.

Biology: The holotype was bred from larvae found in Nepenthes spp.

Distribution: Only known from the type locality.

TRIPTEROIDES (MIMETEOMYIA) OBSCURA* Brug.
Brue, 8S. L., 1934.—Bull. ent. Res., 25: 504 (TL. obscurus).
Type: Single male in British Museum.
Type Locality: Tanah Merah, Upper Digoel R., Dutch New Guinea.

* The termination has been changed to the feminine in agreement with the genus.

274 NOTES ON AUSTRALIAN MOSQUITOES. V1,

DISTINCTIVE CHARACTERS.

Within the group of species in which the outstanding wing scales are long and
narrow the length of the palpi will distinguish this species from all but T. subobscura,
under which species the characters of distinction are discussed. The ninth tergite as
figured by Brug (1934) is apparently very similar to that of 7. subobscura but there
are only five spines on each lobe.

Biology: A nepenthicolous species.

Distribution: Only known from the type locality.

ACKNOWLEDGEMENTS.

I am particularly indebted to Lieut.-Col. W. V. King and his staff of the 19th Medical
General Laboratory, United States Army, for the extensive collection of Tripteroides
which made the present study possible. The following members of the 19th Med. Gen.
Lab. have been individually responsible for various collections: Capts. D. P. Furman and
H. Hoogstraal, Lieuts. L. W. Saylor and J. P. Toffaleti, S./Sgts. W. EH. Brewer, W. B.
Christ, P. L. Douglas, W. R. Fullem, H. A. Levy and W. T. Nailon, Technical Sgt.
W. A. Shelton, Sgt. C. J. Steinhauer, Cpl. H. W. Cook and Lieut. (j.g.) D. Johnson,
U.S.N.R. Other material has been supplied by Major D. O. Atherton, Sgt. D. A. C.
Cameron, Lieut. F. Chippendale, S./Sgt. D. H. Colless, Lieut.-Col. I. M. Mackerras,
Major F. N. Ratcliffe and Major A. R. Woodhill (all of the A.A.M.C.) and Lieut.
J. Forbes of the 220th Malaria Survey Detachment of the U.S. Army. Miss EH. N. Marks,
of the University of Queensland, has at all times been helpful with material and
suggestions. To Miss G. Burns, of the Department of Zoology, University of Sydney,
I am indebted for the care with which she prepared the photographs of wings. f

REFERENCES.
Bancrort, T. L., 1908.—List of the Mosquitoes of Queensland. Ann. Qd. Mus., 8: 1-64.
Brue, 8S. L., 1934.—Notes on Dutch Hast Indian Mosquitos. Bull. ent. Res., 25: 501-19.

DanseER, B. H., 1928.—The Nepentheaceae of the Netherlands Indies. Bull. Jard. bot. Buitenz.,
(3) 9: 249-438.

Epwarps, F. W., 1927.—New Mosquitoes of the genus Rachionotomyia from New Guinea. Nova
Guinea (Zoologie), 15: 352-56.

, 1932.—Culicidae in Wystman’s Genera Insectorum, Fasc. 194.

GRAHAM, D. H., 1929.—Mosquito Life in the Auckland District. Trans. N.Z. Roy. Soc., 69: .
210-24.

HIuu, G. F., 1925.—The Distribution of Anopheline Mosquitoes in the Australian Region, with
Notes on Some Culicine Species. Proc. Roy. Soc. Vict., 37: 61-77.

LANE, J., and CERQUEIRA, N. L., 1942.—Os Sabetineos da America. Arquiv. Zool. Sao Paulo, 3:
473-849. ;

Lee, D. J., 1944.—Atlas of Mosquito Larvae of the Australasian Region. Australian Military
Forces (Restricted) : 1-119.

and WoopHILL, A. R., 1944.—The Anopheline Mosquitoes of the Australasian Region.
Univ. Sydney Dept. Zool., Monogr. No. 2: 1-209.

Luoyp, F. E., 1942.—The Carnivorous Plants. Chronica Botanica Co., Waltham, Mass., U.S.A.

PaIny, R. W., and Epwarps, F. W., 1929.—Mosquitos from the Solomon Islands. Bull. ent.
Res., 20: 303-16.

EXPLANATION OF PLATES XII-XIII.
Plate xii. All x 225.
(a) T. argyropa. (b) T. filipes. (c) T. bisquamata. (d) T. concinna. (e) T. atripes.

Plate xiii, All x 225.

(a) T. microlepis. (b) T. splendens. (c) T. bimaculipes. (ad) T. collessi. Ce) er:
argenteiventris.

BY DAVID J. LEE. 275

INDEX TO SPECIES OF TRIPTEROIDES MENTIONED IN THE TEXT.
(Synonyms in italics.)

< Page. Page.
alboscutellata Sea WeS einer ALS | dey saiete DOD, kingi ray) Whatetcci yl ey saith hustte bere) Me ame erent Taga DO
apicotriangulata .. .. .. .. .. 222,261 latisquama ee) eh, Gh ey el. ee te Pee DAG
aranoides eee eS Ul ugh aie Pan ES IDI littlechildi Eee HME TORU eH ln i, Sesto
argenteiventris ean onn As snot oO: loneipalpatay Yad Pye eke een tere meyer 2
ALE LODAM Ns aes cal ater end Say wie 29 MMASNESIANE wy eee Wik ease eee) CS oo OG
ACE MRO ae eet Ge ey Se ee Nahe iO microlepis son Mette t,t eget nap Vane 7 NIA MA oe 21774 ())
atripes PE eR UA A deep dhce | ei ay) | Son ADO nepenthicolay way mess) eee st ieremralare
DINMTACUIIPCS Henan e ss Cosh ses Se be are 2S nissanensis St an hae ate ae serch a ER SO
DISGUaAM Aram Seat etee fas) Veta ae Misia toe ede MItidivienteri | Wj Esse oe welorce duis olelng ae lpeyel pare’,
brevipalpis eee Seed see hay Sah eh shes eee AO ODSCUTE A lesen) uke, hedhie seo Mabon Meee Mieweh Bots) Hata,
brevirhynchus ees, mete Ahan hte HOOD) OOCBGWGOKIS - 66 “ao. Go cae a6 ao co 268
brugi MSM ae rere RR els chs. | ius oh are wh tare’ ICO ornata lay pie ay Ml acceis'y Vane ilg Wreted Meee ay mrsliccg basreeroro
ECaledonicamrerwrrcr tek Urania. Th we Lay tab pallida PON cr Skene GR ct nar ctaay Pome | dine far APO, Cy Cp
ceylonensis REWER ToC ecrems 20.40 Woe iat DID papua Mk diet! Mlane-7 he ye tee. Sec hae eew aemeo 4
CODLOSTMRT ace ete Sa one! eee ey Oa 2B DOORS om 6e 86 bu eo 68 -60 BAB
coeruleocephala Bac CRETE cya cece Sey pilosa Nae Becket ¢ Oren TE A Weal Le evonin ARTE SD) D
collessi PR ete A Cncaineetat sovskeit seas we Uap euwia aed punctolateralis eURey ives Tee ea wiaesdy | dee UREN OS
CONCINM AMET WE SE Weise acu eee pitGe wt cee et vores OO purpurata SEG hee MRICS BLN ERE Morn Gie Wa etn ey dk
COMRUSAMEE ene i ims = tot Lk) ed 2 Oh QwUasionnatar we fee ee Sate) eaves Peeeema ont)
digoelensis cea Aten cate ater Ti aocy, we Oe rotumana Lea Meare aa s WA coshe te oy 1. Ruste aa atoeare Ont
GISti ein sere tmp ie ce tth tethered (eh iare:, Gratis wre OAL Simplex’ 3 sev Spe Ue Ne NU Pe ee pee OO
CLS SATIS Miricueee ce (oesiss revere versl | ee) were! hore OOD solomonis ee RSE RT UR NOEL RR RERERA AT ciee FADE.
filipes PE yaad) ive: asap uence cea, ee splendens SE ee ee ee ee at en Hae
fuliginosa RMN. Kp eres on che, ile Loren eho subnudipennis Src Says Listed, Susi» atees area Gill
fusea en Uae im ccser motets. tiie cite w | lohan peraiar subobscura Seca tate ernie cisysOle cose) geese a mmc
fuscipleura PTE ee tas, eal) cnn 7e4O sylvestris ee aA od NANT Soa, SEA. SOLO aS
POE 5 oo DE PT ke ee ee ee eee EY tasmaniensis si = Sigaiso8 wesc (fscv eee ge 126"
SLOT OLOME MN Sa ee rs Sis oe nS vanleeuweni <.0 ses) )sec04 2 ie aero. Sie eae DD

BB

276

OBSERVATIONS ON THE MORPHOLOGY AND BIOLOGY OF THE SUBSPECIES
OF ANOPHELES PUNCTULATUS DONITZ.

By A. R. Woopuitt, Department of Zoology, University of Sydney.*
(Three Text-figures and one Map.)

[Read 26th September, 1945.]

INTRODUCTION.

The interpretation of the various forms occurring within the species A. punctulatus
Doénitz has been, and still is, the subject of considerable discussion.

In a recent paper D. J. Lee and the author (Lee and Woodhill, 1944), on
geographical and ecological evidence, substantiated two subspecies, A. punctulatus
punctulatus Don. and A. punctulatus moluccensis Swell, Swell de Graf., and put forward
the theory that forms which were intermediate between these two were actually hybrids,
which occurred where the subspecies existed together. At the same time it was pointed
out that if the form with the dark proboscis, occurring in the New Hebrides, which
was described by Laveran in 1902 as A. farauti, could not be differentiated from
A. punctulatus moluccensis from Australia and New Guinea, then the former name
would have priority and should be adopted. At the time of publication no detailed
descriptions of the larvae of the New Hebridian and Solomon Islands forms were
available, but recent work by Belkin, Knight and Rozeboom (1945) has provided the
necessary information, since their description of A. farauti shows no significant
difference from A. punctulatus moluccensis. However, the arguments previously used
in relation to the subspecific status of this form still hold, and it is therefore proposed
to adopt the name A. punctulatus farauti Laveran rather than the specific designation
A. farauti Laveran as used by Belkin, Knight and Rozeboom (1945).

The following paper gives the results of some detailed observations at Salamaua —
during May-July, 1944, and also the results of re-examination of larval material from
Australia and New Guinea.

MorPHOLOGY OF THE SUBSPECIES.

In the above-mentioned publication (Lee and Woodhill, 1944) the subspecies are
differentiated as follows:

A. punctulatus punctulatus.

Adult.—The proboscis has the apical two-fifths pale scaled.

Larva.—The outer anterior clypeal hairs of the larva are simple; the posterior
clypeals are simple and 0-2 to 0:5 as long as the inners; the antennae are frequently
pale with dark tips. The inner and middle shoulder hairs are transparent with normally
elongated shafts, and arise from small weakly pigmented tubercles which are not
joined to form a single chitinous plate.

A. punctulatus farauti.

Adult.—The adult is not known to be separable from the subspecies A. punctulatus
punctulatus except by the proboscis being entirely dark scaled.

Larva.—The outer anterior clypeals of the larva are simple or weakly branched; the
posterior clypeals may be simple or have up to three or four branches; they vary from
0-1 to 0-5 the length of the inners; the antennae are usually concolorous. Characteris-

* The work embodied in this paper was carried out while the author was serving in the
Australian Army Medical Corps

BY A. R. WOODHILL. PATE

tically the inner and centre shoulder hairs are heavily pigmented, the shaft of the
jnner hair is flattened, and the basal tubercles are heavily pigmented and either enlarged
or joined to form a single chitinous plate. The appearance of the shoulder hairs, and
particularly their basal tubercles, is the most reliable character for separating this
subspecies from A. punctulatus punctulatus.

Since the above was published a further examination of larvae from Australia and
various parts of New Guinea has shown that there are also differences of some value
in the palmate hair of the first abdominal segment. These differences, which were
originally noted by Lieutenant-Colonel W. V. King, are not constant for the whole range
of the species, but hold in certain areas. In A. punctulatus punctulatus from Milne
Bay, Koitaki, Lae, Salamaua and the Solomon Islands, the first abdominal palmate hair
is weakly developed with from six to thirteen slender non-flattened tapering branches,
i.e., not a typical palmate hair. In A. punctulatus farauti from Cairns, Merauke,
Moresby, Milne Bay, Goodenough Island, Salamaua and the Solomon Islands, the first
abdominal palmate hair is strongly developed with from twelve to sixteen flattened
leaflets.

In the Northern: Territory, however, A. punctulatus farauti has this hair weakly
developed as in A. punctulatus punctulatus (see Fig. 3e).

With regard to the antennal colouration of the larvae, as described above, this is
useful in differentiating the subspecies in certain localities, e.g., Milne Bay, but does
not always hold.

At Salamaua one breeding place was found in which the larvae showed all variations
in antennal colouration, but had typical punctulatus shoulder hairs; some 350 adults
were bred out from this series and these were all A. punctulatus punctulatus. The
breeding place was unusual for this subspecies, consisting of a weedy partly shaded
grass-grown pool, with the larvae occurring under floating grass, and thus resembled
the breeding places of A. punctulatus farauti. On the other hand, all the A. punctulatus
punctulatus larvae found in sunlit muddy pools lacking all macroscopic vegetation,
had pale antennae with dark tips. This would suggest that the antennal colouration
is an environmental character, but definite experiments would be necessary toa
substantiate this.

Breeding Areas and Larval Populations.

The principal breeding areas at Salamaua during May to July, 1944, are shown in
Map 1. Three distinct breeding areas were present, separated more or less completely
by natural features such as sago swamps and ridges. Areas 1 and 3 consisted
entirely of open sunny wheel ruts both old and recent, the water of which usually, but
not always, contained a dense suspension of clay. In the majority of the pools no
aquatic vegetation was present. Heavy breeding took place in these pools from May
to July.

Area 2 included some sunlit wheel ruts, but mainly muddy pools in vegetable
gardens, some of which were shaded by dead palm leaves, dried vines, etc., small grassy
swamps, seepages and drains, frequently partly shaded by vegetation. All areas were
kept under observation for some three months, and some 15,000 adults were bred out
from areas 1 and 3, for insecticide experiments. These larvae and adults were constantly
handled and examined during the course of the experiments, and in addition five
separate samples, each of 100 females, were critically examined.

On no occasion was anything except A. punctulatus punctulatus, as defined above,
observed in either larval or adult stages, and it can be reasonably assumed that almost
pure strains of this subspecies were breeding in areas 1 and 3.

In area 2 light to heavy breeding of A. punctulatus punctulatus and very light
breeding of A. punctulatus farauti was found, and very rarely both subspecies occurred
in the same pool. For reasons stated below, however, it appears obvious that fairly
extensive scattered breeding of A. punctulatus farauti must have been taking place
somewhere adjacent to this area, although only very small numbers of larvae could be
located.

278 OBSERVATIONS ON THE SUBSPECIES OF ANOPHELES PUNCTULATUS,

KELA POINT

a
Graves P+ Pt
a REGIST. SB

Vhs
HO OONOOT Hao g BUCO eAsOn),
A ed

WTP pri

ee
DOOI0G e e

Peege”® e
°
°
e

MANGROVES
POSSIBLE BREEDING
PLACE) OF

P— LIGHT BREEDING OF
PUNCTULATUS
F— LIGHT BREEDING OF
FARAUT|
ROADS

AREA BOUNDARIES

SAGO SWAMP

: P+ HEAVY BREEDING OF
we FOUND. : PUNCTULATUS

HALF MILE

Map i1.—Breeding areas of A. punctulatus punctulatus and A.

punctulatus farauti at
Salamaua.

BY A. R. WOODHILL. 279

Estimation of Adult Population.

A series of night catches was made in the three areas, but in areas 1 and 3
practically no adults were obtained, although the catches were made alongside heavy
breeding places. In area 2, however, a number of very successful night catches were
made, the numbers depending on the weather conditions, and possibly other undeter-
mined factors. Although the temperature at Salamaua at 10 p.m. only varied between
76° and 82°F., with an average of 78°F., and the humidity between 89% and 96%,
with an average of 92%, a slight drop in temperature, accompanied by a slight breeze,
appeared to prevent any movement of adults and to reduce the catches considerably.

The inability to take adults at night in areas 1 and 3, as compared with area 2,
may possibly be explained by the fact (a) that there were no human communities in
areas 1 and 3 and the adults may have dispersed over a wide area on emergence,
whereas in area 2 the catches were made in or adjacent to permanent living quarters of
natives and army personnel, or (0) that catches in area 2 were made not only from the
skin when feeding, but from resting surfaces such as walls of huts and tents which
were illuminated by a dim light, whereas in areas 1 and 3 the catches were made
in the open. :

It is to be noted that approximately 50% of the females taken in area 2 were
resting on walls, etc., and not biting.

As in many other localities, no adults could be found resting in tents or huts
during the day time.*

The number of adult females taken in night catches in area 2 between 26th May,
1944, and 4th July, 1944, and determined as A. p. punctulatus, A. p. farauti and
intermediates are shown in Table 1.

TABLE 1.

Date. Number Catching and Time. Total. Ax p: fan: A. p. punct. Inter.
26. v.44 1 native, 8 p.m. to 10 p.m. #5 ue 39 29 5 5
31. v.44 1 native, 8 p.m. to 10 p.m. ee aot 38 16 11 11

6. vi.44 1 native, 8 p.m. to 10 p.m. Ss as 44 2 13 10
13. vi.44 1 native, 8 p.m. to 10 p.m. is we 11 6 3 2
13. vi.44 4 men, 8 p.m. to 6 a.m. Be ae 266 177 60 29
15. vi.44 2men, 10p.m. to 2 a.m. he - 28 19 5 4
23. vi.44

to
27. vi.44 4 men, 8 p.m. to 10 p.m.* ats ae 48 333 4 11
28. vi.44 4 men, 8 p.m. to 6 a.m. a6 Ses 401 175 143 83
1.vii.44 4 men, 10 p.m. to 12 midnight* a 15 8 4 3
4.vii.44 4 men, 11p.m. to 2am. Ear 3h 87 47 14 26
Total .. Pa Sis ae es Me ao 977 531 262 184
Per cent. Ke ae be is se ou 100 54°3 26°8 18:9

* Bad weather.

It is significant that although light to heavy breeding of A. p. punctulatus was
taking place in area 2, while only very light breeding of A. p. farauti could be recorded,
nevertheless 54% of adults were A. p. farauti as compared with 26% A. p. punctulatus
and 18% intermediates. Intermediates were of all types from those with only a small
ventral pale patch on the proboscis to those with a considerable amount of pale scaling
(see Figs. 1, 1b, lc and 1d). The hourly figures from the two all-night catches are
shown in Tables 2 and 3.

* It should be noted, however, that the resting habits of both subspecies vary considerably
in different localities and circumstances. At Nadzab in July, 1944, large numbers of fed
adults were found resting throughout the day on the walls of native huts. The huts were
surrounded by jungle with some open patches of kunai grass, with a stream, including weedy
backwaters, close by. Of 129 adults collected, 85% were A. punctulatus farauti, 3% A.~
punctulatus punctulatus and 12% intermediates.

Lieutenant Brannigan (unpublished data) has also reported large numbers of adults at
Dumpu resting in tents and huts during the day.

280 OBSERVATIONS ON THE SUBSPECIES OF ANOPHELES PUNCTULATUS,

TABLE 2.
All-Night Catches by Four Men, 13th June. 1944.

Time. Total. A. p. farauti. A. p. punctulatus. Intermediates.

8 p.m. to 9 p.m. aes a 9 8 — 1
9 p.m. to 10 p.m. aS os 10 8 1 1
10 p.m. to 11 p.m. a Si 24 21 2 1
11 p.m. to 12 midnight ot 17 11 4 2,
12 midnight to 1 a.m. .. ae 28 19 6 3
1am. to 2 a.m. Bes ae 62 39 15 8
2am. to 3 a.m. ste de 30 23 5 2
3 am. to 4 a.m. a a 38 23 10 5
4 am. to 5 am. pe an 30 19 7 4
5 a.m. to 6 a.m. ae aa 18 6 10 2

Total Bu, a ate 266 177 60 29

TABLE 3.

All-Night Catches by Four Men, 28th June. 1944.

Time. Total. A. p. farauti. A. p. punctulatus. Intermediates.

6.30 p.m. to 7 p.m. se 35 6 4 2 —
7 p.m. to 8 p.m. a oes 18 9 7 2
8 p.m. to 9 p.m. ae Be 19 9 3 2
9 p.m. to 10 p.m. Bi exe 24 Mt 12 5
10 p.m. to 11 p.m. ie at 25 11 8 6
11 p.m. to 12 midnight .. ses 35 12 16 iG
12 midnight to 1 a.m. ae 50 13 25 12
1am. to 2 a.m. ae Es 55 25 17 13
2am. to 3 am. 2S as 55 30 We 8
3 am. to 4 a.m. Fic akc 48 24 11155 9
4 am. to 5 a.m. as ae 42 19 12 11
5 am. to 6 a.m. ss ae 24 12 4 8

Total a Bre és 401 175 143 83

It will be noted that there was a fairly steady rise during the night, with the peak
period occurring between midnight and 5 a.m. without any significant difference between
the two subspecies. On the other hand, Major Atherton (unpublished data), working
at Lalapipi, found that the peak period of adult abundance of A. p. farauti was round
about 9 p.m.

One all-night catch of A. p. farauti made at Cairns by Major Roberts (unpublished
data) gave a fairly even abundance from 9 p.m. onwards with a slight peak round 1 a.m.

Oviposition from Laboratory Bred Females.

It had been noted previously (Lee and Woodhill, 1944) that at least one species
of Australasian anopheline would deposit fertile eggs when bred in small laboratory
cages, and an attempt was therefore made to establish a laboratory culture of

A. punctulatus punctulatus.*

Larvae were collected in the field and the resulting adults, approximately 200 males
and 200 females, were allowed to emerge in a cage 8” x 9” « 15”, and were kept supplied
with sweetened water and given a blood feed every alternate night. As the males died
off approximately 150 additional laboratory bred males were added during a period of
ten days. Under these conditions 170 eggs were deposited in four separate batches
from the 7th to the 13th day after the females emerged. From these eggs about 50
adults were obtained, but these did not deposit eggs.

It would appear that copulation under these conditions only takes place between
a very small percentage of individuals and that laboratory cultures would be very
difficult to maintain. It was felt that considerable information could be obtained if
A. punctulatus punctulatus could be crossed with A. punctulatus farauti in this way.

* As far as is known to the author, Lieutenant F. B. Bang of the U.S. Army Medical
Corps was the first to establish a laboratory culture of A. punctulatus punctulatus.

BY A. R. WOODHILL. 281

As it was not possible to obtain sufficient farauti larvae at Salamaua, the author
suggested to Major I. M. Mackerras of the Medical Research Unit at Cairns that the
attempt should be made there. Some progress has been made along these lines but no
definite results are yet available.

1a

1b

id

1é€

2a

2b

Figs. 1-2b.—Proboscis colouration in intermediate females and their progeny. 1. Inter-
mediate female No. 24. la to le. Progeny of No. 24. 2. Intermediate female No. 20. 2a and 2b.
Progeny of No. 20. All x 54.

A-K-W

282 OBSERVATIONS ON THE SUBSPECIES OF ANOPHELES PUNCTULATUS,

Morphology of Adults and Larvae from Area 2.

In this area, as described above, both A. punctulatus punctulatus and A. punctulatus
farauti were breeding, and collections of adult mosquitoes gave 18% intermediate forms.

In studying the morphology the method used was to obtain eggs from single females
caught in the field and to breed these through, thus obtaining a series of fourth stage
larvae and adults from each female.

The female parents were classified as A. punctulatus punctulatus or A. punctulatus
farauti according to the definitions given above, and all those showing variations in
the scaling of the proboscis were classified as intermediates.

Variations in the palpal colouration of parent and progeny were also noted.

The index used consisted of the distance apart oi the basal tubercles divided by
the distance apart of the bases of the inner and middle shoulder hairs. An index of
0-1 to 0:3 indicates that the basal tubercles are enlarged and almost touching, a
condition which occurs sometimes in A. punctulatus farauti, whereas an index of 0-4 to
0:7 denotes that the tubercles are small and widely separated, i.e., A. punctulatus
punctulatus type.

The following is a summary of adult and larval characters found in the progeny
of single females:

No. 34. A. punctulatus punctulatus.—Palps with apical half of 3rd segment com-
pletely white; progeny, 10 males, 28 females, 43 larvae. Of the 28 females, 22 had palps
resembling the parent, while the remaining 6 had the white apical half of the 3rd
segment interrupted by a few dark scales or a dark ring. All had the proboscis normal
for the subspecies. Of the larvae, 38 had the bases of the shoulder hairs weakly
developed and widely separated (index 0:4 to 0:7), 4 had the bases joined on one side
only, and 1 had these joined on both sides. In all specimens the 1st abdominal palmate
hair was weakly developed, i.e., 6 to 9 branches in the form of slender non-flattened hairs.

No. 45. A. punctulatus punctulatus.—Palps with apical half of 3rd segment only
faintly interrupted by a darker patch; progeny, 6 males, 15 females, 22 larvae. Of the
15 females, 6 had the apical half of the 3rd palpal segment completely white, while
9 had this portion interrupted by dark scales or a dark band; all had the proboscis
normal for the subspecies. Of the larvae, 20 had the bases of the shoulder hairs weakly
developed, with an index of 0:4 to 0:7, while 2 had the bases of the shoulder hairs joined
on one side only. All had weakly developed 1st abdominal palmate tufts as described for .
No. 34.

No. 38. A. punctulatus farauti—Palps with white apical portion interrupted by a
wide black band, proboscis normal for the subspecies; progeny, 1 male, 9 females,
11 larvae. All the female progeny were similar to the parent and all larvae had the
bases of the shoulder hairs joined and the ist abdominal palmate tufts well developed,
with 16 to 18 flattened leaflets.

No. 35. A. punctulatus farauti.—Palps and proboscis as in No. 38; progeny, 5 males,
18 females, 33 larvae. Of the female progeny 16 had the 3rd palpai segment similar to the
parent, while 2 had this segment black except for a narrow white apical ring; all had the
proboscis completely black except for the labella. Of the larvae, 31 had the bases of the
shoulder hairs joined on both sides, while 2 had the bases joined on one side and separate
on the other; all had the ist abdominal palmate hair well developed with flattened
leaflets (see Fig. 3d).

No. 20. Intermediate (Fig. 2)—Proboscis black, except for a minute pale patch
ventrally adjacent to the labella, palps with white apical portion of 3rd segment
interrupted by a black band; progeny, 3 males, 19 females, 4 larvae. The female
progeny in this series showed an extraordinary range of proboscis colouration varying
from forms identical with farauti to those closely resembling punctulatus (see Figs.
2a and 2b).

BY A. R. WOODHILL. 283

The third palpal segment in 17 specimens had the white apical portion interrupted
by a dark band, but in 2 specimens these segments were completely dark except for a
narrow white apical ring. This is a character usually found more frequently in farauti
than punctulatus, but nevertheless these 2 specimens show a considerable amount of

pale scaling in the proboscis.

3d

Figs. 3a-3e.—Structure of shoulder hairs and 1st abdominal palmate hairs in A. punctulatus
a, b, and ec. Variations in shoulder hairs of larval progeny of female No. 24

larvae.
Fig. 1). e. First abdominal palmate hair of larval progeny of female No. 24.
palmate hair of larval progeny of female No. 35. (A. punctulatus farauti from Salamaua. )

a, b, and c x 190, d x 416, e x 390.

(see
d. First abdominal

284 OBSERVATIONS ON THE SUBSPECIES OF ANOPHELES PUNCTULATUS,

The 4 larvae had the bases of the shoulder hairs joined, but the 1st abdominal
palmate tufts were weakly developed with fine non-flattened branches, as in punctulatus.

No. 24. Intermediate (Fig. 1).—Proboscis and palps as in No. 20; progeny, 2 males,
8 females, 31 larvae. The females in this series showed a range of variation in proboscis
colouration similar to the progeny of No. 20, with one specimen practically
indistinguishable from punctulatus, but none identical with farauti (see Figs. la to
le). Of the larvae, 5 had the bases of the shoulder hairs joined on both sides, 8 had
the bases joined on one side and almost touching on the other, while the remaining
18 had the bases of the hairs separated on both sides with an index varying from
0-2 to 0-6, thus showing all variations in the shoulder hairs from farauti to punctulatus.
The 1st abdominal palmate tufts in all the larvae were weakly developed with from
9 to 13 non-flattened branches (see Figs. 3a, 3b, 3c and 3é).

No. 37. Intermediate—Proboscis and palps as in No. 20; progeny, 4 females, 20
larvae. All 4 females were similar to the parent. Of the 20 larvae, 18 had the bases
of the shoulder hairs joined, one had the bases joined on one side and separate on
the other, while one had the bases separate on both sides with an index of 0-4 to 0:5,
i.e., Similar to punctulatus. All larvae had weakly developed ist abdominal palmate
tufts.

No. 50. Intermediate—Proboscis and palps as in No. 20; progeny, 14 larvae. Of
the larvae, 13 had the bases of the shoulder hairs joined on both sides, 1 had the bases
joinéd on one side and separate on the other (index 0-4), and all had the 1st abdominal
palmate tuft weakly developed.

No. 22. Intermediate.—Proboscis mainly pale apically, but with a dark patch
projecting ventrally into the pale area, 3rd palpal segment mainly pale apically without
a distinct black band; progeny, 8 larvae. Of these, 7 had the bases of the shoulder hairs
joined on both sides, while 1 had these separated (index 0-3) on one side. All had the
ist abdominal palmate tufts weakly developed.

No. 43. Intermediate.—Proboscis with irregular pale and dark patches in the apicai
area, but mainly pale resembling punctulatus fairly closely; progeny, 10 larvae, 4
females. Of the larvae, 8 had the bases of the shoulder hairs joined on both sides, while
2 had them joined on one side and separate on the other (index 0-2), while the 1st
abdominal palmate tufts in all specimens were weakly developed.

Table 4 gives a summary of progeny of single females classed according to the
shoulder hairs, and 1st abdominal palmate tufts and female proboscis and palps. The
term “annulipes-like” palps indicates that the apical portion of the 3rd palpal segment
is entirely white, and ‘faint dark band” refers also to this portion of the 3rd segment.
In the first column ‘“punctulatus” or “farauti’” indicates that the proboscis was
indistinguishable from that particular subspecies, while “intermediate” refers to any
form of irregular dark and light scaling.

It will be seen that specimens 34 and 45 gave rise to larvae the majority of which
had all the characters of A. punctulatus punctulatus, but that 7 specimens showed some
A. punctulatus farauti characters.

All the adult progeny resembled A. punctulatus punctulatus.

Specimens 388 and 35 gave rise to larvae and adults all of which showed
A. punctulatus farauti characters.

The remaining intermediate parents gave rise to larval and adult progeny, which
showed a complete range of variation from A. punctulatus farauti to A. punctulatus
punctulatus, except in the first abdominal palmate tufts, which all showed the 4.
punctulatus punctulatus character (see Fig. 3e).

Of special interest is the progeny of the intermediate female No. 24, illustrated in
Figs. la to le and 3a to 3c. It will be noted that although the female parent had only
a very small ventral pale patch on the proboscis, some of the adult progeny were
indistinguishable from A. punctulatus punctulatus, while the shoulder hairs of the
larvae showed a complete range of variation from the A. punctulatus punctulatus to the

285

R. WOODHILL.

BY 7A

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sdjvd OJVIPIUIIOJUT
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sdjed oyvIpoul1oquy
a 8 Z = I 8 = Is 9 éL 8 G 1é se TeulLou
sdjed OyVIPSUl1oquy
= 6. LT z = 6. = a7 <= = = F ¥ as [eulou
sdjed azVIpPoUIoqUy
= 8I —= 8L ane 81 S& = — as G TS S& peutou sdjed wynnwwy
—— 6 = 6 == 6 II iz =e == = IL Il Teuldou sdjed ynnwy
9 6 a == GT GL == CG 06 cae G = GG ae ; pueq
yep Juries ‘sngoynpund
GG 8 = = 8% 8% = SF 8& = t 1 SF : sdjed ox]T
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qualed 9[vulay Jo
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286 OBSERVATIONS ON THE SUBSPECIES OF ANOPHELES PUNCTULATUS,

A. punctulatus farauti type. Again, intermediate female No. 20 gave rise to some adult
progeny (Fig. 2a) with a completely black proboscis, i.e., the A. punctulatus farauti
type.

DISCUSSION.

The theory that the intermediate forms are hybrids between A. punctulatus
farauti and A. punctulatus punctulatus appears to be supported by the field observations
detailed above, since as far as could be ascertained only A. punctulatus punctulatus was
breeding in areas 1 and 3 (Map 1), whereas both subspecies were breeding together in
area 2 where the intermediate forms were obtained. It is significant also that in areas
where only A. punctulatus farauti occurs, e.g., the mainland of Australia and at
Merauke, no intermediates have been recorded from very extensive collections. Inter-
mediates have been taken from most localities in New Guinea where A. punctulatus
punctulatus and A. punctulatus farauti occur together, the percentage of these varying
from 80% at Salamaua (Gilmour, unpublished data) to only a few recorded specimens
at Milne Bay (Allman, unpublished data). The fact that no intermediates occurred in
many thousands of adults bred from open sunny pools in areas 1 and 3 indicates that
their breeding habits probably resemble those of A. punctulatus farauti.

The difference in breeding habits of A. punctulatus farauti and A. punctulatus
punctulatus is emphasized by the fact that in area 2 only light breeding of A. punctulatus
farauti could be located, but adult counts showed over 50% of this subspecies; as
previously recorded by many observers in coastal New Guinea, A. punctulatus
punctulatus larvae are found in all the obvious open sunny breeding places, while
A. punctulatus farauti larvae are usually hidden and far more difficult to locate.

The breeding experiments from individual females also indicate that the inter-
mediate forms are hybrids between the two subspecies since both larval and adult
progeny of intermediate females show all variations from the A. punctulatus punctulatus
to the A. punctulatus farauti type. Since it has been found possible to maintain
continuous laboratory cultures of A. punctulatus punctulatus (Lemerle, unpublished
data), it is to be hoped that workers in this field will attempt to cross A. punctulatus
punctulatus with A. punctulatus farauti on a large scale and thus finally demonstrate
whether inter-fertility occurs between the two subspecies.

In view of the recent work by Owen (1945) and by Belkin, Knight and
Rozeboom (1945) in the Solomon Islands, it must be pointed out that the findings of
these authors in relation to the “punctulatus complex” in that area do not apply in
certain areas in New Guinea and Australia. Reference to Figs. 1-3 will show that
female No. 24 has_ proboscis colouration identical with that described for
A. koliensis Owen, while other individuals which are the progeny of this same female
parent have the proboscis either identical with A. punctulatus punctulatus or inter-
mediate between that subspecies and A. kKoliensis. Again a single female (Fig. 2) with
a proboscis of the type described for A. koliensis gave rise to progeny, some of which
(Fig. 2a) had a proboscis identical with A. punctulatus farauti, i.e., completely dark.

Owen also states that A. koliensis can usually be separated from A. punctulatus by
the absence of a small dark spot on the costal margin of the wing between the basal
and median dark spots; in the progeny of two individual females from Salamaua
(Nos. 34 and 45) with probosces of the A. punctulatus punctulatus type, 7 out of 38
and 7 out of 21 respectively, had this black spot absent, and in many hundreds of
specimens of both A. punctulatus punctulatus and A. punctulatus farauti examined from
New Guinea and Australia this character was found to be extremely variable. Again,
Belkin, Knight and Rozeboom separate A. farauti larvae from those of A. koliensis and
A. punctulatus by the presence of a true palmate tuft on abdominal segment I and the
fusion of the tubercles of prothoracic hairs 1 and 2. Referencé to Figs. 3a, 3b and 3c
shows that the larval progeny of female No. 24 (with koliensis type proboscis) had
shoulder hairs showing all stages between widely separated and completely fused
tubercles. Also the larvae of A. punctulatus farauti from the Northern Territory of

BY A. R. WOODHILL. 287

Australia (where there are no forms with any pale scaling on the proboscis) all have
hair 1 of abdominal segment I with narrow hair-like branches. The branching of the
outer occipitals (used as a distinguishing character for A. koliensis) is also found to
be extremely variable for A. punctulatus punctulatus and A. punctulatus farauti in
New Guinea and Australia, varying from 1 to 5 branchés in punctulatus and from 2
to 6 in farauti.

During the course of examining some thousands of specimens of the subgenus
Myzomyia from various widely separated parts of the Australasian Region from the
Solomon Islands to Morotai, and including the whole of Australia, the outstanding fact
which has been brought out is the extreme variability of such characters as the relative
length of pale and dark bands on the palps, the number of black patches on the costal
margin of the wing, the amount of scaling on the abdomen, the number of branches in
the posterior clypeal and sutural hairs and the form of the shoulder hairs and first
abdominal palmate hairs. Very frequently these characters are found to be quite
constant in a restricted area, but when specimens from widely separated areas within
the Australasian Region are examined, variations in the above characters almost
invariably occur.

It would seem therefore that A. koliensis Owen exhibits constant characters in the
Solomon Islands, but that specimens from New Guinea which are morphologically
identical with A. koliensis, cannot be considered as such since they give rise to
A. punctulatus punctulatus and A. punctulatus farauti in their progeny.

In view of the above it would surely be more logical to regard A. koliensis as a
subspecies of A. punctulatus rather than as a distinct species.

ACKNOWLEDGEMENTS.

The author’s thanks are due to Mr. D. J. Lee for helpful criticism, and to the Director
General of Medical Services, Major-General S. R. Burston, for permission to publish
this paper.

REFERENCES.

BELKIN, J. N., KNIGHT, K. L., and RozmBoom, L. E., 1945.—Anopheline Mosquitoes of the
Solomon Islands and New Hebrides. J. Parasit., 31 (4).

LEE, D. J., and WOODHILL, A. R., 1944.—The Anopheline Mosquitoes of the Australasian Region. ,
Publ. Univ. Sydney, Dept. Zool. Monogr. No. 2.

OWEN, W. B., 1945.—A New Anopheline from the Solomon Islands, with Notes on its Biology:
J. Parasit., 31 (4).

288

NOTES ON NEW SOUTH WALES ORCHIDS: A NEW SPECIES AND
SOME NEW RECORDS.

By the Rev. H. M. R. Rupp, B.A.
(One Text-figure. )

[Read 31st October, 1945.]

1. A New Species of Thelymitra from Brunswick Heads.

THELYMITRA PURPURATA, Nl. Sp.

Planta gracillima, 25-45 cm. alta, caule paullum flexuoso, saepe rubro-purpureo.
Folium angustissime lineare, ad inflorescentiam ascendens; bracteae parvae. Flores
1-7, extra purpurei, intus azurei, in pedicellis brevibus. Sepala petala et labellum
aequalia, 10-17 mm. longa, raro patentia. Columna purpurea; lobi penicillati librati
crinibus pallidis; mitra bifida cristis aequalibus flavis tribus, et circa cristam posteriorem
glandibus fimbriatis coccineis. Anthera obtusa apice subter loborum penicillatorum
bases.

A very slender plant 25-45 em. high, with a slightly flexuous stem, often reddish-
purple except at the base. Leaf very narrowly linear, but channelled, ascending as high
as the middle of the inflorescence. Bracts small. Flowers 1-7, dull-purplish outside,
azure-blue inside, usually with a rudimentary bud above the highest flower. Pedicels
and subtending bracts short; ovary a little more than half as long as the perianth.
Sepals and petals opening very shyly, all equal with the labellum, 10-17 mm. long.
Column bright purple or bluish-purple, the penicillate lobes long, horizontal at least till
the flower is past maturity, blue or purplish with paler upturned hair-tufts. Hood
bipartite, the opening in front quite conspicuous; a broad pale yellow crest behind, and
a narrower one on each side of the partition in front, the crests all equal in height.
The posterior crest almost surrounded by a double fringe of bright red fimbriate glands,
continued as a red band to the front of the hood. Anther obtuse, the apex just lower
than the bases of the penicillate lobes. Stigma large.

Brunswick Heads, ix, 1945, F. Fordham.

I received this plant from Mr. Fordham some years ago, but the specimens, and
others collected later, were too much damaged when I was able to attend to them, to
make it clear whether they should be accepted as a variety of JT. ixioides Sw., or not.
This year excellent specimens were available, and I think it would be quite wrong to
include this beautiful little Sun-orchid in Swartz’s species. Apart from the remarkably
distinctive colouring, the structure of the column differs in important respects. It is,
I think, the most ornate and attractive column of any species of Thelymitra known to
me. In my herbarium there are good specimens of 7. ixioides from Byron Bay, a few
miles south of Brunswick Heads; but except that they are small, these conform to the
type almost precisely, and could never be confused with the plant described above.

2. Some New Records.

Corybas diemenicus (Lindl.) Rupp and Nicholls.—In these Procrerprnes, liii, 1928,
85, Rupp and Nicholls, in reviewing the Australian species of Corysanthes R.Br.,
referred to R. D. Fitzgerald’s unpublished plate of a species found by the late A. G.
Hamilton at Guntawang near Mudgee, which Fitzgerald proposed to call C. Hamiltonii.
After examining the plate and Hamilton’s specimens in the Sydney and Melbourne
Herbaria, we expressed the view that this plant was really Lindley’s Corysanthes

BY H. M. R. RUPP. 289

diemenica, now Corybas diemenicus.. Dried specimens of this genus are notoriously
difficult to determine; but I think we can now reasonably claim that our decision in
regard to Hamilton’s plant has been confirmed. In August, 1945, Mr. Johnson, a school
teacher in the Wellington district—a little further west than Guntawang—sent down
a living plant in flower, which I immediately recognized as a typical OC. diemenicus.
It is rather remarkable that this species, which occurs commonly with C. fimbriatus and
others in the southern States, has never been recorded in New South Wales except on
the far Central Western Slopes, where it does not appear to be accompanied by any
of its relatives.

Fig. 1.—Thelymitra purpurata, n. sp. A, Plant. 3B, Column from the side. C, Column
from the front. D, Column from the back. E, Diagram of column from above. pc, posterior
crest; ac, anterior crests; f, fimbriae; ht, hair-tufts. (B to E all enlarged.)

Caladenia clavigera Cunn.—I received a specimen of this in September, 1945, from
Mr. G. W. Althofer, of Nindethana Nursery, Dripstone. Although this species seems
to be extremely rare in New South Wales, Cunningham’s type locality was the Vale of
Clwydd, near Lithgow. Dripstone is well over 100 miles further west.

Messrs. G. W. and P. Althofer have collected a number of orchids during the current
season (1945), of which there are no definite previous records from the Central Western
Slopes of New South Wales, viz., Thelymitra aristata Lindl., T. nuda R.Br., Diuris lineata

290 NOTES ON NEW SOUTH WALES ORCHIDS.

Messmer, D. flavopurpurea Messmer, D. platichila R. D. Fitzg., D. aurea Sm., Prasophyllum
odoratum Rogers, P. gracile Rogers, Chiloglottis formicifera R. D. Fitzg., Calochilus
Robertsonii Benth., Acianthus reniformis (R.Br.) Schltr., Caladenia filamentosa R.Br.,
C. Fitegeraldii Rupp, C. angustata Lindl., Pterostylis rufa R.Br., P. Woollsii R. D. Fitzg.
They have also sent in living plants of Pterostylis Boormanii Rupp, which was described
from dried material in the National Herbarium of New South Wales.

Glossodia major R.Br.—Specimens were received in September, 1945, from Miss
M. Poynder, collected at Willawong, near Murringo, in the Young district. So far as I
can ascertain, the species has not been previously recorded from that area.

291

CATALOGUE OF REPTILES IN THE MACLEAY MUSEUM.

PART I. SPHENOMORPHUS PARDALIS PARDALIS (MACLEAY) AND SPHENOMORPHUS
NIGRICAUDIS NIGRICAUDIS (MACLEAY).

By STEPHEN J. COPLAND, B.Sc.
(Plate xi; seven Text-figures. )

[Read 28th November, 1945.]

Contents.

Page

I. Introduction oe Be eR. i Sta PE cae, Terme RD te 291

Il. Sphenomorphus par qalis nantes (Macleay) 5, 6 Fin a Ooh Bee LEM At ee eM or Et ne 292

Ill. Sphenomorphus pardalis erro, n. subsp. as Rist y Unies Wiech: vy cacip th et is dant ee ae one 298

IV. Sphenomorphus nigricaudis nigricaudis (Macleay yh ie Sayeed Nisa (PIRES Soar Saas 299

V. Sphenomorphus nigricaudis elegantulus (Peters and Dore Ae ete oes) Ce ceraamecee 305

ViemViierationwand srelationship sof forms) “ae =. 4.) este ae |e sel as | eee 310

VII. Acknowledgements aaa AS «1 POMS EOY POSE AES le LCs URE Samet ws Be eee mance anand ammeter Lats 310
VIII. Bibliography wi8 gO Na GN ene Sains ale Maen Se eres LO Mr nie A ea Aine ton CLT”

I. INTRODUCTION.

There is no need to stress the importance of the reptilian material in the Macleay
Museum at the University of Sydney. The collection of approximately 2,000 specimens
is particularly rich in material from northern Queensland and Western Australia. The
great bulk of the collection has remained practically untouched since the end of last
century. Its range and historic importance from its association with William Macleay
would entitle it to intensive treatment, but the single most pressing need is for
comprehensive treatment of the type specimens. Some original descriptions were so
inadequate that they raised questions of nomenclature which have remained vague for
nearly 70 years. Overseas herpetologists have been particularly hampered, and Loveridge
may be quoted in this connection. He says (1934, p. 248), “No stability of nomenclature
in Australian herpetology can be hoped for until some authority examines the types
(where still extant) and definitely settles the status of the many names so lavishly
proposed by those earlier Australian workers Macleay and De Vis. Longman has done
much work in this direction, but I would plead for one comprehensive study of every
species described. I have attempted to synonymize some sixteen of them in this present
paper and have revived several of their species which had been relegated to the synonymy
by other workers. Doubtless much remains to be done in both directions. The
descriptions, more particularly the earlier ones, of both these authors—Macleay and
De Vis—were so scanty and meagre that it is often difficult to decide with any confidence
what action to take regarding their disposition.” Earlier Boulenger (1904, p. 80) in
a note on Hinulia pardalis said inter alia: “the lizard which bears this name was so
imperfectly described by Macleay ... that when reviewing the Scincidae in 1887, I
could refer to it only in a footnote’. Although his descriptions are meagre, analysis
of the two forms dealt with in this paper vindicate Macleay’s good eye for species.
It is intended to catalogue all reptiles in the Macleay Museum. Two objects will be
kept in view. The first will be confined to giving adequate descriptions of all holotypes,
the second will be to discuss their geographical range and systematic relationships as
completely as possible. Specimens other than types will in general be listed briefly
with annotations. It is proposed to issue the catalogue in small sections for the sake
of convenience. This method will also enable descriptions of holotypes and other
essential matters to be distributed promptly.

CC

292 REPTILES IN THE MACLEAY MUSEUM,

This paper deals with two of Macleay’s species—Hinulia pardalis (here Spheno-
morphus pardalis pardalis) and Mocoa nigricaudis (here Sphenomorphus nigricaudis
nigricaudis). Previous authors have placed Lygosoma (Hinulia) elegantulum Peters
and Doria, and Mocoa nigricaudis in the synonymy of S. pardalis. It is shown that
S. pardalis is specifically distinct from both L. elegantulum and S. nigricaudis. Lygosoma
atromaculatum Garman is included in the synonymy of S. pardalis pardalis. It has
been found necessary to describe a subspecies of S. pardalis. Peters and Doria’s
L. elegantulum is specifically synonymous with S. nigricaudis, which has priority, but
retains subspecific rank. Standard descriptions are given of S. pardalis pardalis and
S. nigricaudis nigricaudis. Systematic points of difference between the four forms and
geographical distribution have been dealt with. All available references in literature
have been noted.

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Fig. 1.—Map showing localities where specimens of Sphenomorphus pardalis and
Sphenomorphus nigricaudis dealt with in this paper have been collected.

II. SPHENOMORPHUS PARDALIS PARDALIS (Macleay). Pl. xi, fig. 1.

Hinulia pardalis Macleay, 1877, p. 63; Boulenger, 1887, p. 209; ? Boulenger, 1904,
p. 80. Lygosoma atromaculatum Garman, 1901, p. 8; Zietz, 1920, p. 208. Lygosoma
pardalis Zietz, 1920, p. 208. Sphenomorphus pardalis Loveridge, 1934, p. 352. _Spheno-
morphus atromaculatus Loveridge, 1934, p. 353.

Holotype No. MR 21 in the Macleay Museum, labelled ‘‘Lygosoma (Hinulia) pardalis,
Macleay. Barrow Island, N.E. Aust.”

Macleay’s original description (1877, p. 63), which is especially valuable for its
colour notes made soon after capture of the holotype, is republished:

“Hinulia pardalis.

Moderately robust; tail acutely pointed, and about the length of the body; nasal
plates not contiguous and large, with the nostril small and in the middle; fronto-nasals

BY STEPHEN J. COPLAND. 293

not contiguous; interparietal small; other plates as in the last species;* ear opening
small and oval without denticulations; scales on the back in four series; colour, pale
olive on the back with numerous black spots, whitish on the sides, with very many
black spots and blotches, and yellowish white on the under surface.

One specimen, about 7 inches long, from Barrow Island, N.E. Australia.”

An extended redescription of the holotype follows. Although the head scales are
sometimes abnormal from fusion, cutting off of small scales or grooving leading to
incomplete separation, there is no doubt as to the arrangement and shape of scales.
This is because in no case are both members of a pair, or paired sides of the same
azygous scale, malformed. .

Description of Holotype.—Rostral moderately high, area visible from above equal
to about half that of the frontonasal, strongly concave sutures with nasals and slightly
‘shorter, much less concave ones with ist supralabials; the nearly straight junction
with the frontonasal is equal to about half the width of the frontal. Nasals large
with nostrils oval and sub-central; left scale a rough quadrilateral with the three
rounded convex sides against rostral, frontonasal (including small scale divided off from
it) and anterior loreal (and small scale divided off from it), fourth side straight and
horizontal against 1st supralabial; right nasal rounded, in contact with rostral, fronto-
nasal, 1st supralabial, also the abnormal 2nd supralabial with which part of the
anterior loreal has fused. No supranasals. Frontonasal large and malformed, about
two-thirds the area of the frontal with which it forms a straight suture approximately
two-fifths the width of the latter scale, also in contact with rostral, nasals and
prefrontals; on the left side it touches the anterior loreal and the small scale cut off
from it; on the right it is in contact with the abnormal 2nd supralabial and abnormal

SS

SS

SOS

aS

3

Figs. 2-—3.—Dorsal and lateral views of head of holotype of Sphenomorphus pardalis
pardalis. Actual head length, 12 mm., width, 10 mm.

posterior loreal; a small scale has been cut from it on the left anterolateral border
and another partly divided off from the left lateral angle. Prefrontals large, left
pentagonal in contact with frontonasal, frontal, 1st supraciliary, anterior and posterior
loreals and just touching 1st supraocular, right quadrilateral with posterior relationship,
the same as that of its fellow, but anteriorly in contact with only frontonasal and
posterior loreal. Frontal large, kite-shaped, half as long again as broad, long straight
postero-lateral sides against 1st and 2nd supraoculars, rounded posterior end against
frontoparietals, blunt anterior end against frontonasal and nearly straight anterolateral
sutures with prefrontals, just separated from 1st supraciliaries. Frontoparietals paired,
both large and malformed, right larger than left, long sinuous sutures with parietals, -
other contacts with frontal, 2nd, 3rd and 4th supraoculars and interparietal, left fronto-
parietal practically divided transversely and right fused to median posterior corner of
2nd supraocular. Interparietal kite-shaped, not much more than one-third the length
of the frontal (40 per cent.), rounded behind, pointed anteriorly and laterally, long

*“FHinulia atrocostata.’? Only two additional series of scales are dealt with: “supraorbital
region of 4 plates and a little elevated; the first five upper labials equal and nearly square,
the sixth and seventh larger and pentagonal”’.

294 REPTILES IN THE MACLEAY MUSEUM,

contacts with parietals and short ones with frontoparietals. Parietals are the largest
head shields, arranged in a V enclosing the interparietal and frontoparietals, each
anteriorly in contact with 4th supraocular, 9th supraciliary and 2nd postocular, long
lateral border against upper secondary temporal, right posteriorly in contact with two
nuchals, left with one nuchal and a scale about half the size of a nuchal interpolated
between 1st nuchal, parietal and upper secondary temporal; suture between parietals
slopes backwards towards the left. There are three pairs of large nuchals, the anterior
one on the left being largely separated from the parietal by the interpolated scale. Seven
supralabials, anterior four roughly oblong, slightly higher than long, upper margins
forming a fairly straight line with nasal, loreals, lower preocular and presubocular,
the fourth being capped by the last-named scale, posterior three rough pentagons, lower
margins straight and horizontal, anterior and posterior straight and vertical, the other
two sides meeting in a point dorsally, suboculars between 4th and 5th and 5th and 6th,
primary temporal between 6th and 7th, lower secondary temporal and two small post-
labials behind 7th; 5th under centre of eye; size in decreasing order 6, 7, 5, 4, 3, 2, 1.
Primary temporal squarish, subequal in area with 6th supralabial, posterodorsal border
against upper secondary temporal, anterodorsal against 2nd and 38rd postoculars, two
lower between 6th and 7th supralabials, anterior angle touches posterior subocular and
the posterior angle the tertiary temporal. The upper secondary temporals are very
large scales approaching the parietals in size, oblong in shape, sutures in order of
decreasing length with parietals, lower secondary temporal, 1st nuchal (on the left
the interpolated scale), primary temporal, tertiary temporal, 2nd postocular and a body
scale, the last three contacts being much shorter than the others. Lower secondary
temporal only slightly smaller than upper, more squat, higher than long, in contact with
the other three temporals, 7th supralabial and a postlabial. Tertiary temporal long
and band-like, long axis approximately vertical. It and the lower secondary temporal
are divided on the right side. Body scales begin behind the nuchals, interpolated scale
on left side, upper secondary temporal, tertiary temporal and postlabials. The two
loreals are practically normal on the left except for a small scale divided off the upper
margin of the anterior one; the small anterior loreal lies between nasal, frontonasal,
prefrontal, posterior loreal, and 1st and 2nd supralabials, only narrowly touching the
ist; the posterior, which is nearly twice the size of the anterior, has a very long
suture with the prefrontal and is also in contact with anterior loreal, 2nd and 3rd
supralabials, preoculars, and 1st supraciliary; on the right side the posterior borders of
the posterior loreal are normal but the anterior loreal has disappeared as an entity,
portions being fused to posterior loreal, 1st and 2nd supralabials, frontonasal and
prefrontal; fusion with the posterior loreal and frontonasal makes these two scales
continuous, but the others have well-developed sutures. Upper and lower palpebral
series abut against their respective preoculars. The upper preocular has also long
sutures with the lower preocular and ist supraciliary and short ones with posterior
loreal and upper accessory palpebral; it is divided on the left side. The upper preocular
as only half the size of the lower, which is in contact with upper preocular, posterior
loreal, 3rd and 4th supralabials and presubocular, which is wedged between lower
preocular, 4th supralabial and the most anterior subocular. The subocular chain is
ill-defined, consisting of about eight scales, three large between dorsally-directed points
of 4th and 5th and 5th and 6th supralabials and capping 6th supralabial; the other
scales are smaller and either join or cap these three. The’ postoculars are three small
scales, the 1st anterior and against 8th supraciliary and lower palpebral chain; 2nd
and 3rd lying in front of primary temporal; 2nd, which is at least twice as large as
each of the others, is also in contact with upper secondary temporal, parietal and 9th
supraciliary. Of the nine supraciliaries, the 1st is by far the largest, the Sth and 9th
ranking next; the 1st is a rough oblong between 1st supraocylar, prefrontal, posterior
loreal, upper preocular, upper accessory palpebral and 2nd supraciliary; the next six
scales are irregular; the 8th and 9th are subequal in size and lie between 4th supra-
ocular, postoculars and parietal; the upper palpebral series abuts against the Sth.
There are four large supraoculars, the 2nd being the largest, the frontal is in contact

BY STEPHEN J. COPLAND. 295

with the 1st and 2nd, the frontoparietal with the 2nd, 3rd and 4th, and parietal with
the 4th; the scales are somewhat irregular and the sutures between the anterior two
are abnormal or incomplete. The lower eyelid is scaly, its rim being formed by about
eight equidimensional or elongated scales of the lower palpebral series over about 11
scales, two or three in the centre being the largest. Infralabials five, all elongated.
Large mental in contact with two supralabials when mouth is closed, one infralabial
and the large postmental. Three pairs of large chin-shields, lst and 2nd pairs separated.
by an azygous, kite-shaped scale, 38rd pair by three scales. Ear oval, about half length
of eye, without denticulation, separated by two or three scales from 7th supralabial.

Scales in 24 rows at midbody, lateral and ventral scales subequal, two dorsal rows
markedly larger. Caudal scales much larger than those of body especially a dorsal
row. Two enlarged preanal scales, each twice the size of a ventral scale at midbody.
Seales from above vent to parietals, 57. Habitus compact, body only very slightly
depressed. Snout shorter than the space between eye and ear. The distance between
the end of the snout and the forelimb is contained 1:54 times in the distance between
axilla and groin. Tail, which is apparently intact, thick and tapering to a fine point;
slightly longer than head and body; broken twice near body since being preserved. Limbs:
short and reasonably powerful, well separated by about the length of the forearm and
manus when adpressed. The 3rd and 4th fingers and toes are by far the longest. Lamellar
formula for fingers, 4, ? (both 2nd fingers mutilated), 12, 11, 5. There are about 36
rounded tubercles on the palm, arranged in rough transverse rows. Lamellar formula
for toes, 6, 11, 18, 20, 11. There are about 45 rounded tubercles on the sole, the
proximal row being large and pointed.

Measurements of Holotype in mim.

Snout-vent an ee Abas (4 Head, length .. hs pat emDe,

Tail me ae Ae ia enter Head, width .. He eo

Snout-forelimb .. ike 56) eee) Forelimb, length ae so) al8s

Axilla-groin ee = ee 40) Hindlimb, length bas so PD
Width of body =e 56 abibes)

Colour: Ground colour of head, body and tail is light olive. There are no lines
or stripes, only black dots averaging about half a scale in size, very few filling an
entire scale. These black dots are scattered singly and in groups, where they are on
different scales. They are lightly sprinkled over the dorsal surface of the head, being
inclined to margin scales such as the supraoculars. From the head to above the
forelimbs there is the vaguest suggestion of four irregular longitudinal lines; about a
third of this area is black. The body is dorsally about half black, two or three large
black spots being commonly aggregated. Spots on the tail have about the same
intensity, but here there are two or more longitudinally disposed dots on each scale.
The side of the head is heavily barred vertically. The dorsolateral areas are practically
clear from the head to above the vent. Below these clear spaces there is a fairly dense
aggregation of large black spots along the flanks. Half way down the sides the spots
become separate and fade out ventro-laterally in a series of small dots. Neither a
clear space nor a marked aggregation of dots occurs on the tail, which is marked much
the same laterally and dorsally. The underside from the throat, which is sprinkled with
black, to the vent shows no trace of black and is light olive much as the back. The
tail below is fairly clear but dotted at intervals.

Garman’s type description of his Lygosoma atromaculatum (1901, p. 8), which is
here placed in the synonymy of VS. p. pardalis, is given for purposes of comparison.

“Horm similar to that of L. isolepis Boul. Body elongate, slightly depressed; limbs
short, rather weak, not meeting by the length of the arm when adpressed; feet penta-
dactyl; tail one and one-half times as long as head and body, thick, round, tapering
regularly. Distance from snout to fore-leg contained one and one-half times in the
distance from axilla to vent. Snout short, shorter than the space between the eye and
the ear. Lower eyelid scaly, transparent. Rostral hexagonal, wider than high, truncate,

~

296 REPTILES IN THE MACLEAY MUSEUM,

in contact with the frontonasal. Nostril pierced in a single nasal; no supranasal.
Nasal quadrangular, in contact with the first labial; postnasal in contact with the
second labial; loreal in contact with second and third labials. Labials seven, fifth and
sixth below the eye. Frontal one and one-half times as long as wide, broadly in
contact with the frontonasal and with the anterior two pairs of supraoculars; prefrontals
small; frontonasal broader than long, octagonal; frontoparietals moderate, little larger
than the interparietal; parietals large, meeting behind the interparietal. Three to four
pairs of nuchals, twice as wide as the shields behind them. A large shield and a much
smaller one at the outer side of each parietal. Four supraorbitals, second widest. Five
or six broad shields between the eye and the ear. Seven or eight supraciliaries. Mental
shield large, broad, in contact with two labials and a submental. Anterior submental
broader than long, in contact with five shields, followed on each side by four broad
submentals, the anterior pair of which meet on the median line, the second pair are
separated by a single small scale, and a third pair are separated by three scales. Har
opening subelliptical, oblique, little smaller than the eye, with several hardly noticeable
lobules on the anterior border. Scales smooth, in twenty-four rows around the body,
dorsals larger and laterals little smaller than the ventrals; a pair of enlarged preanals.
Below the tail the scales are somewhat larger than those on the upper surfaces. Rostral,
nasals, first labial and mentals have in most cases the appearance of being thicker than
the other head scales or of having retained the slough. Digits weak, slightly compressed;
subdigital lamellae forming a low keel, nineteen under the fourth toe.

Bronzed olive on the back, more or less lightly sprinkled with black spots which
become more numerous toward and on the tail and on the limbs. Belly and lower side
of tail uniform whitish. Scales of sides and lower surfaces of head and throat with
black spots, those of labials and submentals most intense. Entire flanks closely spotted
with small black spots; in cases the spots of sides and back become longitudinal
streaks. On some individuals the back is more thickly covered with spots which are
smaller forward and on the back of the head, and each labial bears a white vertical

bar in the middle, the black spots being situated on the sutures and covering a portion
of each scale.

Differs from L. isolepis Boul. and L. elegantulum Pet. & Dor. in the smaller
number of scales.

Barrier Reef, Australia; G.B.R. Exp.: Queensland; Mr. Olive.” :

Of the approximately 86 characters dealt with in Garman’s description of
S. atromaculatus, 76 agree exactly with those of S. p. pardalis. The remaining 10
represent at most slight differences and are discussed hereunder.

Description of Cotypes of S. atromaculatus. ; Holotype of S. pardalis.
1. “tail one and one-half times as long as Tail is only slightly longer than head and
head and body”. body. Tail may have been damaged.

This character is almost always very
variable in the genus.

2. “lower eyelid ..., transparent’’. The central enlarged scales of the lower
eyelid are at least translucent.
3. ‘“frontoparietals . ., little larger than Each frontoparietal twice the size of the
the interparietal’’. interparietal.
4. “seven or eight supraciliaries’’. Nine supraciliaries, but only three large,

the remainder small. A variable character
in the genus.
5. “anterior submental .. ., followed on Three pairs of chin-shields follow the post-
each side by four broad submentals”’. mental, not four. Garman described three
pairs of chin-shields, calling them sub-
mentals, exactly as they occur in the
type of pardalis. His “four” was prob-
ably a lapsus for three unless he included
the unpaired postmental, which he called
anterior submental, as the fourth sub-
mental. 3
6. “ear-opening .. ., oblique’’. Bar opening vertical.

BY STEPHEN

Description of Cotypes of S. atromaculatus.

J. COPLAND.

Holotype of S. pardalis.

7. “ear-opening . . ., with several hardly
noticeable lobules on the anterior
border’’.

8. “rostral, nasals, first labial and mentals
have in most cases the appearance
of being thicker than the other head
seales or of having retained the
slough”’.

No. trace of lobules, but when scales dry
slightly during examination the borders
give the impression that lobules are
present.

There is little or no difference in the
appearance of these scales. The presence
of sloughing would probably depend on
the season at which specimens were col-
lected.

297

, nineteen under Twenty in pardalis. A form may be charac-
terized by a narrow range in the number
of lamellae, but there is never an exact,
invariable number of lamellae.

The underside is uniform, but now greenish,
probably partly due to preservation.
Macleay, who examined the specimen
shortly after it had been collected, said
“yellowish white on the under surface’’.

9. “subdigital lamellae .
the fourth toe’’.

10. “belly and lower side of tail uniform
whitish”’.

Of the points of difference only numbers 3 and 6 appear to be of any significance,
and the last is probably only an individual difference. The relationships of the fronto-
parietals to the interparietal may be expected to fluctuate to some extent even in
neighbouring populations. Unless constant and prominent, they would not justify
subspecific differentiation, especially when swamped by the correspondence of 85 other
characters out of 86 considered. Supposing the original mainland stock to have been
practically homogeneous when the coastal islands were cut off by submergence of
low-lying coastal areas, say 10,000 years ago, slight differences must have developed
between isolated populations. It is indeed remarkable that under the circumstances,
specimens collected on separated islands should show such little variation. The over-
whelming weight of evidence is that Garman’s Lygosoma atromaculatum is conspecific
with Macleay’s Hinulia pardalis.

Locality Records of S. p. pardalis.

In this and the following lists the original reference is given when possible followed by the
collector’s name (when available) in brackets and date of collection.

Barrier Reef: Garman (1901, p. 8) 2 cotypes of Lygosoma atromaculatum (M.C.Z.
6475) (A. G. Mayer) 1896.

Barrow Island, N.E. Australia: Macleay (1877, p. 63) holotype of Hinulia pardalis.

Coen: Loveridge (1934, p. 353) under Sphenomorphus atromaculatus, 23 specimens
(M.C.Z. 35412-34) (BP. J. Darlington) 1932.

Cooktown: Loveridge (1934, p. 353) “3 cotypes” of Lygosoma atromaculatum (M.C.Z.
6478) (EH. A. Olive) 1896.

‘Except for Zietz (1920, p. 208), who lists it from Queensland and the Barrier
Reef, I have been unable to find other references to S. atromaculatus beyond its
treatment in Loveridge’s 1934 paper, which is invaluable to all Australian herpetologists.
Loveridge, who had 28 specimens under examination, two cotypes (M.C.Z. 6475) from
the Barrier Reef, collected by A. G. Mayer; three other cotypes (M.C.Z. 6478) from
Cooktown, collected by E. A. Olive; and 23 specimens (M.C.Z. 35412-34) collected at
Coen in 1932 by P. J. Darlington, says (1934, p. 353): “Midbody scale-rows 24 (every
individual counted); frontonasal forming sutures with the rostral and frontal; scales
bordering the parietals posteriorly on right and left sides respectively 2 + 2 (in 13
skinks), 2 + 3 (in 10), 3 + 2 (in 2), 3 +3 (in 3); adpressed limbs do not nearly meet.
Largest skink (No. 35412) measures 142 (63 + 79) mm. This skink is very similar to
S. pardalis and must be extremely difficult to distinguish without comparative material.
It is well named, for the aggregation of black markings along the flanks are, perhaps,
its most distinguishing feature. The unusual constancy in a skink of a fixed number
of midbody scale-rows is interesting; in this connection it may be noted that a single
pardalis was also taken at Coen but was eliminated by its larger size and absence of
characteristic atromaculatus markings quite apart from its 26 midbody scale-rows. It
will also be noted that there is a single skink with 24 midbody scale-rows referred to

298 REPTILES IN THE MACLEAY MUSEUM,

pardalis. Here again I have no doubts as to its correct relegation to that species.
Possibly atromaculatus has but recently been subject to speciation.”

III. SPHENOMORPHUS PARDALIS ERRO, nh. Subsp. PI. xi, fig. 2.

Diagnosis: Closely allied to Sphenomorphus pardalis pardalis but separated by the
bright reddish-brown ground colour (instead of pale greenish-olive), and to a lesser
degree by slimmer build, greater length of interparietal, and other scale characters dealt
with in the following notes and table.

Holotype. No. R 6352 in the Australian Museum, Sydney, from the old collection,
locality unknown. The single paratype No. R 6373 appears to have been collected at
the same unidentified locality.

In both specimens the scales are clear cut and show no sign of the tendency towards
malformation so evident in the holotype of S. p. pardalis, where it is evidently due to
some genetic instability.

R 6352 and R 6373 resemble each other in scale characters almost as if they were
twins and there is the strongest resemblance to S. p. pardalis. Holotype and paratype
have been treated together in the following notes, and where S. p. erro differs from
S. p. pardalis the character of the latter subspecies in question is included in brackets.
The two subspecies are identical in all points not mentioned. This is also the case
where it has been thought advisable to mention other characters.

Table of Measurements in mm. and Other Characters of Holotype and Paratype of 8. p. erro, with Those
of S. p. pardalis for Comparison.

R 6352 R 6373 MR 21

Snout-vent sie ve ae ae an 59 56 72
Tail bid sy as Be es Pe 83 +* 58+ 77
Snout-forelimb .. as at Af Bic 21 19 26
Axilla-groin ee re oe uA ee 35 31 40
Head, length .. oe ®. a we 10 9-5 12
Head, width .. ac Ne Bic aa 9 8 10
Forelimb, length Se a3 ot 6 11:5 11 13
Hindlimb, length to ee ate a 18 17 20
Width of body = ae bs Be 10 8:5 11-5
Axilla-groin/Snout-forelimb .. ie Ae 1:67 1°63 1:54
Tail/Body as ae Ms ae 1:41+ — 1:07
Suture of frontonasal with frontal to width

of frontal ae Bs rhe ae 1/3 1/3 * 2/5
Length of interparietal to length of frontal 55% 60% 40%
Nuchals, left and right sides. . sii ae 3-3 3-3 3-3
Seales touching postero-lateral border of

parietals,; left and right Rae 2-4 3-4 3-3
Supraciliaries .. 5 ae ib. is 9 9 9
Infralabials ao we oe 5 3 5 5 5
Infralabials in contact with postmental on

each side .. aie te se Le 1 1 1
Number of scales from above vent to parietals 60 54 57
Lamellae beneath fingers ae ae a) 4565, LODO, 4, 7, 10, 10, ? greg ab TTS 15)
Lamellae beneath toes. . ue Ee ae toh dO ae aieyae GGL (hs aby tse al) Gaal, Te AO), La
Midbody scale rows .. mi Bey Ls 24 24 24

* Practically complete, extreme tip bifid.
7 Including upper secondary temporal, interpolated scales and nuchals.
£15 on short right toe.

In both R 6352 and R 6373 the width of the suture of the rostral with the frontonasal
equals ; the width of the frontal (4), and the area of the rostral visible from above
is equal to at least 2/3 that of the frontonasal (about 3). The nasal is in contact with
9/10 of the upper margin of the 1st supralabial. Prefrontals just touch the 1st supra-
oculars. The lower border of the posterior loreal is against the 2nd and 3rd supralabials.
R 6373 has the three postoculars very small. The primary temporal in R 6352 is slightly
smaller than the 6th supralabial, in R 6373 it is little more than half its size (subequal).
The two specimens are identical in colour and markings. The bright reddish-brown
ground colour resembling that of Sphenomorphus nigricaudis elegantulus is in striking

BY STEPHEN J. COPLAND. 299

contrast to the pale greenish-olive of S. p. pardalis. The dorsum is more sparingly
sprinkled with black than in S. p. pardalis. Black markings are aggregated along the
flanks but again are less concentrated than in S. p. pardalis. There is a greater
tendency towards vertical barring of the sides between the ear and the forelimb than
in the nominate subspecies. This is more pronounced in R6373. The whole ventral
surface, except for short bars along the sutures between the infralabials, is of much
lighter brown than the dorsum. Although the ground colour is that of S. n. elegantulus,
the pattern is much like that of S. p. pardalis. A topotype of S. n. elegantulus (R 9599)
lacks the dense concentration of markings along the sides, is lighter ventrally, and more
heavily spotted and cross-barred across the dorsum anteriorly from the head to the
length of the forelimb behind the shoulders.

I was reluctant to describe this subspecies because of the unknown origin of the
specimens, but there was no alternative. S. p. erro is strikingly different in colour
to S. p. pardalis, yet there is no doubt that it belongs to the same species. In handling
specimens of the two forms, one is impressed by the close similarity in scalation
except for comparatively minor differences already mentioned. S. p. erro is sharply
separated from S. n. nigricaudis and S. n. elegantulus by the lower number of midbody
scale rows and the fact that the postlabial is in contact with only one infralabial instead
of two (see Figs. 6 and 7). The fact that the only two specimens in the Australian
Museum agree in the number of midbody scale rows and the postlabial contact indicates
that they are representatives of a population normally possessing these characters,
and immensely lessens the possibility that they are exceptional varieties of a population
possessing other characters. I am confident that this subspecies will be rediscovered—
probably in north Queensland. Its present homeless condition has been suggested by
its subspecific name—the Latin noun for wanderer—placed in apposition.

IV. SPHENOMORPHUS NIGRICAUDIS NIGRICAUDIS (Macleay). Pl. xi, fig. 3.

Mocoa nigricaudis Macleay, 1877, p. 63. Homolepida crassicauda Barbour (non
Duméril), 1914, p. 204.

Cotypes. Nos. MR376-380 in the Macleay Museum labelled ‘“Lygosoma (Mocoa)
nigricaudis Macleay. Darnley Island’’.

MR 378 is here designated the lectotype, the remaining four specimens becoming
paratypes.

It appears to the author that it is as essential to choose a single specimen in
redescription where cotypes are involved as to designate a holotype in an original
description. Davis and Lee (1944, p. 18) put the» matter clearly: ‘Where an early
worker used a series of specimens as the basis of his original description without
designating one as type (a practice which has ied to considerable confusion and for the
perpetuation of which there can be no excuse), these may be regarded as syntypes or
cotypes. In cases such as the preceding the onus is on the subsequent reviewer to
select one specimen, the lectotype, to act for the future as the true single type.”

Dunn (1934, p. 171) says: “If the cotypes of a species turn out to represent two
different forms some revisionary restriction of the cotype series is necessary. In no
other situation is discrimination between cotypes necessary, nor has it any legal
sanction.” The difficulty is that reviewers cannot know what future refinements there
may be in systematic technique or requirements. All possibility of confusion is avoided
when a single specimen, which obviously can at any time belong to only one form, is
chosen. Mayr (1942, p. 15) is definite on this point. Where the author of a species
has used cotypes and distributed some of them, as in the special case given by Dunn,
a worker would be guilty of unethical conduct if he selected a lectotype without
consulting the author if available. This position does not apply here where all cotypes
have been kept together.

Darnley (or Erub) Island (9° 30’ S, 148° 42’ E) is a high, wooded, fertile, volcanic
island in Torres Strait, near the northern tip of the Great Barrier Reef.

Following is Macleay’s original description (1877, p. 63), mainly important, as in
the case of S. p. pardalis, for the early observations on colour.

300 REPTILES IN THE MACLEAY MUSEUM,

“Mocoa nigricaudis.

Har opening nearly round, without denticulation; rostral plate rounded above;
nasals not contiguous; fronto-nasals nearly contiguous; supraorbitals four; fronto-
parietals two, of the same size as the interparietal; scales of the back in about six
series; colour, reddish brown above with a few scattered black spots—these become
very dense towards the apex of the tail, giving it a black appearance; the under
surface is greyish yellow; the labial plates are spotted with black; body moderately
robust; limbs weak; toes of fore feet short; total length, 9 inches.

Hab. Darnley Island.”

The scheme of the following redescription of the lectotype is identical with that
used earlier for S. p. pardalis.

Description of Lectotype—Rostral moderately high, area visible from above equal
to nearly half that of the frontonasal, strongly concave sutures with nasals and much
less concave, nearly straight, ones, half the length of those with the nasals, with ist
supralabials, the convex suture with the frontonasal is equal to nearly half the width
of the frontal. Nasals large, with nostrils oval and slightly behind the mid-line, convex
sutures with rostral, frontonasal, anterior loreal and ist supralabial, in contact with

ZA
ZA
4

LES

hy

ra
ws
(2

shy

Fig. 4.

B
Sims

Figs. 4-5.—Head of lectotype of Sphenomorphus nigricaudis nigricaudis. Actual head
length, 13 mm.. width, 11 mm. 4. Dorsal view. 5. Lateral view.

BY STEPHEN J. COPLAND. 301

about two-thirds the upper margin of the latter scale. Frontonasal large, at least
two-thirds the area of the frontal with which it forms a nearly straight suture only
one-fifth the width of the latter scale, also in contact with rostral, nasals, prefrontals
and anterior loreals. Prefrontals large, in contact with frontonasal, frontal, Ist supra-
ciliary, anterior and posterior loreals, just touching 1st supraoculars. Frontal large,
kite-shaped, half as long again as broad, long straight postero-lateral sides against 1st
and 2nd supraoculars, rounded posterior end between frontoparietals, blunt anterior end
against frontonasal, and slightly concave antero-lateral sutures with prefrontals, just
separated from 1st supraciliaries. Frontoparietals paired, large, and symmetrical except
that the suture between them slants backwards towards the. left, strongly indented
against frontal, short transverse suture with 2nd supraocular, long slightly concave
antero-lateral border against 3rd and 4th supraoculars, sinuous contact with parietal
and sinuous slightly shorter one with interparietal. Interparietal large, kite-shaped,
considerably more than half the length of the frontal (60 per cent.), rounded behind,
pointed anteriorly and laterally, long, straight contacts with parietals, and little shorter,
sinuous, mainly concave ones with frontoparietals. Parietals are the largest head
shields, rather irregular in shape, twice as long as wide, each anteriorly in contact
with frontoparietal, 4th supraocular, 10th supraciliary and 2nd postocular, the contact
with the 4th supraocular being very short and that with the 2nd postocular little
longer; separated by interparietal except for a short suture one-quarter the length of
the interparietal; left postero-lateral suture against upper secondary temporal, anterior
nuchal and a scale interpolated between these two, right border similar but in contact
with two interpolated scales and just touching a second nuchal. There are five distinct
nuchals on the left side, the scales are irregular on the right, with only two well-marked
nuchals, which correspond to the first and third on the left. Seven supralabials, Ist,
2nd and 3rd with concave posterior sutures, other four convex, all higher than long,
1st with dorsal peak between nasal and anterior loreal, 2nd straight against the two
loreals, 3rd straight and capped by lower preocular and presubocular, 4th to 7th pointed
dorsally, 4th and 5th between suboculars, 6th between suboculars and primary temporal,
7th between primary temporal and lower secondary temporal, two postlabials behind
7th; 5th under centre of eye; size in decreasing order, left side, 6, 7, 4, 3, 1, 5, 2, right
side, 6, 7, 5, 3, 4,1, 2. Primary temporal squarish, considerably smaller than 6th supra-
labial, two lower sides between 6th and 7th supralabials, antero-dorsal against 2nd and
3rd postoculars, and postero-dorsal against upper secondary temporal, posterior angle
touches tertiary temporal and anterior the last of the suboculars. The upper secondary
temporals are large scales, each about half the size of a parietal, roughly oblong, long
sutures with parietal, lower secondary temporal, shorter with body scales (three
contacts on right, two on left), primary and tertiary temporals, and very short with 2nd
postocular. Lower secondary temporal about equal in size to the upper, nearly square,
long sutures with upper secondary temporal, 7th supralabial and the tall, upright,
band-like tertiary temporal, shorter lower border against upper postlabial and in front
narrowly in contact with primary temporal. Body scales begin behind the nuchals,
interpolated scales, upper secondary temporals and postlabials. The two loreals are
large and about equal in size, anterior roughly lens-shaped with one long, smoothly
convex border against frontonasal, prefrontal and posterior loreal, other border more
irregular and in three steps against nasal and ist and 2nd supralabials; posterior
loreal quadrilateral, posterior side against upper and lower preoculars, upper against
1st supraciliary and prefrontal, other two sides against anterior loreal and 2nd supra-
labial. Upper and lower palpebral series abut against the upper preocular, which is
also in contact with lst supraciliary, posterior loreal, lower preocular and upper
accessory palpebral, which is wedged between the palpebral chain and lst and 2nd
supraciliaries. The lower preocular is twice the size of the upper and lies between it,
the posterior loreal, 3rd supralabial and presubocular; its posterior angle touches the
lower palpebral chain. The presubocular is wedged between the lower preocular, 3rd
supralabial and the most anterior subocular. The suboculars are reasonably prominent
and consist of five or six scales, all in contact capping the 4th, 5th and 6th supralabials

302 REPTILES IN THE MACLEAY MUSEUM,

or sending down points between them; there are other smaller scales above the main
row. The postoculars are three small scales, the Ist anterior and against the 9th
supraciliary and a small, rounded scale lying against the end of the lower palpebral
chain, the 2nd and 8rd lying in front of the primary temporal; the 2nd, which is
larger than the others and subequal in size to the 10th supraciliary, is in contact with
the primary temporal, upper secondary temporal, parietal, 10th supraciliary and ist
postocular. Of the 10 supraciliaries, the 1st is by far the largest, the 10th and 9th
ranking next; the intermediate six are often little more than nodules; the ist is
narrowly separated from the frontal and lies between the 1st supraocular, prefrontal,
posterior loreal, upper preocular, upper accessory palpebral and 2nd supraciliary; the
9th and 10th lie between the 4th supraocular, parietal, and lst and 2nd postoculars;
the upper palpebral series ends against the 9th. There are four large supraoculars, the
2nd the largest; the frontal is in contact with the 1st and 2nd, the frontoparietal with
the 2nd, 3rd and 4th, and the parietal is narrowly in contact with the 4th. The lower
eyelid is scaly, its rim being formed of about ten scales interlocking with those below.
Infralabials seven, all much the same in depth but more elongated posteriorly. Large
mental in contact with just half the lower border of the 1st supralabial when the mouth
is closed. Broad postmental in contact with two infralabials on each side, the anterior
chin-shields and mental, making seven scales in all (see Fig. 6). Three large pairs of
chin-shields, 1st in contact, 2nd separated by a small triangular scale, and 3rd pair by
three small scales.

Ear opening oval, greater diameter equal to two-thirds the length of the eye, without
denticulation, separated by three scales from 7th supralabial.

Scales in 28 rows at midbody, dorsal scales considerably larger than ventral, lateral
seales much smaller again. Caudal scales dorsally much the same size as those on
the body until quite near the tip of the tail; laterally much larger; ventrally very
large, especially the median row of transverse scales, which begins about the length
of the hindlimb behind the vent. Two large preanals, each twice the size of a ventral
scale at midbody. Scales from above vent to parietals, 58. Habitus compact, body
slightly depressed. Snout shorter than the distance between eye and ear. The distance
between the end of the snout and the forelimb is contained 1:48 times in the distance
between axilla and groin. Tail thick, tapering to a fine point, apparently intact,
slightly longer than body. Limbs short, reasonably powerful, separated when adpressed
by about the length of the manus. Length of fingers in decreasing order 4 = 3, 2, 5, 1;
of toes, 4, 3, 5, 2, 1. Lamellar formula for fingers, 5, 8, 10, 12, 6. There are about
40 small, rounded at base, pointed tubercles on the palm and four or five large ones
edge the wrist. Lamellar formula for toes, 5, 8, 14, 17, 10. There are about 50 rounded
tubercles on the sole, the proximal ones larger and arranged in two or three rows.

Measurements of the lectotype are given with those of the paratypes.

Colour: Dorsal surface of head and body uniform brown, about 20 small blackish
spots occur on the back in the area 10 scales before and 15 behind the forelimbs. The
spots average about % the area of a scale and are nearly uniform in size. The underside
of the head, body and tail (except for the terminal fifth) is uniform light brown,
now little different to the dorsal ground colour. The blackish spots on the supra- and
intralabials, suboculars, preoculars, supraciliaries and temporals do not exceed 25 in
number. There are scattered small black spots on the sides of the body, mainly on
the dorso-lateral region before the forelimbs, but descending laterally behind them.
The posterior half of the area between axilla and groin is clear. There are about 25
spots on each side. A very few spots occur on the dorsum of the proximal fifth of the
tail, then the next two-fifths is sprinkled. The terminal two-fifths is black dorsally
and laterally. Only odd spots occur laterally on the proximal fifth of the tail. The
dorsal surfaces of the limbs are spotted with black.

Variation in Paratypes.—The series of five (lectotype and four paratypes) is very
uniform and there is even a strong family likeness, but there is some variation. The
length of the interparietal in MR376 agrees with that of MR37S in being equal to
60 per cent. of the length of the frontal; in the other three specimens it is 70 per cent.

BY STEPHEN J. COPLAND. 303

The interparietal is subequal in area to a frontoparietal in MR378 and MR 379; as
long but narrower in the other three cases. In MR 376 the suture between the parietals
slopes backwards towards the right instead of left as normally. The nuchals have a
strong tendency to be more numerous and regular on the left side. The following
table summarizes the relations to the scales following the parietals, and also the
arrangement of nuchals.

Seales touching Postero-

Nuchals. lateral Border.*

Left. Right. Left. Right.
MR 376 5 27 4 3
MR 377 4 3 3 4
MR 378 5 2t 3 5
MR 379 4 3 3 4
MR 380 4 4 3 4

* Including upper secondary temporal, interpolated scales and nuchals.
7 2nd corresponding to 4th on left.
{2nd corresponding to 3rd on left.
Although the 6th and 7th are always the largest two supralabials and the Ist and
2nd generally the smallest, these scales vary somewhat irregularly as indicated in
the following table. which gives sizes in decreasing order.

MR 376 MR 377 MR 378 MR 379 MR 380
Left Side 5% 6735412 6735421 6743152 6735421 7635412
Right Side ee 6735241 6735412 6753412 6734512 7631254

The 6th supralabial narrowly touches the primary temporal in three cases. On the
right side of MR 376, as in MR 378, the contact is long, but on the left the two scales
are separated by the large last subocular which touches the 7th supralabial. MR 377
and MR 380 have the lower border of the posterior loreal equally in contact with the
2nd and 3rd supralabials on the right, but not on the left side, where it touches only
the 2nd. MR379 has the right posterior loreal narrowly in contact with the 3rd
supralabial as well as a long suture with the second. The anterior loreal is partly
divided on the left side in MR376. In MR376 the lower preocular is in contact with
the 2nd supralabial as well as the 3rd. The right presubocular in MR 379 touches the
4th as well as the 3rd supralabial. MR377 and MR380 have their suboculars much
more prominent than in MR378. MR376 and MR 377 have nine supraciliaries: these
scales are most strongly developed in MR380. There are six infralabials in MR 376
and MR379. There may be as many as six scales separating the ear from the 7th
supralabial, but in this case the posterior three are very small. All specimens have
28 midbody scale rows. Number of scales from above vent to parietals is 56 (twice),

Measurements of Lectotype and Paratypes of Sphenomorphus nigricaudis nigricaudis in mm.
+ = eS

MR 376 MR 377 MR 378 MR 379 MR 380

Snout-vent .. 75 78 76 (Al 78
Tail : 78 + 28+ 90 84+ 96
Snout-forelimb a; ae Bt ae 26 27 27 26 25
Axilla-groin .. as os NAS B oe 40 44 40 39 42
Head, length Ste te 7: ps ss 112-5 13 13 12 14
Head, width = or oa sh Li 10-5 11 11 ; 10 T1355)
Limbs, length forelimb .. Sie A at 13 14 13 Zid 14
Limbs, length hindlimb .. ae we ios 21 23 24 20 24
Width of body Bid a) ne th. a 13 13 14 12 14
Axilla-groin 3% “i as oe ne = = = = are
Snout-forelimb ts ae ae at ae 1-54 1:63 1-48 1-50 1-68
Tail/Snout-vent .. 25 a ae oe + — 1:18 —- 1323

304 REPTILES IN THE MACLEAY MUSEUM,

57 and 58. MR377 has had the tail broken off about 15 mm. behind the vent and the
regenerated portion, a stub about 10 mm. long, is only about one-third the diameter of
the tail it replaces. MR880 has had its tail snapped off but preserved. Number of
lamellae under the 4th toe is MR376, 20; MR8377, feet on both sides mutilated;
MR 379, 18; MR 380 with 17 agrees with the lectotype. Accidents, probably attacks by
birds, have caused the loss of, or damage to, three of the five tails in the series, the
loss of the left manus and the ends of two toes in MR 377 and four left toes of MR 376.

Variation in colour is not very striking. All specimens have the uniform reddish-
brown dorsal ground colour. MR376 and MR 377 are more distinctly spotted anterior
to the forelimbs than MR 378. In MR 376 and MR 377 the black dots give the suggestion
of six or seven cross bars, and these are continued laterally. MR 379 is almost undotted
dorsally but has four distinct vertical bars between the ear and forelimb on each side.
The same lizard has ill-defined narrow lines along the sides of the body. All specimens
have the labials and adjacent scales somewhat spotted. Except in MR 377, where it is
missing, the distal third to two-thirds of the tail, not the base as mentioned by
Loveridge (1934, p. 353), is blackish with a dense aggregation of pigment. This is
most conspicuous in MR 380, which otherwise almost duplicates MR 378 in colour and
markings.

Locality Records of S. n. nigricaudis.

Darnley Island: Macleay (1877, p. 68) type locality of Mocoa nigricaudis; and Barbour
(1914, p. 204) under Homolepida crassicauda.
South-East Cape, New Guinea: Nos. A 5654 and A 5682 in the Australian Museum
(“through Rev. Macfarlane from teachers and natives, 1879’).
As far as I can ascertain there is no South-East Cape in New Guinea, but it is almost
certain that the specimens came from the neighbourhood of East Cape, where Macfarlane is
known to have collected extensively.

Table of Measurements in mm. and Other Characters of 3S. n. nigricaudis in the Australian Museum.

A 5654 A 5682

Snout-vent Ay Ag as ae ie ae a ae 80 73
Tail ts ue ys ane a8 a ie A at 58+ 105
Snout-forelimb .. ae ve sae a a ne a 27 24
Axilla-groin ne Sh 5 aa nfs one Se Re 42 40
Head, length Ls ou Ss ate a be oe Ce 13 13
Head, width Hes A a 6 Ae ae Aa 2M 11-5 11
Forelimb, length .. ae one as eye $e is ze 14 15
Hindlimb, length .. Me te se oO a5 a a 23 23
Width of body .. a ys an ts aa ES 13-5 13
Axilla-groin/Snout- forelimb Be, it bo aie i Se 1-56 1-67
Tail/Body .. , ae — 1-44
Width of suture of Rare! with fontonacal to width of frontal oe 1/3 1/3
Upper margin of 1st supralabial in contact with nasal _ .. ais 2/3 2/3
Width of suture ‘of frontonasal with frontal to width of frontal a 1/5 1/5
Length of interparietal to length of frontal .. ae ae Ae: 65% 80%
Number of nuchals, left and right sides ag 5-4 4-4
Number of scales touching postero-lateral sides of aamietell left

and right Ba es ae see Be ae Hie Sa 2-3 2-3
Supraciliaries ae yc ai es ie ae fs ak 10 10
Infralabials. . ve ae 56 Ate Ae a a 6 6
Lamellae beneath 4th foe, ay oe sts on 3 te 20 19
Midbody scale rows ey eit au ee ye ne 28 28 is

* Including upper secondary temporal, interpolated scales and nuchals.

The lower border of the posterior loreal is in contact with the 2nd supralabial only.
Habitus of these lizards and the Darnley Island holotype and paratypes is very close.
In colour they closely resemble the types except that they are considerably more heavily
spotted, especially A 5682. This specimen has the distal third of the tail dark though
not as conspicuously so as MR378. The distal third of the tail in A 5654 has been
damaged.

BY STEPHEN J. COPLAND. 305

Differences between S. nigricaudis nigricaudis and S. pardalis pardalis——These
closely allied forms are yet separated in so many characters that a list of these can
only help treatment and comparison.

The posterior border of the rostral makes sweeping curves, first concave with the
nasals and then convex with the frontonasal, as it passes between the supralabials in
nigricaudis. In pardalis the contact with the frontonasal is nearly straight.

Contact of the frontonasal with the anterior loreal is much longer and has a quite
different outline in nigricaudis, the length of the contact of the frontonasal with the
anterior loreal being twice that of the prefrontal with the same scale. In pardalis
these proportions are reversed.

The interparietal is much better developed in nigricaudis than in pardalis, its
length compared with the length of the frontal being 60 per cent. (in the lectotype
and one paratype, 70 per cent. in the other three paratypes) against 40 per cent.

The length of the suture between the parietals is equal to 25 per cent. of the
length of the interparietal in nigricaudis; 60 per cent. in pardalis.

The primary temporal in nigricaudis is much smaller than in pardalis.

The upper secondary temporal is relatively smaller in nigricaudis.

The anterior loreal in pardalis is much smaller than the posterior; they are of
equal size in nigricaudis.

In nigricaudis the posterior loreal is not in contact with the 3rd supralabial as
it is in pardalis.

The lower preocular in nigricaudis is widely separated from the 4th supralabial;
it is in contact in pardalis.

Nigricaudis has seven infralabials; pardalis five. Their outlines are also distinctive.

In nigricaudis the mental is in contact with half the margin of the 1st supralabial
when the mouth is closed; in pardalis it is in contact with all the 1st and portion of
the 2nd.

In nigricaudis the postmental is in contact with seven shields (including two
infralabials on each side); in pardalis it is in contact with five shields (one infralabial
on each side).

Nigricaudis has 28 scale rows at midbody; pardalis 24.

Nigricaudis is the more sturdily built and compact lizard.

The limbs are stronger and better developed in nigricaudis than in pardalis,
especially the hindlimbs which are longer and markedly more robust.

Fingers and toes in pardalis are more slender and more compressed than in
nigricaudis.

Differences in colour are very striking. The following three are perhaps the most
important: (a) the ground colour of nigricaudis is reddish-brown, that of pardalis pale
olive; (0) the back and sides of nigricaudis have only a few scattered black spots, the
general impression of uniform colouration being scarcely affected except for the distal
third of the tail, while pardalis is heavily spotted and blotched with black dorsally and
laterally on all parts of the head, body and tail; (c) pardalis is without the characteristic
aggregation of black on the tail, which also extends to the underside, of nigricaudis.

V. SPHENOMORPHUS NIGRICAUDIS ELEGANTULUS (Peters and Doria). Pl. xi, fig. 4.

? Hinulia striatula Giinther (non Steindachner) part, 1875, p. 11; 1877, p. 413.
Lygosoma (Hinulia) elegantulum Peters and Doria, 1878, p. 344; Zietz, 1920, p. 208;
Kinghorn, 1931, p. 89. Lygosoma elegantulum Boulenger, 1887, p. 235; ? Oudemans,
1894, p. 140; Boulenger, 1895, p. 29; Broom, 1898, p. 643; de Rooij, 1915, p. 182.
? Hinulia pardalis Boulenger, 1904, p. 80.

Peters and Doria’s type description of their Lygosoma (Hinulia) elegantulum (1878,
p. 344) has been translated.

“A Lygosoma reddish-brown above, black spotted; underside yellowish; ear opening
quite smooth; prefrontals separated; posterior of each pair of frenals trapezoidal; four
supraorbitals; body scales in 26 or 28 series, eight rows on the back; rather short feet.

306 REPTILES IN THE MACLEAY MUSEUM,

Length from tip of snout to base of tail 65 mm., head 16 mm., forelimb 15 mm., hand
with 4th digit 6 mm., hindlimb 22 mm., foot with 4th digit 11 mm.

Hab. North Australia, Somerset. (D’Albertis.)

A species akin to H. elegans Gray and dH. fasciolata Ginth. It is of the same
colour, reddish-brown dorsally spotted with black, but without the bands which unite
with the lateral bars. The scalation of the head does not differ essentially from that of
H. elegans. The ear, which has its margin perfectly smooth, seems smaller than in
the last species. The scales of the body are smooth and shining as in H. elegans, but
they are larger. In fact in our species we count 26-28 longitudinal rows, while in
H. elegans and fasciolata there are 32-33. The two median rows on the back have
larger scales and the two median preanals are also noticeably enlarged. Those of the
tail are very large, perfectly smooth, and form towards its base 15 longitudinal series.
Those of the middle inferior row are larger than the others.

That which is really noticeable in this species is the proportions of the limbs. In
H. elegans the forelimb placed along the snout reaches to the anterior margin of the
eye, but in our species only to the angle of the mouth. The hindlimb laid along the
trunk only reaches a little more than half the distance between groin and axilla.

The underside is whitish.

It is a species discovered by D’Albertis. He collected two specimens at Somerset,
Cape York.”

Two specimens in the collection of the Australian Museum have been examined.
One, a topotype of Lygosoma (Hinulia) elegantulum Peters and Doria, R 9599, taken
by Melbourne Ward at Cape York near Somerset, has been invaluable for comparison
with S. n. nigricaudis. A specimen from the Murray Islands, collected by Hedley and
McCulloch, is discussed.

Table of Measurements in mm. and Other Characters of S. n. elegantulus.

R 9599 R 4512

Snout-vent he of Et. : AS at AA MS 56 48
Pally cus ote KA ok = Sts os BS a ads 38 +
Snout-forelimb oe oF se ae oy ais Pee ins 19 18
Axilla-groin ae ve a ms Sts teats Es ba 35 23
Head, length Ee ae mA By ae be an Ae 10 10
Head, width ae ays she se at ys 2.2 Ho 8 7
Forelimb, length .. wa a a aS oy a5 aa 13 10:5
Hindlimb, length 4g ne ate ae a Ae ss 17 17
Width of body .. Ms a Lt aa ae ay 10 28)
Axilla-groin/Snout- forelimb os 5 i ae ss ae LO 1:84 1-28
Tail/Body .. : Be 1:38 _
Width of suture of Poetralll with fr Ontanaeal to width of frontal oe 1/4 1/3
Contact of 1st supralabial with nasal a by 2/3 2/3
Width of suture of frontonasal with frontal to width of eel, Be 1/5 1/8
Length of interparietal to length of frontal .. as ac ah 75% UY
Number of nuchals, left and right .. oe 3*-3* 4-4
Seales touching postero-lateral border of parietals,; left al right 4-3 4-3
Supraciliaries ae sf Bs oh of ax a ae 9 9
Infralabials . ae ate age aU: y: abs of 6 6
Lamellae beneath 4th io we she ae Ae: as ae 18 20
Midbody scale rows Pr faa a8 of oe aS ae 28 30

* Rather irregular.
+ Including upper secondary temporal, interpolated scales and nuchals.

The lower border of the posterior loreal is in contact with the 2nd supralabial only.
Colour of R 9599 is rich reddish-brown dorsally and on the sides of the head and tail.
Lighter brown on sides of body. The underside is whitish except for a few short
black bars along the margins of the infralabials. The head dorsally is almost unflecked,
but dotted with black along the posterior sutures of the parietals and on the temporals
and supra- and infralabials. About a dozen interrupted and rather irregular bars
formed of heavy black spots run across the back from behind the head to the length
of a forelimb behind the shoulders and extend half way down the sides. Posterior to

BY SLEPHEN J. COPLAND. 307

this area the back of the body is almost uniform brown. The tail is spotted except
for the proximal quarter, but there is no dense aggregation of pigment anywhere.
The sides are practically immaculate except for the anterior bars.

R 4512 is rather a puzzling specimen. It is hard to place, being a juvenile with its
tail broken off and the colour bleached. Banding was probably heavy across the
forepart of the body and places it near S. n. elegantulus. This is in spite of the
fact that the Murray Islands are only about 30 miles from Darnley Island, the type
locality of S. n. nigricaudis. The two sets of islands, however, were never joined,
being of volcanic origin, and must have been colonized separately. Geographical and
zoological aspects of the Murray Islands bearing on this matter are dealt with in the
fascinating book by Yonge (1930, pp. 183-199, et al.). Although R4512 has been
tentatively assigned here to S. n. elegantulus, definite identification must await other
material from the Murray Islands giving information unavailable from this single,
damaged, bleached and juvenile specimen.

Figs. 6-7.—Ventral views of head of lectotype of S. n. nigricaudis and holotype of S. p.
-pardalis. Arrangements of the scales are characteristic and apply to the two subspecies of each
species. M, mental; P, postmental; C1, C2, C3, 1st, 2nd and 38rd pairs of chin-shields; L1 and
L2, 1st and 2nd infralabials.

Retention of S. elegantulus as a subspecies of S. nigricaudis appears to be well
justified. This step is supported by the fact that neither Peters and Doria in their
original description (1878, p. 344) nor Boulenger in the first extended description (1887,
p. 235), which was followed closely by de Rooij (1915, p. 182), noted black on the
tails of their specimens. This most conspicuous feature of S. n. nigricaudis, which even
now is striking in specimens collected nearly 70 years ago and indeed so impressed
Macleay, as shown by hig specific name, could not have been overlooked. Except for this
omission Peters and Doria’s rather generalized description applied exactly to NS. n.
nigricaudis. SS. n. nigricaudis also differs from 8S. n. elegantulus, on the basis of
Boulenger’s description, in the following characters. The distance between the end of
the snout and the forelimb in 8S. n. nigricaudis is contained from 1:48 to 1:68 times
(average 1:57) in the distance between axilla and groin (NS. n. elegantulus, 1:60 to 2-00).
Two specimens of S. n. nigricaudis have nine supraciliaries, three 10 (S. n. elegantulus,
eight, rarely seven or nine). S. n. nigricaudis has from two to four pairs of nuchals.
but there is generally a different number on each side (8S. n. elegantulus, three or four
pairs). The length of the tail to the length of head and body is 1:18 to 1:23 in S. n.
nigricaudis (1:33 to 1:50 in S. n. elegantulus).

Locality Records.

The following records given by various authors almost certainly apply for the most part
to S. n. elegantulus, some probably to S. x. nigricaudis and 8S. p. erro, but it is most improbable
that any apply to S. p. pardalis.

Bara Bara: de Rooij (1915, p. 183) under Lygosoma elegantulum.
Bégowré River: de Rooij (1915, p. 183) under Lygosoma elegantulum.

DD

308 REPTILES IN THE MACLEAY MUSEUM,

Bloomfield River: Loveridge (1934, p. 352) under S. pardalis.

Bogadjim: de Rooij (1915, p. 183) under Lygosoma elegantulum.

Burnett River ?: Oudemans (1894, p. 140) under Lygosoma elegantulum.

Chillagoe district: Broom (1898, p. 648) under Lygosoma elegantulum.

Coen: Loveridge (1934, p. 352) under S. pardalis.

Ferguson Island: Boulenger (1895, p. 29) under Lygosoma elegantulum (A. S. Meek).

Fly River: Boulenger (1887, p. 235) under Lygosoma elegantulum (Rev. S. Macfarlane).

Haveri: de Rooij (1915, p. 183) under Lygosoma elegantulum.

Inawi: de Rooij (1915, p. 183) under Lygosoma elegantulum.

Islands of Torres Straits: Giinther (1877, p. 413) under Hinulia striatula.

Lake Barrine: Loveridge (1934, p. 352) under S. pardalis.

Lake Sentani: de Rooij (1915, p. 183) under Lygosoma elegantulum.

Lankelly Creek, MclIlwraith Ranges: Loveridge (1934, p. 352) under S. pardalis.

Lizard Island ?: Boulenger (1904, p. 80) under Hinulia pardalis (A. E. Finckh) 1901.

Merauke: de Rooij (1915, p. 183) under Lygosoma elegantulum.

Moroka, 2,300 feet: de Rooij (1915, p. 183) under Lygosoma elegantulum.

Mount Carbine: Loveridge (1934, p. 352) under S. parddlis.

Mount Spurgeon: Loveridge (1934, p. 352) under S. pardalis.

Mount Victoria: de Rooij (1915, p. 183) under Lygosoma elegantulum.

Murray Island: Boulenger (1887, p. 235) under Lygosoma elegantulum (Rev. S.
Macfarlane).

Rocky Scrub, MclIlwraith Ranges: Loveridge (1934, p. 352) under S. pardalis.

Rossel Island: de Rooij (1915, p. 183) under Lygosoma elegantulum.

Somerset: Peters and Doria (1878, p. 3844) type locality of Lygosoma (Hinulia)
elegantulum.

Sorong: Barbour (1912, p. 91) under Sphenomorphus elegantulus.

Sunday Island ?: Glinther (1875, p. 11) under Hinulia striatula, and Boulenger (1887,
p. 235) under Lygosoma elegantulum (J. B. Jukes).

Waigeu: de Rooij (1915, p. 183) under Lygosoma elegantulum.

Specimens from Waigeu, Begowre River, Lake Sentani and Merauke were examined
personally by de Rooij; authority for his other records is not given.

All localities mentioned are shown on the map, except five. Three of these given
by de Rooij as from New Guinea—Bara Bara, Haveri and Moroka—have not been
traced. Bara Bara may be Bira Bira (10° 39’ S, 150° 20’ E) near the eastern extremity
of New Guinea, but cannot be Bara (3° 9’ S, 126° 8’ EF) on Boeroe or Buru Island to the
west of Ceram; and Haveri and Moroka are believed to be near the headwaters of the
Aroa River west of Mount Victoria. The other two are the Burnett River (25° 10’ S,
152° 6’ EH) in south Queensland, and Sunday Island (16° 20’ S, 123° 10’ E) off the coast
of Western Australia.

Giinther (1875, p. 11) placed specimens from New South Wales and Sunday
Island, off the coast of Western Australia, and later (1877, p. 413) other specimens from
islands in Torres Straits, in Hinulia striatula Steindachner. Lizards from these three
localities are almost certainly respectively Sphenomorphus tenuis tenuis (Gray), S.
isolepis isolepis (Boulenger) and S. nigricaudis elegantulus. There is a strong
probability, however, that the lizards recorded as coming from Sunday Island were
collected elsewhere. Boulenger (1877, p. 235) identified two lizards from “Sunday
Island ?” as Lygosoma elegantulum. These two specimens appear to be the ones on
which Gunther based his record. Boulenger also had specimens from the Fly River (1),
Murray Island (7), and islands of Torres Straits (3). Lizards from these localities
were collected by the Rev. S. Macfarlane. The “Sunday Island” specimens were
presented by J. B. Jukes. Boulenger questions this last record. On the preceding
page (p. 234) he had erected Lygosoma isolepis to include five specimens from Nicol
Bay, Swan River and Australia, previously placed in Hinuwlia tenuis and H. striatula.
Other specimens placed under H. striatula he identified as Lygosoma elegantulum.

Oudemans (1894, p. 140) identified a lizard from the Burnett River in southern
Queensland as Lygosoma elegantulum. Although he said it agreed exactly with
Boulenger’s description except for having 30 midbody scale rows instead of 28, the
specimen probably belongs to one of the closely allied species.

Boulenger (1895, p. 29) notes an individual collected by A. S. Meek on Ferguson
Island.

BY STEPHEN J. COPLAND. 309

Broom (1898, p. 643), in his paper on the lizards of the Chillagoe district, says of
Lygosoma elegantulum: “This beautiful Lygosoma is not uncommon. Though the large
majority of the lizards are found chiefly in the limestone district, all the specimens I
obtained of this form were from the granite region where they were found under
fallen timber.”

Barbour, in giving the first Sorong record (1912, p. 91), says: “A single specimen
from Sorong, Dutch New Guinea, agrees well with Boulenger’s description. Nevertheless,
even a good diagnosis alone, without specimens for comparison, is never very con-
vincing; and if this Sorong specimen were compared directly with authentic examples
from Queensland or British New Guinea, it would very possibly: be found to be different
from SNS. elegantulus. I cannot find any record that it has been previously taken in
this region of Papua. It is known from the type locality, and British and German
New Guinea.”

Loveridge (1934, p. 352) under the heading of Sphenomorphus pardalis (Macleay)
gives the following synonymy, locality records and remarks:

“Hinulia pardalis Macleay: Mocoa nigricaudis Macleay: Lygosoma (Hinulia)
elegantulum Peters and Doria: Homolepida crassicauda Barbour (not of Duméril).

1 (M.C.Z. 9485) Darnley Island, T.S. (H. L. Clark) 1913: 1 (M.C.Z. 10199) Bloomfield
River, Q. (Australian Mus.) 1914: 1 (M.C.Z. 35403) Lake Barrine, Q. (P. J. Darlington)
1932: 3 (M.C.Z. 35404-6) Rocky Scrub, McIlwraith Ranges, Q. (P. J. Darlington) 1932:
2 (M.C.Z. 35407-8) Lankelly Creek, Q. (P. J. Darlington) 1932: 1 (M.C.Z. 35409) Mt.
Spurgeon, @. (P. J. Darlington) 1932: 1 (M.C.Z. 35410) Mt. Carbine, Q. (P. J.
Darlington) 1932: 1 (M.C.Z. 35411) Coen, Q. (P. J. Darlington) 1932.

The Bloomfield River specimen was received as Omolepida crassicaudum; that
species, however, has 22 midbody scale-rows. Midbody scale-rows 24-30 (24 in one
Rocky Scrub skink only, 30 in the Mt. Spurgeon skink only), average 27; frontonasal
forming sutures with the rostral and frontal; usually 3 (2-4) but often an azygous
arrangement of scales bordering the parietals posteriorly, such as 2 on one side, 3
on the other, or 2 and 4 in No. 35410; adpressed limbs do not nearly meet; lamellae
beneath the fourth toe 16-20, average 18. Largest skink (No. 35407) measures 186
(68 +118) mm. I follow Zietz (1920, p. 208) in referring elegantulum to the synonymy;
judged by a comparison of the descriptions the course seems justifiable. I venture
to add nigricaudis on the strength of our No. 9485 which is a topotype and does not
differ in any structural character but only in details of colouring. It lacks the
concentration of dots on the base of the tail which caused Macleay to name it
nigricaudis but it is certainly conspecific with the rest of our series.”

Confusion of S. elegantulus (Peters and Doria) with S. pardalis (Macleay) may be
traced to two sources, the first being the scantiness of Macleay’s original description.
The second occurred between 1901 and 1904. In these ProcrEpInes, xxvi, 1901, p. 214,
under Notes and Exhibits we find: “Mr. S. J. Johnston exhibited a collection of lizards
procured by Mr. A. E. Finckh of the University of Sydney, on Lizard Island, during
his visit to the Barrier Reef in the early part of this year. It comprised four
species, represented by about 35 specimens, namely, Lygosoma pardalis, Macl., .. .”

One of these lizards was sent to Boulenger, who says (1904, p. 80) under the
heading of Hinulia pardalis: “The lizard which bears this name was so imperfectly
described by Macleay in 1877 (Journ. Linn. Soc. N.S.W., ii, p. 62), from a collection
made at Katow, New Guinea, that, when revising the Scincidae in 1887, I could refer to
it only in a footnote (Cat. Liz., iii, p. 209) appended to the general synonymy of the
genus Lygosoma. I have now received, through the kindness of Mr. 8S. J. Johnston,
of the Technological Museum, Sydney, a specimen collected by Mr. A. EH. Finckh on
Lizard Island, Queensland, which, he informs’ me, he has compared with the type of
Hinulia pardalis in the Macleay Museum, University of Sydney, and found identical
with it. From this specimen I conclude that H. pardalis is the same as Lygosoma
elegantulum, Peters and Doria (Ann. Mus. Genova, xiii, 1878, p. 344), and, as the
former name has priority, I propose in future to designate this rather common species
as Lygosoma pardalis, Macleay.”

310 REPTILES IN THE MACLEAY MUSEUM,

I can only suppose that a mistake was made when the two specimens were compared
in the Macleay Museum. A rapid examination might have led to characters such as
the number of midbody scale rows, relationships of the interparietal, postmental and
other scales being overlooked, and the conspicuous differences in markings and colour
(as they appear now and as they were originally noted by Macleay for S. pardalis and
Peters and Doria for S. elegantulus) may have been regarded as variable and so
unimportant. Possibility of a mistake is made more probable by the fact that it is
unlikely that Mr. Johnston had specimens of S. elegantulus for comparison. The
alternative is that S. pardalis occurs on Lizard Island. In this case Boulenger would
have made the error of confusing two species. With comparative material at his
disposal and in view of his systematic methods in the Catalogue of Lizards and
elsewhere, this possibility must be regarded as remote. With Boulenger’s authority it
was inevitable that the misunderstanding would continue until a thorough re-examination
was made of Macleay’s type.

S. elegantulus was synonymized with S. pardalis by Zietz (1920, p. 208) and
Loveridge (1934, p. 352).

VI. MicrRATION AND RELATIONSHIP OF FORMS.

Concerning the evolution and migration of the two species dealt with in this paper,
nothing but profit can be gained by presenting an hypothesis in accord with the known
facts, which may serve as a framework for future research, provided that the hypothesis
is regarded purely as an hypothesis. It appears probable that the common stock, which
resembled the present S. n. elegantulus fairly closely, arose in western New Guinea.
One wave of migration spread into the area now occupied by S. n. nigricaudis, where
it differentiated into that race. Another wave colonized north Queensland. While part
of the population remaining in Cape York Peninsula remained practically unchanged,
another became isolated and evolved into a form with a reduced number of midbody
scale rows, a characteristic arrangement of scales of the lower jaw, and aggregation
of pigment along the flanks. This form retained the colouring of the parent stock and
is represented by S. p. erro. Representatives of this population became cut off on
Barrow Island during the positive movement of sea-level contemporaneous with the
growth of the Great Barrier Reef. With the hastening of selection and evolution of
pure lines associated with isolation of small populations on small islands, S. p. pardalis
was evolved. It is of course quite probable, and may square with problems of isolation .
much better, that north Queensland was colonized twice; the first wave becoming
differentiated into the S. p. erro stock before the second invasion of individuals still
close to the commen S. n. elegantulus stock from the New Guinea reservoir. If it
later proves true that the Murray Islands race is S. n. elegantulus, and lack of
adequate material for examination at present makes this uncertain, there may have
been independent colonization of these islands in the Darnley-Murray Islands area.
Transport by canoe or floating timber must have been haphazard, and the degree
of chance is enhanced by the fact that the direction of the prevailing winds changes

during the year. In summer the north-west monsoon replaces the south-east Trade
winds.

VII. ACKNOWLEDGEMENTS.

I wish to acknowledge help and advice from Professor W. J. Dakin and Professor
H. A. Briggs, of the University of Sydney; Dr. A. B. Walkom, Mr. J. R. Kinghorn and
Mr. W. A. Rainbow, of the Australian Museum; and Mr. T. Iredale, formerly of the
Australian Museum. I have to thank Professor A. H. McDonald and Mrs. R. A. Shaw,
of the University of Sydney, for translations of Latin and Italian respectively, and
Miss A. G. Burns, of the Department of Zoology, University of Sydney, for the
photographs. The Land Administration Board, Brisbane, kindly forwarded details of,
and a plan showing the position of Barrow Island.

BY STEPHEN J. COPLAND. Apibit

VIII. BIBLIOGRAPHY.

BargBour, THOMAS, 1912.—A Contribution to the Zodgeography of the Hast Indian Islands, No. 1.
Mem. Harv. Mus. Comp. Zool., 44: 1-168.

, 1914.—On Some Australasian Reptiles. Proc. biol. Soc. Wash., 1914, 201-205.

BouLENGER, GEORGE ALBERT, 1887.-—-Catalogue of the Lizards in the British Museum (Natural
History), 2nd Ed., London. Vol. 3.

—, 1895.—On a Collection of Reptiles and SBatrachians from Ferguson Island,

D’Entrecasteaux Group, British New Guinea. Ann. Mag. nat. Hist., (6)16:; 28-32.

———_—, 1904.—Note on Hinulia pardalis of Macleay. Ibid., (7)14: 80.

Broom, R., 1898.—On the Lizards of the Chillagoe District, North Queensland. Proc. LINN.
Soc. N.S.W., 22 (3): 639-645.

Davis, CoNSETT, and Len, D. J., 1944.—The Type Concept in Taxonomy. Aust. J. Sci., 7 (1):
16-19.

DE Rooig, Neuuy, 1915.—The Reptiles of the Indo-Australian Archipelago, Lacertilia, Chelonia,
Emydosauria. Leiden. Vol. 1.

DuNN, E. R., 1934.—Systematic Procedure in Herpetology. Copeia No. 4: 167-172.

GARMAN, SAMUEL, 1901.—Some Reptiles and Batrachians from Australasia. Bull. Muws. comp.
Zool. Harv., 39: 1-14.

GUNTHER, ALBERT, 1875.—The Zoology of the Voyage of H.M.S. Hrebus and Terror. London.

— , 1877.—Description of Three New Species of Lizards from Islands of Torres Straits.
Ann. Mag. nat. Hist., (4) 19: 413-415.

KINGHORN, J. R., 1931.—Herpetological Notes, No. 2. Rec. Aust. Mws., 18 (38): 85-91.

LOVERIDGE, ARTHUR, 1934.—Australian Reptiles in the Museum of Comparative Zodlogy,
Cambridge, Massachusetts. Bull. Mus. comp. Zool. Harv., 77 (6): 2438-383.

MACLEAY, WILLIAM, 1877.—The Lizards of the ‘‘Chevert” Expedition. Proc. LINN. Soc. N.S.W.,

2 (1): 60-69.

Mayr, ERNST, 1942.—Systematics and the Origin of Species. Columbia University Press,
New York.

OUDBEMANS, J. TH., 1894.—Hidechsen und Schildkréten, Zoologische Forschungsreisen in
Australien und dem Malayischen Archipel ... Richard Semon, 5. Denkschr. med.-naturw.

Ges. Jena, 8: 127-146.
PETERS, W., and DoriA, G., 1878.—Catalogo dei Rettili e Batraci raccolti da O., Beccari, L. M.

D’Albertis e A. A. Bruijn nella sotto regione Austro-Malese. Ann. Mus. Stor. nat. Genova,
13: 323-450.

YONGE, C. M., 1930.—A Year on the Great Barrier Reef. Putnam, London and New York.
Zietz, KF. R., 1920.—Catalogue of Australian Lizards. Rec, 8. Aust. Mus., 1 (3): 181-228.

EXPLANATION OF PLATE XI.
Figs. 1-4.—Dorsal views of the four races.
Fig. 1.—Sphenomorphus pardalis pardalis.
Fig. 2.—S. p. erro.
Fig. 3.—Sphenomorphus nigricaudis nigricaudis.
Fig. 4.—S. n. elegantulus.
Body lengths of specimens (including head) are 72, 59, 76 and 56 mm. respectively.

[Photos.—Miss A. G. Burns. ]

el2

STUDIES ON TROMBIDIIDAB (ACARINA).
SOME OBSERVATIONS ON THE BIOLOGY OF THE MICROTROMBIDIINAE.
By R. V. Soutrucott, M.B., B.S.
(Three Text-figures. )

[Read 28th November, 1945.]

The Australian members of the subfamily Microtrombidiinae Thor 1935 have
recently been restudied by Womersley (Rec. S. Aust. Mus., viii (2), 1945, 293-355)
and he described the larva of Camerotrombidium simile (Hirst 1928). Boshell and
Kerr (Revist. Acad. Colomb. Cienc. Exact. Fis. Quim. y Nat., xvii, 1942, 110-127) have
described the larva of their adult species Manriquia bequaerti B. and K. 1942, which
Womersley (1945, 294) refers to the genus Hchinothrombiwm Womersley 1937. Other
correlations of adults and larvae in this subfamily have been suggested in Europe,
apparently on less secure grounds; and also for some Australian material (see
Womersley, J. Linn. Soc. Lond., Zool., xl, 1936, 114-115).

In this paper is placed on record that in an Australian species of this subfamily,
Microtrombidium hirsutum Womersley 1945, the larval stage is suppressed, the eggs
hatching direct to nymphs. As far as I am aware, such a life history has not previously
been reported in the Trombidiidae, although well Known for other Acarina. The egg
and nymph are described and figured here. In addition an outline drawing of the adult
of the closely related Microtrombidium karriense Womersley 1934 is given.

MICROTROMBIDIINAE Thor 1935.

Ottoniinae Thor 1935; Womersley 1942. Microtrombidiinae Thor 1935; Womersley
1937, 1945.

MIcCROTROMBIDIUM Haller 1882.

Jber. Ver. Wurttemb., xxxviii, 1882, 322. Genotype, M. purpureum Haller 1882;
[non] Trombidium pusillum Hermann 1804 (see Willman, Zool. Anz., cxxxi (9-10),
1940, 255).

MICROTROMBIDIUM HIRSUTUM Womersley 1945. Figs. 1, B—-H, 2, 3.

Rec. S. Aust. Mus., viii (2), 1945, 312. Microtrombidium karriensis Wom. 1937,
Ibid., vi (1), 87 (part, locality record only).

PAGES IMIS, 5 IB, 1a, This has recently been separated by Womersley from
M. karriense Wom. 1934. Figures of the dorsal setae, for comparison with those of the
nymph, are given.

Egg: Fig. 1, B-D. Red. Spheroidal to ovoid or somewhat irregular. Surface smooth
or very slightly wrinkled, not patterned. Average size about 350u long by 300 across.
See further under notes on biology.

Nymph: Figs. 1, G, H; 2, 3. Red. Body cordate as in the adult, legs somewhat
more thick-set. Body somewhat flattened dorsoventrally; body length (to anterior end
of crista) 4004 average (varied from 350-4504”); width 300u average (varied from
250-3504). Crista 834 long from anterior tip to centres of sensillae. Sensillary setae 2,
simple, filiform, 804 long; centres of sensillae 16% apart. Eyes 2 + 2, on shields, behind
middle of crista. Dorsal setae long, slender, heavily ciliated, increasing gradually in
length posteriorly and laterally over the dorsum, 18-52u long. Ventral setae similar

BY R. V. SOUTHCOTT. 313

to dorsal. Genitalia with 2 pairs of suckers. Palpi stout with stout accessory claw,
with simpler arrangement of setae than adult; only one pectine of setae present on the
palpal tibia, this on the dorsal side. One stouter seta (or spine) present at about
the centre of the ventral surface of the palpal tibia. Chelicerae falciform, inner
edge serrate. Legs comparatively more thick-set than in adult; I 4304 long, II 315uy,
III 300u, IV 3804 (all lengths including coxae and claws); 2 claws to each tarsus,
strong, simple, falciform. Tarsus I 1174 long by 68u high; metatarsus I 59 greatest
length by 37u high; a number of simple sensory setae are present on the legs, especially
terminally, in addition to the normal ciliated setae.

E ‘ GS.
3 a PD y
(aoe ey LY E of
G A |
B \

Seu TH@rT-

Fig. 1.—WMicrotrombidium karriense Wom. 1934, and Microtrombidium hirsutum Wom. 1945.
A, M. karriense, entire, outline, ventral; B-H, M. hirsutum. B, C, Egg, lateral; D, Egg, dorsal;
H, Anterior dorsal seta of adult; F, Posterior dorsal seta of adult; G, Anterior dorsal seta of
nymph; H, Posterior dorsal seta of nymph.

Biology: The adult (ACB 31) was captured by myself in a damp situation at
Waterfall Gulley, in the Mt. Lofty Ranges, South Australia, on 24th August, 1938. It
was confined to a tube with a small amount of damp clay and sand from the same
locality. Initially the tube was examined weekly. Up till 24th September, 1938, the
adult remained plump, and became active on stimulation; no eggs nor larvae yet seen
in the tube.

1.x.38. Adult appears somewhat shrunken; a cluster of 6 red translucent eggs
present, which are very slightly wrinkled on the surface. The eggs large in
comparison with the adult mite. Adult active on stimulation. Tube wet.

8.x.38. Some lighter patches appearing in the eggs.

9.x.38. Eye-spots of embryos clearly visible (not shown in text-figures).

16.x.38. Leg somites of one embryo visible. Eggs now examined at least once
daily.

314

18.x.38.
24.x.38.
25.x.38.
26.x.38.
ZOO Se
5.xi.38.
12.xi.38.
Zilexae 38s

STUDIES ON TROMBIDIIDAE,

Text-figures of eggs made.

The first egg hatched out, toa nymph. This nymph removed and mounted.
Four more nymphs hatched out. Adult active, well.

Adult and one nymph removed and mounted.

Tube now dry. At least one nymph seen alive.

Two nymphs seen wandering around tube.

One nymph only seen alive.

Tube dry. No mites seen alive.

Ni

1

x

SS wy
aN

elec SAY
SS

CS
SS

IS
NaS
5
aS
S)

SouTtHhcort

Fig. 2.—Microtrombidium hirsutum Wom. 1945. Nymph, freshly emerged, dorsal aspect.

a

santa int

BY R. V. SOUTHOOTT. 315

9.xii.38. Contents of tube emptied out and examined carefully. Four dead nymphs
were found, and several scraps of egg-skins. No unhatched egg found. The
nymphs and egg-skins were mounted. No signs in any way referable to larvae
were found. (Thus 6 eggs and 6 nymphs are accounted for.)

SESS
SNL

vos

DG i.

Sou
ao i
v : ial a

S = ES ~ SN x

>

SoutHcote

Fig. 3.—Microtrombidium hirsutum Wom. 1945. Nymph, freshly emerged, ventral aspect.

EE

316 STUDIES ON TROMBIDIIDAE.

Remarks: It is considered that the statement made here regarding the absence of
a larval stage, for this species at least, is firmly established. The time for development
in the egg, from laying to hatching, was 28-5 + 3-5 days (for 4 of the 6 eggs).

I have had the opportunity of comparing the adult specimen above with the type
of M. hirsutum, and other type material, for which I wish to thank Mr. Womersley.
The adult agrees completely with the type of M. hirsutum. The considerable reduction
and simplification of the palpal spines and setae of the nymph is of some interest from
the point of view of generic classification.

MICROTROMBIDIUM KARRIENSE Womersley 1934. Fig. 1, A.

Microtrombidium karriensis Womersley, Rec. S. Aust. Mus., v. (2), 1934, 191.
Microtrombidium (M.) karriensis, karriense Womersley, Ibid., vi (1), 1937, 87. Micro-
trombidium karriensis Womersley, Ibid., viii (2), 1945, 310.

This species has recently been redescribed by Womersley. As the type is now in
a damaged condition, an outline drawing of the ventral surface of the entire mite,
made some time ago, is given here.

A CONTRIBUTION TO A STUDY OF THE PHYSIOLOGY OF DECAY IN APPLES.

By Mary Casu, M.Sc.
(From the Department of Botany, University of Sydney.)

(Ten Text-figures. )

[Read 28th November, 1945.]

INTRODUCTION.

Fungal decay and physiological breakdowns are responsible for the wastage which
occurs in apples during storage and it is the object of this paper to discuss briefly some
of the factors relating to fungal decay.

The moulds most frequently responsible for damage in apple storage are Penicillium —
expansum and Gloeosporium album.

P. expansum is a typical wound parasite and most frequently gains entrance through
mechanical injuries such as stem punctures, insect injuries and necrotic tissue. The
lesions are soft and watery, light brown in colour, and are not restricted to any one
position but may occur on the stem, cheek, or calyx ends, or may develop as core rots
arising from spores that have penetrated the open calyx canal. About 80% of the
storage decay has been attributed to this blue mould soft rot. The fact that P. expansum,
given a favourable medium, can make a much better start at 0°C. than any other fungus
helps to explain its common occurrence in cold storage (Brooks and Cooley, 1917).

Gloeosporium album produces what is known as the ‘bull’s eye rot’. The rot
develops as a flat or concave, fairly firm, lesion, chestnut-brown in colour. The lesion
has a typical zoned appearance with a well-defined margin and the flesh beneath the
rot is dry and leathery. Like the blue mould, lesions produced by G. album may occur
in any position on the apple. Acervuli situated concentrically around the point of
infection appear when lesions are mature. This rot develops more slowly than the blue
mould soft rot and is prevalent in the later part of the storage life.

Experiments on these moulds carried out by the writer and discussed in this paper
are set out in two sections.

Under Section 1 the following factors were examined for the effect they produced
on the rate of decay in apples under storage conditions: variety of apple, maturity at
picking, district and locality of orchard.

Under Section 2 a study was made of the effect of the following factors on the
growth rate of P. expansum and G. album: (a) hydrogen ion concentration; (0b)
temperature; and (c) age of spore.

SECTION 1.
MATERIAL AND METHODS.

The investigations on fruit were carried out at the Food Preservation Laboratory
of the Council for Scientific and Industrial Research, Homebush, where storage rooms
with properly controlled temperatures were made available for the storage of the fruit
during the years 1941 and 1942. All fruit on arrival at the Food Preservation Laboratory
was stored immediately at 34°F. until removal for inoculation.

In 1941, the fruit was obtained from Batlow and Orange. The varieties used were
Delicious and Granny Smith and there were three periods of picking: first, second, and
third; the first and third picking refer to early and late maturity.

~All inoculation procedure was standardized and although inoculations took place
over a period of time, spores of similar age were used in all inoculations. The spores

318 THE PHYSIOLOGY OF DECAY IN APPLES,

of P. expansum were taken from cultures which had been incubated at 20°C. for
10 days; G. album from cultures incubated at 20°C. for 21 days. The method of
inoculation was a modification of that developed by Granger and Horne (1924). A
sterilized cork-borer was inserted obliquely into the apple to a specified depth, then
withdrawn lifting up a triangular flap of tissue for the insertion of the inoculum (250
spores per loop). The flap was then closed and sealed with sterile paraffin. The fruit
was then wrapped and packed in cases and stored at 32°F. and 40°F.

The apples were sampled five at a time and the rate of decay was measured by
Gregory and Horne’s (1928) method of radial advance.

During the following year (1942) a more detailed examination of fruit from one
district was carried out using the same technique. For this investigation early and
late maturity fruit were obtained from the following orchards at Orange: (a) Delicious
fruit from the orchards of Messrs. Fox-Martin, Scott, Tonking and Coote; (0) Granny
Smith fruit from Scott’s orchard.

The inoculated fruit was stored at 40° F. in 1942.

EXPERIMENTAL RESULTS.

There was a general tendency for the fruit at the late maturity stage to be more
susceptible to invasion by both fungi. The effect was more striking in fruit from
Orange, and where maturity differences were pronounced, the effect was observed at
32°F. and 40°F. An example of the effect of late maturity picking on the rate of
rot development is illustrated in Fig. 1.

fe VANTUIRS YC
z = SHE es AOF
ee O=2ZN0
ty IN BIRD)
GC 26
pass
{20 Pn ( ,
a og Sale
ie
es i) i EM © |
O - at ae
ms phe Page
yO ras Bee (Oe
© : yee ans
& 05 ae Cra
00 | | {

O 0) 20 SO 40 50 oe) 70 & 90 100 Ke)

Tle EN (DIANE S

Fig. 1.—The effect of maturity on the radial advance of P. expansuwm in Delicious apples
from Orange.

Delicious fruit rotted at a faster rate than the Granny Smith fruit. This varietal
effect was most pronounced in the fruit from Batlow inoculated with G. albwm stored
at 40°F. (Fig. 2).

It was observed that, in fruit from Batlow, greater differences occurred in the rate
of decay between varieties than in the same variety at different maturities. In fruit
from Orange the differences due to maturity were more pronounced than the varietal
effect.

BY MARY CASH... 319

= VALORES 0 W/ MEY eqn \/

U X“IST — DELICIOUS
2 O-e ND ---- GRANNY SMITH
im 25 N= SY (20)
LJ

©

&

=

®)

<

#

<

(a)

<

ag

0 20 40 60 880 00% FI2Z0" 146

TIME IN DAYS

Fig. 2.—The effect of variety on the radial advance of G. album at 40°F.—Batlow.

IGO 180

A district-effect was observed in the Delicious fruit: fruit from Batlow showed a
marked tendency to decay at a faster rate than fruit from Orange (Fig. 3). No such —
district-effect was shown by the Granny Smith fruit.

O MATURITY DISTRICT

Z = Sef — BATLOW
= O- 2ND ORANGE
S 2:5 [S\= SRD ye
z

>

O

x

RADIAL

6 @e 40 60 8 [OO GIZOnN aS Om CO en SOM COOR a ae®

TIMES Ne DAS

Fig. 3.—The effect of district on the radial advance of G. album at 40°F.—variety, Delicious.

320 THE PHYSIOLOGY OF DECAY IN APPLES,

Data obtained from the investigation carried out in 1942 indicated that maturity
differences were not as marked as varietal differences. There was, however, the same
general tendency for the late maturity fruit to be more susceptible to fungal invasion
than the early maturity.

Locality of orchard had no effect on the growth of P. expansum. Early maturity
fruit from Scott’s orchard, inoculated with G. album, rotted at a slower rate than fruit
from the other orchards. Differential rotting was not apparent in the late maturity
fruit (Fig. 4).

3-0 a

LATE MATURITY
FOX MARTIN X
(ten ©)

2-5

= ee) SCOR == 6
Z 05 GOO Ee A
LJ

©)

VL

<< 3-0

>

a

tL 25

I) ao)

<x

@Qeeers

<I

(ag

TIME IN AS

Fig. 4.—Radial advance of G. album in Granny Smith and Delicious apples.

With a view to determining the cause of this differential rotting between varieties
and maturities, the following possibilities were investigated: (a) hardness of tissue;
and (0b) pH of sap.

To determine whether hardness was correlated with susceptibility to decay, an
examination was carried out on the hardness of the apples. This was measured by
the resistance of the cells to a standard plunger. No significant difference between
varieties and maturities was seen in the hardness of the fruit.

The pH of the expressed juice of apples of all maturities and varieties was
determined. Differences in hydrogen ion concentration between maturities were not

IN @QCULAR DIVISIONS
* Wn o N = oy) e) > wa fo)
oO .o) ro) ° _9o fe) O° ° ° °

Ww
oO

)

o
ft

ELONGATION OF GEINM WUE S

pb
is)

4 6

BY MARY CASH. 321

GRAPH T
SORENSENS CITRATE
KEY TO pH SERIES

G-- 4-0
A-- 3°8
Oo O58
0-3-0 :
X26
AF
4 6 8 10 12 14 16 1S 20 22 24 26 28
p
GRAPH IL

MCILVAINE’S STANDARDS

aN
is>)
0
ro)
f
a
a
©
%
ro)
iw)
i)
N
bh
ow)
(6)
N
o

GRAN ml L000
ACL “AGE MINORU aS)

10 l2 14 16 18 20 ae 24 26 28

8
EME ENE OURS

Fig. 5.—Elongation of germ tubes of P. expansum in relation to hydrogen ion concentration.

322 THE PHYSIOLOGY OF DECAY IN APPLES,

significant but the hydrogen ion concentration of Granny Smith juice (pH 3-6) was more
acid than that of Delicious juice (pH 4:0). Successful attack is due partly to chemical
changes in the cell such as hydrogen ion concentration, sugar content and pectin. It
seems possible therefore that this difference in hydrogen ion concentration observed
between varieties may account for the differential radial advance of the rots. This
aspect was therefore made the subject of further experiment (Section 2).

GRAPH JI MCILVAINES STANDARDS

Su

pH SERIES

30

2S)

20
= 15
=

10
TE,
= 5;
”
WJ
Zz
O
5 GRAPH IL Hel = Ke aS aRies
VO s9 ae SAO
[ln 5
eo = 3:8
nee 26 pH SeRIES
if 35 4:0
s
< . 3:
O

WEE YING DAYS

Fig. 6.—The growth of P. expansum in relation to hydrogen ion concentration.

BY MARY CASH. 323

SECTION 2.
(a) The Effect of Hydrogen Ion Concentration on Growth.

Studies of the moulds in culture comprised both germination and growth
experiments.

To determine whether hydrogen ion concentration accounted for differences in the
rate of decay between varieties, the fungi were grown at pH values closely approxi-
mating to the hydrogen ion concentration of the apple juices. The moulds were grown
in potato-dextrose broth at the following pH range: 2-6, 3-1, 3:6, 3-8, 4:0 and buffered
by Mcllvaine’s Standards, or Sorensen’s Citrate, or HCL-KCL Mixtures.

GRAPH I MC ILVAINE’S STANDARDS
45
40
35)
(30
(@)
25 Wie ye ©) pH
KM 220)
ee © 3-8
4s © 376
=
3:
>3 16) : |
Peo
5
Zz
iD SS Ee (ara oe 20 Zod de 40
uJ
E GRAPH I Hiei Kem HiaiURES
md]
O
U
LL
O
~
LJ
=
LJ
=
<
OQ

iM ENE eDAGS

Fig. 7.—The growth of G. album in relation to hydrogen ion concentration.

324 THE PHYSIOLOGY OF DECAY IN APPLES,

Sterile plaster of Paris discs were placed in Petri dishes containing 20 c.c. of
buffered broth. The growth rate was measured by recording the diameter of the
colonies and the discs were marked with pencil in mm. to facilitate readings (Cash,
1942). Each disc was inoculated in the centre with a 1 mm. mycelial square and the
experiment was carried out in replicates of five at each pH value. To eliminate the
effect of staling on growth, the discs were transferred daily under aseptic conditions
to Petri dishes containing fresh culture solution of the same pH.

Results—The examination of spore germination of P. expansum showed that pH
did not affect the latent period (Fig. 5, Graphs I and II). Irregular sprouting of the
germ tubes and an increase in the latent period at pH 2-6 in the HCL-KCL buffer
series suggest that unfavourable conditions were due to the type of buffer rather than
to unfavourable pH (Fig. 5, Graph III). Germination studies on G. album were not
carried out.

The growth of P. erpansum in Mcllvaine’s Standards series was independent of pH
but a well-defined optimum was observed at pH 3:6 in the HCL-KCL buffer series
(Fig. 6). G. album showed a maximum growth at pH 4-0 in all buffer types with a
consistent decline on the acid side (Fig. 7).

Normal growth took place at pH 2:6 in the HCL-KCL buffer series but the cultures
did not grow in MclIlvaine’s Standards series at a similar pH. This indicates the
necessity for using more than one buffer type to determine whether the effect is
chemical or due to hydrogen ion concentration.

(6) The Effect of Temperature on Growth.

The technique used for observing the latent period, percentage germination and
elongation of the germ tubes was to spread a dilute spore suspension over agar films

(Oc
{
Wy
QO
SC AG
er
x- PDA

ul) 30
a o— BROWNS AGAR
> 25
7 (0)
(oC
uJ
© 1S
Cao
&

5 ~(__ PENICILLIUM
oO O
D O 4 8 12 16 20 24 28
O
iz

TEMPERATURE

Fig. 8.—The effect of temperature and media on the latent period of P. expansum and
G. album.

BY MARY CASH. 325

corresponding to the type of media to be used in the growth studies. For the
examination of the effect of temperature on growth, colonies were grown in Petri
dishes containing 18 c.c. of 2% potato-dextrose agar, malt agar or Brown’s agar at
1:3°C., 3°C., 11°C., 14°C., 20°C. and 26°C. The experiment was carried out in replicates
of five at each temperature and growth was measured by recording the diameters of
the colonies.

Results.—The results given in Fig. 8 show that the temperature has no effect on the
latent period of G. album. As the temperature decreases there is an increase in the
latent period of P. expansum. In all cases 100% germination was obtained. Fig. 9

= PIDA
O- BROWNS AGAR
A- MALT AGAR

P EXPANSUM

G. ALBUM

RAVE OF GROW ITE

—_—_—
SS

—
a
_—

0 S) 10 15 20 AS) 30

MEMRERATURE SIN ~DEGCREES CENTIGRIADE

Fig. 9.—Growth rates of P. expansum and G. album on different media and at different
temperatures.

shows that, within the range of the experiment, optimal conditions for the growth of
P. expansum were at 20°C., whereas the optimal growth rates of G. album varied slightly
with the type of the medium and were dependent on time.

(c) The Effect of Age of Spore on Germination at Different Temperatures.

Germination studies were carried out with the spores of P. expansum and G. album
of different ages to determine whether the age of the spore lying on the fruit could
affect the initial rate of penetration into the fruit. Spores were obtained from cultures
of P. expansum of the following ages: 5, 42, 104 and 217 days, and from G. album
64, 132, 179 and 212 days old. All cultures used to determine the effect of age of
spore on germination had been stored at 3° C. Germination studies were carried out
aveleseCn 147C) and 25°C.

Results —wWithin the experimental range, spore age did not affect the latent period
of either P. expansum or G. album. Germination was 100%. Optimal rates of germ
tube elongation of P. erpansum were observed in the youngest spores with a progressive
decline as the spores age (Fig. 10). The spores of G. album showed a maturation
effect, the optimum rate for the elongation of the germ tubes varying with the
temperature.

This examination on the germination of spores of different ages may possibly
suggest that spore age does not affect initial penetration into the fruit but may play
a part in the rate of rot development. It is possible that the different rates observed
in the elongation of the germ tubes may be an initial effect only and requires further
investigation.

co

26 THE PHYSIOLOGY OF DECAY IN APPLES,

Or, PE PANS UM

x Cs MLM!

O 20 40 GO 80 100 =|20 40°. |G) 180°) *200mNs220mmN2z40

Ros Of ILONGATION OF GERM TUBES

ACE {Oh * SFORES IND AS

Fig. 10.—Effect of spore age on the rate of elongation of germ tubes of P. expansuwm and
G. album at different temperatures.

DISCUSSION.

Data obtained from these investigations present no adequate explanation for the
differential decay observed between varieties. Results of experiments on the growth
of P. expansum in culture at different hydrogen ion concentrations were not consistent
with the data obtained on the radial advance in the fruit. Optimal growth rates for
this fungus were observed at pH 3:6 in the HCL-KCL buffer series. This pH value
closely approximates to the pH of the Granny Smith juice (pH 3-5), yet the rate of
decay was faster in the Delicious fruit. At a hydrogen ion concentration similar to
that of the Delicious fruit, P. expansum showed a decrease in the growth rate in
culture. It appears, therefore, that other factors are responsible for differences in
the rotting of fruit by P. expansum. ;

The differential rates of radial advance in apples by G. album showed a close
correlation with growth in culture at different hydrogen ion concentrations, and,
therefore, hydrogen ion concentration may also possibly offer an explanation of the
differential rate of decay observed between varieties. The pH values found in apples
of the same maturity showed so much variation that it was impossible to attribute
differences in radial advance between maturities of the same variety to differences in
hydrogen ion concentration.

It has been found by other investigators (Wilkinson, 1938) that resistance to
invasion is influenced by manurial treatment. Wilkinson found that nitrogen manuring
had a marked effect on the incidence of bull’s eye rot, low nitrogen manuring showed
a 10% wastage due to G. album and a high nitrogen manuring a 31% wastage.
Though no data were obtained on the manurial treatments used on the orchards
at Orange it was found that, irrespective of maturity, the locality of the orchards had

BY MARY CASH. 327

no effect on the rate of decay due to blue mould. Fruit from Scott’s farm inoculated
with G. album rotted at a lower rate than fruit from the other orchards, but it is
doubtful whether this would be a manurial effect since this effect was not observed in
fruit at the late maturity. However, for both types of rots, the varietal effect was more
pronounced than any differences in orchard locality which may have been due to
manuring. The chemical composition of the fruit may influence the rate of decay, but
so far no chemical analyses have been carried out in conjunction with this investigation.

Since spore age has no effect on the latent period and percentage germination, it
may be assumed that spores of these moulds of any age, on the surface of the fruit at
the time of storage, are all capable of infection.

SUMMARY.

The effect of some environmental factors on the rate of decay of Delicious and
Granny Smith apples in storage has been studied and the following conclusions are
presented:

1. It seems that district from which the fruit comes and variety of fruit have a
marked effect on the rate of radial advance of the rot. Fruit at late maturity showed a
general tendency to rot at a faster rate than fruit at early maturity.

2. Several suggestions have been put forward which may help to account for the
differential decay observed between varieties but no completely satisfactory explanation
can be offered at present.

3. Inherent qualities of the apples such as hardness of the tissue and skin colour
have no measureable effect on the rate of decay.

4. Insufficient data prevent interpretation of the orchard locality and district
effects but it is possible that nutritive conditions may influence rate of decay.

5. Although the studies in the physiology of P. expansum and G. album are
incomplete the data obtained indicate that germination of the spores of G. album
is not affected by temperature, but with a decrease in temperature, an increase in
the latent period of P. expansuwm was observed.

6. Within the range tested, optimal conditions for the growth of P. expansum were
observed at 20°C. The position of the optimum of G. album was found to be dependent
on time.

7. Hydrogen ion concentration has a well-defined effect on the growth of G. album.
Within the experimental range a maximum was found at pH 4:0, and there was a
progressive decline in the growth rate as the culture media became more acid. The
growth of P. expansum in broth buffered by MelIlvaine’s Standards was found to be
independent of pH but an optimal effect was observed in the HCL-KCL series at
pH 3-6. This indicates the importance of using more than one buffer type.

8. Age of spore was found to have no effect on the latent period and percentage
germination of P. expansum and G. album at different temperatures.

ACKNOWLEDGEMENTS.

The writer wishes to express her thanks to Professor Eric Ashby in whose
department this work was carried out, to Dr. J. R. Vickery for permission to use the
facilities of the Division of Food Preservation, Council for Scientific and Industrial
Research, Homebush, and to Dr. John McLuckie and Dr. Lilian Fraser for their helpful
criticism in the preparation of the manuscript.

REFERENCES.

Brooks, C., and Coouey, J. S., 1917.—Temperature Relations of Apple Rot Fungi. J. Agric. Res.,
8° (4): 139-165. ;

CasH, Mary, 1942.—A Simple Method for Determining the Growth Rates of Fungal Colonies at
Different Hydrogen Ion Concentrations. Aust. J. Sci., 4 (4): 135.

GRANGER, K., and Horne, A. S., 1924.—A Method of Inoculating the Apple. Ann. Bot., 38 (149):
213-215.

Grucory, F. G., and Horns, A. S., 1928.—Fungal Invasion of Apples. Proc. Roy. Soc. Lond. B.,
102: 427-466.

WILKINSON, E. H., 1938.—Information on the Prevalence of Fungal Spots and Rots of Apples
in Cold Store at Long Ashton. Abstr. 363. A.R. Agric. and Hort. Res. Stat., Long Ashton.

328

THE DIPTERA OF THE TERRITORY OF NEW GUINEA. XIII.
FAMILY TABANIDAE. PART I. THE GENUS CHRYSOPS.

By Frank H. Taytor, F.R.E.S., F.Z.S., School of Public Health and Tropical
Medicine, University of Sydney.

(Plate x; two Text-figures. )

[Read 28th November, 1945.]

The present contribution, the first of a series on the family Tabanidae, like some
previous papers of this series, is not confined to species from the Territory of New
Guinea. It has been decided, after due consideration, on account of recent activities
in these areas, to give descriptions and illustrations of species from the Molucea Islands
eastward to the Bismarck Archipelago and the Admiralty Islands.

There is a fairly considerable literature extending over a long period. Fabricius
in 1805 appears to have been the first author to name a Tabanid from the northern
part of the Australian Region, then follow Macquart (1838), Francis Walker (1848 to
1866) based on the Wallace Collections, Doleschall (1858), van der Wulp (1868),
Megnin (1878), Bigot and Roeder (1892), Ricardo (1913), de Meijere (1915 and 1917),
and finally Schuurmans Stekhoven (1924 and 1926).

I desire to extend my very sincere thanks to the late Dr. F. W. Edwards and
Mr. H. Oldroyd for sending to me on loan the undetermined specimens of Tabanidae
contained in the British Museum (Nat. Hist.) from the Territory of New Guinea and
Papua, in the main collected by Miss L. E. Cheesman, also to Mr. N. D. Riley, Keeper,
Department of Entomology, for many courtesies, and finally, to Dr. John Smart of the
same department for checking the series of drawings made by Mr. Arthur Smith from
the types and/or named specimens in the British Museum (Nat. Hist.).

The following descriptions are based on those of Schuurmans Stekhoven.

CHRYSOPS ALBICINCTA van der Wulp. Plate x; Text-fig. 1.

Tijdschr. Ent., xi, 1868, 103, pl. 3, fig. 6; Osten Sacken, Ann. Mus. Stor. nat. Genova, xvi,
1868, 418; Ricardo, Rec. Ind. Mus., iv, 1911, 377; Nova Guinea, ix, 1914, 406;
Schuurmans Stekhoven, Treubia, vi, Suppl. 1926, 22.
©. Head: Front broad, black, denuded; callus transverse, nearly reaching the eyes.

broader than long, shining black to dull red-brown, half-moon shaped, grooved, somewhat
prominent between the antennae; ocellar plate rectangular, shining black, nearly
reaching eyes, covered with black hairs; cheeks havanna-brown, shining, pubescence
brownish-black, with some white hairs close to the base of the palpi; beard white:
clypeus shiny, ochraceous-brown, with sparse brownish hairs; black or reddish-brown
patches on the cheeks at the upper edges of the facial callus; facial callus with a
shallow indentation at its upper border in the middle; antennae: segments almost equal
in size, from two to two and a half times as long as the head, first segment with annuli
faintly indicated, ochraceous-yellow, darker toward the slightly swollen apex which is
pale havanna-brown, with sparse black pubescence, second segment with five annuli,
in some lights apparently six, dark havanna-brown, black haired, third segment bare,
first annulation broadened near the base, with four faint annulations, shining black at
base, rest dull brown, the four apical annuli deep velvety black; palpi short, reddish-
brown, with black hairs.

BY FRANK H. TAYLOR. 329

Fig. 1.—Chrysops albicincta van der Wulp. A, Front showing frontal callus. B, Head in
profile with antenna. C, Wing showing pattern. (A and B same magnification. )

Thorax black, with scanty black hairs on disc, the posterior border of the scutum
with a narrow line of golden-yellow hairs, prolonged along the lateral borders of the
scutum as a falciform tuft; pteropleuron with golden-yellow hairs; scutellum black;
pleurae shining black to blackish-brown, with black hairs; lobe of the mesopleuron
distinctly tinged red-brown; the shoulders sparsely covered with black hairs.

Wing as illustrated, veins yellow-brown, except on the markings where they are
havanna-brown; halteres red-brown.

Legs: Fore-coxa black-brown, covered with brownish hairs, trochanter black-brown,
femur yellow to reddish-brown, black haired, tibia and tarsus olive-brown, black haired;
mid-coxa and trochanter black-brown, black haired, femur tawny, black haired, tibia
havanna-brown, black haired, swollen, tarsus straw-yellow, black haired; hind-coxa black
with velvety brown apical border, white haired, trochanter red-brown, femur reddish-
brown to black, sparsely covered with yellowish-brown hairs, tibia as in mid-leg, less
stout, black haired.

Abdomen black, first segment chocolate-brown, except two small white median spots,
nearly touching the scutellum, and separated by a very narrow longitudinal stripe,
posterior border black, second segment broad, chocolate-brown, with a yellowish-white
ring basally, surrounding the entire segment, widening to the lateral borders of the
tergite; ventrally it becomes more brownish-white and covers also a part of the first
segment. Some specimens show a brownish-white longitudinal stripe in the middle
of the black band of the second segment. Dorsum elsewhere black, venter blackish-
brown to black, covered with black hairs except on the pale spots where they are white.

Length: 7-5 mm.

Habitat: Salawati Island (type); Netherlands New Guinea, numerous localities.

The above description is from Schuurmans Stekhoven. He notes that a female
specimen from South New Guinea has the front grey tomentose and grey haired.
Ricardo states that the wing markings of this species are typical and distinct from
any Oriental species.

Ricardo describes the wing as having a broad transverse band with a hyaline sinus
in the fifth posterior cell, and an apical spot, the apex beyond being grey, with the
dark colouring in fore border and in the basal cells reaching to about two-thirds of
the length in the first one, and half the length in the second basal cell. Schuurmans
Stekhoven’s specimens mostly had smaller brown spots in the basal cells with the grey
colouring absent in some of them.

330 DIPTERA OF THE TERRITORY OF NEW GUINEA. XIII,

The illustrations in Text-figure 1 were made by Mr. Arthur Smith from a specimen
in the British Museum (Nat. Hist.) determined either by the late Major EH. EH. Austen
or Miss Ricardo.

The location of type is unknown to me. Specimens in the following institutions:
Natura Artis Magistra Museum, Holland, Buitenzorg Zoological Museum, N.EH.I., Leyden
Museum of Natural History, Holland, and the British Museum (Nat. Hist.).

Fig. 2.—Chrysops atrivittata Sch. Stekh. a, Front of head. b, Antenna. After Schuurmans
Stekhoven.

CHRYSOPS ATRIVITTATA Sch. Stekh. Plate x; Text-fig. 2.
Treubia, v, 1924, 317, pl. v; op. cit., vi, 1926, Suppl., 26, pl. 1, fig. 4; text-fig. 8.

9. Head: Eyes unicolourous mauve, when moistened they show three distinct dark
patches; front broad, nearly same width throughout, very slightly narrower at vertex,
yellowish-grey tomentose sooty in some places; callus dark black-brown in type, not
reaching the eyes, varying from red-brown to black-brown in other specimens, shining,
convex to the posterior end, pointed between the antennae, bearing a shallow flat groove,
which divides the callus into two unequal halves, surrounded by a whitish-grey margin;
ocellar triangle with rounded sides, and with a moderately dense covering of long
yellow-brown hairs, similar hairs in the space between the eyes and the ocellar triangle;
excavation behind the callus is white haired; antennae: first segment yellow-brown,
darker bismarck-brown toward apex, indistinctly 6-7 ringed, barely visible at base, |
distinct toward apex, black-brown hairs more or less arranged as festoons, corresponding
with the rings, short and appressed basally, longer and denser intermixed with some
white hairs apically, especially on the three apical rings, apex of first segment with
short yellow pubescence in some specimens; second segment dark olive-brown with a
black tint nearly as long as the first segment, slightly broader, five ringed, sometimes
four or six, black hairs as in the first segment, longer on the distal lateral border
than on the proximal border, with some mixed white hairs; third segment broader
than the second, shorter than the first and second but longer individually, with five
distinct rings, velvety-brown basally, almost black-brown apically, basal annulation
longer than those apically together, fifth annulation is nearly as long as the third and
fourth, apically blunt; palpi with first segment brownish-black, very short triangular,
covered with long black hairs, second ochraceous yellow-brown, at least four times as
long as the first, ending in a blunt point, pubescence mainly yellow-brown with a few
black hairs; proboscis with long labella with mixed long and short dark brown hairs
ventrally, reddish-ochre-brown dorsally, ventrally more blackish-brown; cheeks and face
yellowish-pollinose except on the two shining dark olive-brown streaks on the lateral
border of the clypeus, median part of clypeus bright ochraceous, pubescence long, yellow,
beard bright brownish-yellow-white, cheeks elsewhere nearly devoid of long hair.

Thorax with a median dull rusty-brown stripe, latter darkened laterally, sides
reddish-brown to black with a violet hue, pubescence short and sparse varying from

BY FRANK H. TAYLOR. 331

bright to dark brownish-yellow, posterior border of scutum with a broad line of golden
hairs ending laterally in a triangular tuft of golden-yellow hairs; pteropleuron golden
haired, pleurae olive-brown with golden-yellow hairs, mesosternum blackish-brown
eovered with blackish-brown pubescence; scutellum dark reddish-brown with a median
transverse very narrow black spot fringed with bright yellow-brown short pubescence,
halteres with a dull bismarck-brown stem and dark blackish-brown knob.

Wings as illustrated in Plate x.

Legs: Coxa of fore-leg whitish tomentose, shining black basally, the middle swollen,
covered with black hairs, distal lateral border with yellowish-brown hairs, proximal
lateral border bright oak-brown, with a reddish hue, with brownish-black hairs mixed
with some white ones, femur rusty brown, darker than trochanter, densely covered
with black hairs dorsally, lateral distal border with a row of erect, long, black hairs
decreasing in length to the base, tibia dilated slightly over the middle, oak-brown,
densely covered with short appressed hairs, tarsals one to four same colour as tibia,
fifth tarsal black, some erect, relatively long, dirty white hairs on its hind border,
elsewhere black haired, mid-leg with coxa and trochanter both short, dark brown, hairs
scanty, femur same colour as in fore-leg, knee yellow, dorsal surface black haired,
sides with a fringe of black hairs increasing in length toward the apex, tibia black-
reddish-brown, slightly dilated, densely covered with black hairs, ventral surface with
black hairs, mixed with some white ones basally and apically, first tarsal very long,
nearly as long as two to five together, mainly black haired, a number of short appressed
yellow hairs on basal third, lateral margins yellow haired, tarsals two to four yellow
haired with an apical ring of black hairs, some black hairs on the proximal lateral
border, fifth tarsal black, with five long white hairs apically; posterior leg: coxa swollen
with yellowish-brown hairs, mixed with some black ones, trochanter short, narrow,
femur dark yellow-brown densely covered with black hairs, some longer yellowish-white
ones being scattered over the whole surface, tibia shining dark brown, basally swollen,
black haired, attenuated, long, longer than two to five together, brownish-yellow, black
haired dorsally with some intermixed yellowish-white hairs laterally, ventral surface
mainly yellowish-white haired, apical ring black haired, two to four yellowish-brown,
black haired, ventral surface with a black haired apical band, elsewhere yellow haired,
fifth black on dorsal half, ventral surface yellowish-brown with the apical ring black
haired.

Abdomen with four longitudinal stripes, joined on the second and third segments,
first segment hazel-brown with two black spots, second yellow, laterally black with a
small yellow spot on each side, segment three with four longitudinal black stripes
connected in pairs posteriorly; the enclosed areas are yellowish-brown, all black stripes
interrupted at the borders of the segments, median stripes broader than the laterals,
remaining segments similar, colour of segments increasingly dark toward the apex,
first segment with brownish-yellow hair, spots black haired, second with yellowish-white,
markings black haired, third mainly black haired, with some yellowish-white ones,
remaining segments black haired except for the apical and basal yellow fringe; venter
yellowish-white haired on first two segments, remainder reddish-brown with black
hairs, some mixed yellow hairs on the seventh.

Length: 9-2 to 11:0 mm.

Habitat: Buru Island, Rana (type), Koentoeroen Marsh.

Related to Chrysops designata Ricardo but may be readily distinguished.

Type and other specimens in the Natura Artis Magistra Museum, Holland.

CHRYSOPS SIGNIFER Walker. Plate x.

Proc. Linn. Soc. Lond., v, 1861, 276; Osten Sacken, Berl. ent. Z., xxxvi, 1882, 97; Ricardo,
Ann. Mag. nat. Hist., (7) ix, 1902, 376; Schuurmans Stekhoven, Treuwbia, v, 1924,
324; op. cit., vi, 1926, Suppl., 43.

6. Head: Eyes with large facettes on the upper half and are mainly yellow-brown,
lower half black, facettes small; ocellar plate with small erect hairs; antennae rising
from piceous pits, first segment dorsally coffee-brown at base, remainder blackish, black

FF

332 DIPTERA OF THE TERRITORY OF NEW GUINEA. XIII,

haired, indistinctly eight ringed, second segment black with eight indistinct rings, first
five narrow, last three broad, swollen at the borders, first and second segments equal
in length, third segment bare, black, slightly longer than the second, incrassate at base,
five distinct annulations, the first as long as two to five inclusive, apex bluntly rounded;
clypeus shining black, with a narrow median yellow-grey tomentose stripe; subcallus
yellow tomentose; cheeks golden-yellow with similarly coloured hairs, beard the same
colour; proboscis piceous, labella with short brown-black hairs; palpi black, very short.

Thorax: Scutum black, median line with short golden-yellow hairs, posterior border
with dense golden-yellow hairs, lateral tufts golden-yellow, shaped as a ? mark;
scutellum piceous.

Wings as illustrated in Plate x.

Legs: Fore-leg, coxa piceous with erect black hairs, femur dark chocolate-brown,
black haired, lateral border with short hairs, distally the hairs are long and erect, tibia
somewhat swollen basally, chocolate-brown at base, rest black, hairs black, tarsals
black, black haired, mid-leg with femur and tibia similar, black haired, first tarsal
with a row of black hairs along the proximal lateral border, remaining tarsi whitish
with a brownish hue, apices with black hairs, ventral surface with dense black hairs,
hind-leg with femur and tibia black covered with long, erect hairs, first tarsus dirty
yellowish-brown, black haired, tarsi two to four similar, fifth tarsus of all legs black.

Abdomen: First segment posteriorly with a narrow black transverse band touching
the anterior border of the second segment in the median line and curving backwards
laterally to the middle of the lateral margin, first segment yellow elsewhere, with
brownish-yellow pubescence, second segment clear yellow, yellowish-white haired with
a crescent shaped blackish-brown band apically not reaching the lateral border of the
segment, third with a broad concave blackish-brown hoof-shaped band drawn out to and
touching the crescent-shaped pattern of the second, lateral borders of this segment
yellowish-white with yellowish-white hairs, both segments with an enclosed curved
triangular yellow spot with sharp basal angles, all dark markings with blackish-
brown hairs, segments four to seven terracotta-reddish-brown, fourth and fifth with
blackish median indistinct spots, pubescence brown intermixed with yellow hairs;
venter: fourth segment with a brownish-red almost black transverse band, a blackish
stripe connecting this band with the dorsal marking on the third segment, seventh
entirely black haired, fifth and sixth oakleaf reddish-brown, pubescence the same, first
three segments yellow-haired.

Length: 10-5 mm.

Habitat: Batjan Island (type); Buru Island.

Closely related to Chrysops cincta Bigot.

Type, ¢, in the British Museum (Nat. Hist.).

EXPLANATION OF PLATE X.

Chrysops albicincta van der Wulp. (Top.)
Chrysops atrivittata Sch. Stekh. (Middle.)
Chrysops signifer Walker. (Bottom.)

(All illustrations after Schuurmans Stekhoven.)

STUDIES IN THE METABOLISM OF APPLES.
VI. PRELIMINARY INVESTIGATIONS ON THE RESPIRATION OF SLICED APPLE TISSUE.

By Frances M. V. Hackney, M.Sc., Linnean Macleay Fellow of the Society in
Plant Physiology.
(From the Department of Botany, University of Sydney.)

(Three Text-figures. )

[Read 28th November, 1945.]

INTRODUCTION.

Previous papers of this series (Hackney, 19438, 1944) were concerned with the
respiratory trends of Granny Smith apples at various stages of development and of
maturity.in store. It was shown that the causes underlying these trends were not the
same at all stages of development and maturity. In order to continue the investigations
it seemed advisable to develop a technique for the study of the respiratory behaviour
of sliced apple tissue immersed in distilled water and other media. By supplying
various respiratory inhibitors and possible respiratory substrates to the sliced tissue,
information might be obtained about the nature and relative importance of the reactions
involved in respiration at various stages cf development and maturity. Similar methods
have been applied to the study of the respiration of carrot tissue by Turner (1938) and
others, of potato tissue by Boswell and Whiting (1938) and others, and of beet tissue
by Bennet-Clark and Bexon (1943). Apple tissue presents peculiar difficulties not met
with in work on other storage tissues, and the technique of other investigators has.
been modified accordingly. The present paper outlines this technique, and includes
results of preliminary investigations on the respiration rates of tissues from various
parts of the fruit.

MATERIALS.

Most of the experiments were carried out on Granny Smith apples of the 1944
season; some experiments were carried out on fruits of the 1945 season. The apples
used were part of the normal commercial (mid-April) picking from a selected orchard
at Orange, New South Wales. They were placed in cool store (1°C.) within a few days
of harvesting. Batches of fruit were removed from store as required and placed in a
room maintained at 21°C. in 1944. Their subsequent behaviour was observed at this
temperature. During 1945 the experimental temperature was 25°C.; facilities for
keeping the fruits at the lower temperature were not available in 1945.

PREPARATION OF TISSUES.

In the preparation of the cut apple tissue, skin and flesh were always treated
separately. The skin was sliced off very thinly, using a razor blade. The average
thickness of the tissue classified as skin was about 0:15 mm. The strips of skin were cut
into pieces about 6 mm. square. In the early experiments the cut skin was subsequently
washed in aerated distilled water for about 20 minutes, and the surplus water was
removed by carefully drying the tissue between filter papers. The skin was then
divided into replicate sets, usually of 1 gm. fresh weight each. In the later experiments
the tissue was placed in the respiration vessels without preliminary washing (see
later section).

(Jt)
wo
NSS

STUDIES IN THE METABOLISM OF APPLES. VI,

Dises of flesh were prepared in the following way: Cylinders of tissue were cut
from the fruit by means of a sharp cork-borer 0-9 cm. in diameter. Portions obviously
bruised were discarded, and the remainder of each cylinder was cut into slices of the
desired thickness (usually 1 mm.) by means of a razor blade. It was shown, by
experiments to be described in a later section, that the small variations in thickness,
due to cutting the slices by hand, were not likely to cause inaccuracy in the measure-
ment of respiration rate. In the early experiments the flesh discs were washed and
dried in the same way as the skin slices. Replicate sets of flesh tissue were usually
of 2 gm. fresh weight (weighed after washing). In later experiments the discs were
not washed before being placed in the respiration vessels.

In experiments where a large number of replicate sets was required, tissue slices
were cut from a number of similar apples and randomized during washing. There was
no significant difference in respiratory behaviour between tissue sets taken from such
randomized lots. In ‘experiments where a small number of replicates was required,
randomized sets of tissue from a single apple were used. Although there was
considerable variability in respiration rate between tissue sets from different apples,
there was good agreement between the respiration rates of tissue sets from the
same apple.

It was not considered necessary to prepare the material under sterile conditions,
as bacterial growth did not occur during the short duration of the experiments (cf.,
Turner, 1938).

The Warburg manometric technique was employed in the determination of the
rates of oxygen uptake and carbon dioxide output of the tissue slices. Each of the
Warburg vessels had two side-arms as well as a small inner receptacle within the
central cavity (see Dixon, 1943, p. 49). Two gm. flesh tissue or 1 gm. skin were
suspended in the experimental medium in the central cavity of the vessel.

In determinations of the rate of oxygen uptake, the carbon dioxide given off by the
tissue was absorbed by means of a small piece of Whatman’s No. 40 (starch-free) filter
paper soaked with 0-3 c.c. of normal potassium hydroxide; absorption was found to
be equally efficient whether the filter paper was placed in the inner receptacle or
in one of the side-arms. The vessels were immersed in a water-bath maintained at
the experimental temperature (21°C. in 1944, 25°C. in 1945), and shaken at approximately
110 complete oscillations per minute.

In experiments where it was necessary to determine the respiration rate of the
whole apple before cutting, the Pettenkofer method was adopted. These experiments
were carried out in a room maintained at the desired temperature.

RESPIRATORY BEHAVIOUR AFTER CUTTING.

It is well known that the damaging of living tissue due to wounding frequently
brings about an increase in the respiration rate. This increase may be due to a variety
of causes.

Turner (1940) found that carrot tissue must be washed in aerated distilled water
for 200 hours or more in order to minimize the preliminary period of abnormally high
respiration. It was not practicable to follow this procedure with apple tissue, as it
lost nearly all its respiratory activity when washed for periods longer than about
30 hours. Flesh tissue washed in aerated phosphate buffer (pH 5:9) did not survive
longer than similar tissue in distilled water.

In the early work apple tissue was washed for 20 minutes (see later section)
and suspended in distilled water for the duration of the experiments. The respiration
rate was approximately the same whether the volume of water used was 1 ¢.¢., 3 ¢.¢.,
or 5 c.c. per gm. of flesh. For purposes of convenience 4:5 ¢c.c. was the volume generally
used.

Shape of the Respiration-Time Curve.—Although variations frequently occurred in
the magnitude of the respiration rate, the shape of the respiration-time curve for
flesh discs in distilled water was always the same at the experimental temperatures
considered. Soon after cutting, the rates of oxygen uptake and carbon dioxide output

oe ee eA ee

BY FRANCES M. V. HACKNEY. 335

were comparatively high (23-30 mm.*/gm. initial wt./hr.). During the first 2 or 3
hours after cutting, the respiration rate decreased, subsequently reaching an almost
steady level, which was maintained with very little change during the remainder of
the period of observation (a further 4-6 hours). A typical curve for oxygen uptake of
flesh tissue is shown in Figure 1. The shape of the respiration-time curve and the
magnitude of the respiration rate were the same in phosphate buffer as in distilled
water. The shape of the respiration-time curve is discussed further in a later paragraph.
The respiration-time curve for skin did not show the preliminary fall; it either
remained approximately steady or rose slightly during the period of observation.

30

[ mami: Frat hr]

OXYGEN UPTAKE
©)

1 2 3 4 S)
HOURS AFTER CUTTING

Fig. 1.—Typical respiration-time curve for flesh of Granny Smith apple at 21°C. Time of
preliminary washing was 20 minutes. The entire lines denote respiration rate based on the
weight of the sample at the time of setting up in the Warburg vessel; the dotted lines denote
respiration rate corrected to the true weight of the sample at the time of each observation
(see section on effect of immersion in distilled water).

Effect of Immersion of Tissue in Distilled Water.—It was not until many
investigations had been carried out using tissue washed in distilled water that it
was discovered that considerable losses occurred in both fresh and dry weights when
apple flesh was surrounded by distilled water. This behaviour was surprising, as no
comparable loss in weight occurs in carrot, potato or beet tissue.

In order to determine the extent of loss in dry weight during washing, samples of
2 gm. fresh weight each were taken from a large batch of flesh discs after various periods
of washing. The dry weights of these samples were determined. It was found that the
ratio dry wt./fresh wt. decreased as the duration of the washing period increased (see
Table 1).

In a second experiment the changes in fresh weight of a single sample of flesh
discs were observed after various periods of washing. It was found that fresh weight
decreased considerably as the duration of the period of washing increased (see Table 2).
The dry weight of the sample after each period of washing was calculated from the
ratios dry wt./fresh wt. in Table 1.

From these and similar experiments it was evident that during 2 hours in distilled
water the apple flesh lost considerable proportions of both fresh and dry weight. The

336 STUDIES IN THE METABOLISM OF APPLES. VI,

rates of loss were greatest during the first 20 minutes of washing and decreased as
the length of the washing period increased.

TABLE 1.
Changes in Dry Wt./Fresh Wt. of Flesh Tissue after Various Periods of Washing in
Distilled Water.

Period of Fresh Wt. Dry Wt. Dry Wt./Fresh Wt.
Washing. (gm.). (gm.). %)-

0) 2 0-289 14-45

5 min. 53 0-219 10°95
20s ss 0-180 9-00
GON ee 0-146 7-40
20 Sy 0-118 6°50

Unlike the flesh, the skin showed an increase (25%) in fresh weight due to uptake
of water during the first 20 minutes of washing, after which further washing produced
no change.

TABLE 2.
Weights of One Sample of Flesh Tissue after Various Periods of
Washing.

Period of Fresh Wt. Dry Wt.
Washing. (Observed.) (Calculated.)
0 4-00 gm. 0:58 gm.

5 min aodly A Mes) 55
20. Moy Oi 217 eames
GOMess Zid Ones cil) 52
120 ee oil 0-14
16 hr IESENO Ss =

Respiration Rates of Tissue Sets washed in Aerated Distilled Water for Various
Periods —Two experiments were carried out to determine the respiration rates of
comparable sets of mature apple discs which had been washed in aerated distilled water
for various periods. Table 3 shows the results of one of these experiments.

TABLE 3.
Respiration Rate of Flesh of Mature Apple after Various Periods of Washing
(25—-26.iv.44). Material Weighed after Washing, before being Set Up in
Warburg Vessels.

Respiration Rate (mm.?/gm. initial wt./hr.).

Period of
Washing. Replicates.
A. B. C.
0 16-5 16-0 15-5
5 min. 8-0 = aa
15-20 ,, 6-0 — =
120 ,, 5-0 zi =
43 hr. 4-0 = —
18); 5-0 6-0 ae
Zones, 5°5 4-0 =

Discs which were placed in the Warburg vessels with no preliminary washing
settled down to relatively high respiration rates; discs which had been washed attained
lower respiration rates than those which had not been washed; increasing the washing
period beyond 15-20 minutes did not appear to decrease the respiration rate further.
The results of the above experiments led to the adoption of a standard 20 minutes
washing period for tissue used in the earlier investigations.

It is highly probable that the decrease in respiration rate of flesh with increased
time of washing in distilled water (see Table 3) was due in some measure to the
extensive loss of water and dry material which took place during the first 20 minutes

BY FRANCES M. V. HACKNEY. 337

of washing. In samples which had been washed for very short periods, leaching of
solutes from the cells was still proceeding at a relatively high rate when the discs
were transferred to the Warburg vessels (cf., Table 2), whereas in samples which had
been washed for longer periods leaching was slow. In the former samples the amounts
of solutes leached from the cells might have been sufficient to alter the concentration
of the external solution in the Warburg vessels to such an extent as to prevent further
leaching; this might have resulted in the maintenance of the respiration rates of
these samples at relatively high levels. The fact that washing for more than 20 minutes
did not bring about a further decrease in the respiration rate was probably due to the
comparative smallness of the change in the dry wt./fresh wt. ratio which occurred
after 20 minutes (see Table 2). No significant difference was observed in the respiration
rates of tissue sets washed for various periods (from 5 min. to 5 hr.) in calcium chloride
solution isotonic with the cell sap (see following section).

Effects of Immersion of Flesh in Solutions isotonic with Cell Sap—When the effects
of contact of flesh cells with distilled water were observed microscopically, it was
found that many of the cells absorbed water so rapidly that disorganization of the
contents occurred. In view.of this, several experiments were carried out to determine
the effects of immersion of discs of flesh in solutions of glucose, potassium nitrate and
calcium chloride at various concentrations. It was found that loss in fresh weight
after 2 hours immersion was much less in low concentrations of these substances than
in distilled water. When the osmotic pressure of the surrounding solution was between
11 and 15 atmospheres the loss in fresh weight was very small or negligible after
2 hours. Losses in percentage dry wt. per unit fresh wt. were very small or negligible
after various periods of washing, from 5 min. to 2 hr., in 0:25M calcium chloride
(O.P. = 12:3 at.). The suction pressure of the fiesh cells was between 11 and 15
atmospheres. When the osmotic pressure of the surrounding solution was greater than
15 atmospheres, the tissue lost weight owing to plasmolysis; when the osmotic pressure
of the surrounding solution was less than 11 atmospheres the flesh tissue lost weight
Owing to rupture of the cells following rapid absorption of water from the surrounding
solution. The suction pressure of the skin cells was of the same order as that of the
- flesh cells. When the osmotic pressure of the surrounding solution was lower than 11
atmospheres, the skin tissue gained in weight owing to absorption of water; the cells
of the skin were not disorganized by immersion in solutions of low osmotic pressure.

Several experiments were carried out comparing the respiration rates of samples
of flesh (unrinsed) in distilled water with those of similar samples in 0-25M calcium
chloride. It was found that the shape of the respiration-time curve of tissue in calcium
chloride was similar to that of tissue in distilled water, except that the initial fall was
not as steep in calcium chloride as in water; the final respiration rate per unit initial
weight was almost twice as high in calcium chloride as in water. The tissue sets were
weighed at the close of the experiments. When the respiration rates were calculated
per unit final weight it was found that the respiration rate of tissue in calcium chloride
was only 15-20% higher than that of tissue in water.

There was no significant difference between the respiration rate of skin in 0-25M
calcium chloride and that of skin in water.

In view of the fact that the respiration rate per unit true weight of flesh in water
was so close to that of flesh in calcium chloride, it seemed that the main effect of
immersion in the salt solution was the prevention of the occurrence of changes in
weight during the experiments. In some tissues (e.g., carrot discs, Robertson, 1941)
addition of chloride to the external medium results in chloride accumulation and
increased respiration. This effect does not appear to be of major importance in apple
flesh tissue, and there is complete absence of ‘salt effect’ in the skin.

In spite of its effects on weight loss, substitution of calcium chloride for water
during washing did not greatly prolong the life of the cut flesh. The respiration rate
of discs washed overnight in calcium chloride was no greater than that of discs washed
overnight in distilled water.

338 STUDIES IN THE METABOLISM OF APPLES. VI,

The steepness of the initial fall in the respiration-time curve for flesh tissue in
water was probably due in part to the loss in true weight which occurred after
transference to the Warburg vessel. Respiration rate estimated on the basis of initial
fresh weight would clearly have been erroneous, since fresh weight continued to
decrease in the Warburg vessels. Making allowance for this decrease in weight and
assuming that a proportionate number of cells ceased to respire, the respiration-time
curve still showed a slight fall during the first 2 hours after setting up (see Figure 1).
This fall might have been partly due to some of the cells ceasing to respire, as a
result of disorganization, but not immediately losing their contents, and therefore
contributing a small part of the weight of the tissue. The fact that there was still
a slight initial fall in respiration rate when the discs were immersed in calcium
chloride instead of in water indicated that some factor other than rapid disorganization
of the cells was involved in the determination of the shape of the respiration-time curve.

Experiments with Tissue Slices of Different Thicknesses.—It was considered that
if any fraction of the observed respiration rate of the flesh tissue had been due to
the presence of cut cells, this fraction would have been increased in proportion to the
increase of cut cells relative to uncut cells in a given weight of tissue. Several
experiments were therefore carried out using tissue slices of various thicknesses. If
an increase in the percentage of cut cells had accelerated the respiration rate (cf., work
on other tissues) it might have been expected that this rate would have been highest
in the thinnest discs. It was found, however, that whether the medium was water or
calcium chloride, there was no significant difference in rate of oxygen uptake per unit
fresh weight between tissue sets composed of discs 2 mm. thick and those composed of
discs 1 mm. thick, and that in water, the rate of oxygen uptake of discs 0-5 mm. thick
was slightly lower than that of discs 1 mm. thick. This difference could be largely
accounted for if the damaged cells had lost most of their contents during washing and
were therefore not contributing to the respiration. Table 4 shows the results of a
typical experiment with discs of different thicknesses. From these data it is probable
that no significant percentage of the observed final respiration rate of the tissue was
contributed by cut cells.

TABLE 4.
Respiration Rates of Flesh Discs of Various Thicknesses. The Discs
in Calcium Chloride were not from the Same Apple as Those in Water.

Respiration Rate (mm.?/gm. fr. wt./hr.).

Thickness of Discs

(mm.). In Calcium
In Water. Chloride.
2-0 12:0 10°5
1-0 ile) 9-0
0:5 8-0 8:9

The results presented in the following pages for tissue in water have been corrected
to the true weight of the tissue three hours after immersion (i.e., when most of the loss
in weight had taken place).

RESPIRATION OF TISSUES FROM DIFFERENT REGIONS OF THE APPLE.

Respiration of Tissue from Different Depths in the Flesh—In order to discover
whether there was a gradient in respiration rate between the skin, the cells immediately
under the skin and the cells in the deeper parts of the flesh, the rate of oxygen uptake
was determined under similar conditions for each of the following regions of the
same apple:

(1) skin;

(2) flesh tissue immediately under the skin, to a depth of approximately 3 mm.;
(3) tissue from 4 to 8 mm. under skin;

(4) tissue from 8 to 14 mm. under skin;

(5) flesh tissue more than 14 mm. under skin.

-o-—

aw a “oe

Qe...

BY FRANCES M. V. HACKNEY. 339

There was a very marked difference between the respiration rates of skin and
flesh tissue; there was no significant difference between the respiration rates of flesh
tissue from any of the regions considered.

In a second series of experiments the respiration rate of flesh tissue containing
vascular strands was compared with that of flesh tissue containing no vascular strands.
There was no significant difference in respiration rate between these two types of flesh
tissue.

Respiration of Seeds—The weight of seeds per fruit was no greater than 0:6 gm.
It was observed that the respiration rate in air of seeds freshly removed from the
fruit was of the same order per unit fresh weight as that of the flesh, or less. As the
seeds constitute such a small percentage of the total weight of the fruit, their
contribution to the total respiration of an uncut apple is very small.

Respiration of Skin and Flesh.—The respiration rate of the skin (expressed in
mm.*/gm. fr. wt./hr.) was found to be at least ten times as great (frequently 20 times
as great) as that of flesh from the same apple expressed in the same units. This was
true for Delicious and Jonathan apples as well as for Granny Smiths. When the
respiration rates were expressed in mm.*/gm. dry wt./hr., that of the skin was 4-5 times
as great as that of the flesh. Table 5 gives data for two typical immature fruits. For
Granny Smiths the ratio skin respiration/flesh respiration varied between approximately
the same limits in mature fruits as in immature fruits.

. TABLE 5.
Respiration Rates of Skin and Flesh in Immature Granny Smith Apples.

Oxygen Uptake.
Qum.?/gm. fresh wt:/hr.).

Date of Oxygen Uptake.
Picking. Tissue. Value based (mm.3/gm.
on Weight Corrected dry wt./hr.)
at Time of to Approx.
Setting Up. Final Wt.
5. 1.45 Flesh .. ee 8-2 10-3 117
Skintaaer. Te 150-0 _— 625
19.ii1.45 Flesh .. ss 8-0 10-0 145
Skin =- aS 140-0 — 635

From results of the type shown in Table 5, it has been calculated that the skin
contributes a considerable proportion of the total respiration of the apple. In a typical
fruit, removed from store on 10.vii.45, the respiration of the skin was 150 mm.’/gm.
fr. wt./hr. and that of the flesh was 15 mm.*/gm. fr. wt./hr. The fruit weighed 140 gm.
fr. wt.; the skin constituted 4:3 gm., the remaining 135-7 gm. being composed of flesh,
seeds, etc. Thus the total respiration of the skin was 645 mm.*/fruit/hr., and the
total respiration of the flesh was 2,040 mm.*/fruit/hr. The total respiration of the skin
plus flesh was therefore 2,685 mm./fruit/hr. In this fruit, the skin was apparently
responsible for a little less than one-third of the total respiration. The proportion of the
total respiration contributed by the skin varied in individual fruits. It was nearly
always of the order of one-third, but in rare cases values as high as one-half were
recorded.

The Relation between Skin Respiration and Flesh Respiration throughout Post-
Storage Life—The magnitudes of the respiration rates of skin and flesh varied among
individual fruits. It has been shown (Hackney, 1943) that the respiration rate of the
whole fruit decreases with time after removal from store. The data obtained during
1944 and 1945 show that this decrease in total respiration is probably made up of
decreases in both the respiration of the skin and that of the flesh; the ratio skin
respiration/flesh respiration probably remains constant throughout the post-storage life
of each individual fruit.

In 1944 a strong positive correlation was observed between the respiration rates in
water of skin and flesh from the same fruit. The regression line was Y = 6:136X + 49-58,

340 STUDIES IN THE METABOLISM OF APPLES. VI,

where Y = respiration of skin and X = respiration of flesh, and the regression coefficient
was highly significant (P < 0-001). A similar correlation was indicated in 1945, but
the data were not numerous enough to merit statistical analysis.

The Anatomy of the Skin and Flesh.—The cells of the skin are much smaller and
more tightly packed than those of the flesh. The percentage dry weight per unit
fresh weight is 4-5 times as great for skin as for flesh. Figures 2 and 3 show the
cellular detail of mature apple tissue. Immediately below the cuticle is a layer of
irregular polygonal cells of small diameter (about 20u), each containing a comparatively
small vacuole and dense cytoplasm, in which a large number of chloroplasts is visible.
Figure 2 shows a surface view of the epidermal layer. In cross-section (Figure 3) it
appears to be about as thick as the cuticle. Immediately under the epidermis are about

Fig. 2.—Surface view of the epidermal cells of the Granny Smith apple, showing dense
cytoplasmic contents with many chloroplasts (x 500).

three layers of flattened cells about twice as large in diameter as the epidermal cells,
but slightly smaller in radial thickness. These cells contain relatively dense cytoplasm
with many chloroplasts. The 3-4 layers immediately under this subepidermal region
show a gradual transition from the small, densely packed, chlorophyllous cells of the
subepidermis to the large, loosely packed, non-chlorophyllous cortical cells of which the
greater part of the apple is composed. The tissue used as ‘skin’ in the experiments
described here includes only the first 7-8 layers of cells beneath the cuticle.

The flesh tissue is composed of very loosely arranged cells, having between 10 and
20 times the volume of the cells comprising the skin. Hach of these cells contains a
thin layer of cytoplasm and a large vacuole. The intercellular spaces are large.

The large volume of the individual cells and the relative thinness of their cytoplasmic
layers are probably the cause of the disorganization which occurs on immersion of the
flesh in distilled water or in solutions of low osmotic pressure. The cytoplasmic layers
of some of the cells are probably ruptured by the sudden expansion of the very large
vacuoles consequent upon the absorption of water from the external solution.

The cellular details of Jonathan and Delicious apples were found to be very similar
to those of the Granny Smith. The cuticle of the Delicious apple is thinner than those
of the Granny Smith and Jonathan. Tetley (1930, 1931) described the anatomy of
several varieties of apple grown in England. The Granny Smith appears to be more
similar to the Bramley’s Seedling than to the other varieties examined by Tetley, but
there are differences in anatomical detail between the two varieties; the epidermal cells

of the Granny Smith are much smaller than those of the Bramley’s Seedling (Tetley,
A931).

ny

BY FRANCES M. Vv. HACKNEY. 341

From a microscopic study of the structure of the skin and flesh, it is clear that
the flesh contains a large proportion of vacuolar material, which is not actively
concerned in respiration, and a small proportion of cytoplasm, whereas the skin contains
a small proportion of vacuolar material and a large proportion of actively respiring
cytoplasmic material. A rough estimation was made of the numbers of cells in 1 gramme
of flesh and skin:

Volume of 1 gm. flesh = 11 x 10? mm.* (approx.).

Volume of 1 gm. skin = 9 x 10? mm. (approx.).

Volume of a flesh cell was of the order of 2 x 10-* mm.

Therefore the number of cells in 1 gm. flesh was of the order of 5:5 x 10°.

Fig. 3.—Transverse section of skin and flesh of Granny Smith apple. ARC uticler:
B, Ep‘dermis; C, Subepidermis; D, Transition zone; EH, Flesh (x 500).

342 STUDIES IN THE METABOLISM OF APPLES. VI,

The skin was made up of cuticle plus epidermis plus subepidermis plus transition
tissue. From sections of the type shown in Figure 3 the following calculations were
made:

Volume of cuticle in 1 gm. skin = 100 mm: (approx.).

Volume of epidermal tissue in 1 gm. skin = 66 mm.’ (approx.).

Volume of subepidermal tissue in 1 gm. skin = 300 mm.’ (approx.).

Volume of transition tissue in 1 gm. skin = 450 mm.’ approx.).

Average volume of epidermal cell = 1:8 x 10°° mm.® (approx.).

Therefore number of epidermal cells in 1 gm. skin = (66/1°8) x 10° (approx.).

Average volume of a subepidermal cell = 3:3 x 10°° mm.® (approx.).

Therefore number of subepidermal cells in 1 gm. skin = (300/3°3) x 10°
(approx.).

Average volume of transition cells = 36 x 10° mm.’ (approx.).

Therefore number of transition cells in 1 gm. skin = (450/36) x 10° (approx.).

Thus total number of cells in 1 gm. skin was of the order of 140 x 10°.

The ratio (number of cells in 1 gm. skin)/(number of cells in 1 gm. flesh) was
25/1 approximately. As this was of the same order as the ratio (respiration of 1 gm.
~skin/hr.)/(respiration of 1 gm. flesh/hr.), the difference in magnitude between skin
respiration and flesh respiration might be accounted for by the greater number of
cells per unit fresh weight in the skin than in the flesh; the average respiration rate
per cell is approximately the same in different parts of the fruit. This is analogous
to the constancy of the respiration rate per cell previously shown in developing Granny
Smith apples (Hackney, 1944).

CORRELATION BETWEEN RESPIRATION RATE OF FLESH TISSUE AND RESPIRATION
RATE OF THE WHOLE APPLE BEFORE CUTTING.

During August, 1944, a series of experiments was carried out in which the
respiration rate of flesh tissue was compared with the respiration rate of the whole
apple prior to cutting. Three samples of 25 apples each were removed from store on
1.viii.44, 14.viii.44 and 28.viii.44 respectively. Using the Pettenkofer technique, the rate
of carbon dioxide output (mg./10 Keg./hr.) was determined for each apple during the
48 hours immediately before cutting. Some of the apples were cut within two days
after removal from store; others had been out of store for as long as 13 weeks before
being cut. The respiration rate of the whole fruit was falling steadily during the ~
post-storage life (cf., Hackney, 1943). The rate of oxygen uptake in distilled water of
a 2 gm. sample of flesh tissue from each apple was then determined. Assuming that
the respiratory quotient was approximately 1 (Hackney, 1944), the values for
oxygen uptake of flesh were converted from mm.’/gm./hr. to their equivalents in
mg. CO./10 Kg./hr., for purposes of comparison with the respiration rates of the uncut
apples. Table 6 shows a typical set of results, given by the first sample of apples. Hach
pair of figures refers to a single apple.

When the data from all three batches were considered, it was found that there
was a strong positive correlation between the respiration rate of the whole fruit and
that of flesh tissue cut from it. The regression coefficient was 0:75 and was highly
significant (P << 0-001). A similar correlation was indicated during 1945, but the
data were not numerous enough to merit statistical analysis. It appears that when
the respiration of the whole apple is high, that of its flesh tissue is high; when the
respiration of the whole apple is low, owing to length of time after removal from
store or to individual variability, the respiration of its flesh tissue is also low.

The respiration rate of skin was not determined in this experiment. However, in
view of the correlation observed in other experiments between the respiration rates of
skin and flesh (see earlier discussion), it is highly probable that the respiration rate
of the skin was correlated with that of the whole fruit.

BY FRANCES M. V. HACKNEY. 34

oo

COMPARISON BETWEEN RESPIRATION RATE OF WHOLE APPLE AS DETERMINED DIRECTLY
BY THE PETTENKOFER METHOD AND AS CALCULATED INDIRECTLY BY THE
WaARBURG MErHop.

In order to compare the respiration rate of the whole apple after cutting with that
of the whole apple prior to cutting, it was necessary to determine the respiration rate
of the cut skin as well as that of the flesh.

TABLE 6.

Comparison between Respiration Rate of Whole Apple and that of
Flesh Tissue.

Respiration of Flesh Tissue

Respiration of Whole Apple ate zilicaCs
Be PAIL (OE (mg./10 Kg./hr.)
(mg./10 Keg./hr.) (Cale. on True Fresh Wt.

at Close of Exp.

270 373
250 307
225 293
217 346
210 266
210 293
207 266
205 293
200 260
195 266
200 277
195 293
195 266
190 213
160 173
140 227
140 253

On 30.x.45 two apples were removed from store and held at 25°C. The concen-
trations of oxygen and carbon dioxide in their internal atmospheres were determined
by the method described by Trout et al. (1942). It was found that the fruits were
probably at the stage where internal oxygen concentration was limiting respiration
rate (cf., Hackney, 1943). The respiratory drifts of the two fruits were observed,
using the Pettenkofer method. All the skin was then removed from the fruits and
weighed; the remainders of the total weights of the fruits were regarded as ‘flesh’.
The respiration rates of samples of skin and flesh in 0:25M calcium chloride were then
determined, using the Warburg method. From the respiration rates of these samples
it was possible to calculate the respiration rate of the whole fruit. The respiratory
quotient of an intact apple (Hackney, 1944) and of cut apple tissue under the conditions
of the experiment were both approximately equal to unity, i.e., the volume of carbon
dioxide given off was approximately equal to the volume of oxygen taken up. Both
fruits behaved similarly. The following data were obtained with one of them:

Respiration rate of flesh = 7:°3 mm.’®/em. fr. wt./hr.

= (7:3/500) meg./gm./hr.
Total weight of flesh = 125 gm.
Therefore total respiration of flesh = 1:83 mg./fruit/hr.
Respiration rate of skin = 135 mm.*/gm. fr. wt./hr.

= (135/500) meg./gm./hr.
Total weight of skin = 4:34 gm.
Therefore total respiration of skin = 1:17 mg./fruit/hr.
Therefore calculated respiration of whole fruit = 3:0 mg./fruit/hr.

= 233 -mg./10 Ke. /hr.

The observed respiration rate of this fruit prior to cutting was 200 mg./10 Kg./hr.
Thus the calculated respiration rate was 33 units higher than the observed rate. As
Mentioned previously, the apples were probably at the stage where internal oxygen
supply was limiting respiration rate. The internal oxygen concentration was governed

344 STUDIES IN THE METABOLISM OF APPLES. VI,

by the resistance of the skin to the passage of oxygen (Hackney, 1943). When the
tissues were cut and exposed to the air, the oxygen supply was suddenly increased to
approximately 21% (the internal oxygen concentrations of the two fruits prior to
cutting were 7:0% and 6:2% respectively). In order to determine whether the difference
between observed respiration rate prior to cutting and calculated respiration rate after
cutting could be attributed to the change in oxygen supply to the tissues, the skin and
flesh were surrounded with gas mixtures containing higher percentages of nitrogen
and lower percentages of oxygen than are normally present in air. Table 7 shows the
calculated respiration rates of the two fruits in these gas mixtures.

TABLE 7.
Comparison of Respiration Rates of Whole Fruits (mg./10 Kg./hr.) obtained by Pettenkofer and
Warburg Methods; Internal Oxygen Concentrations prior to Cutting; Effects on Calculated
Respiration of Lowering Oxygen Supply.

Resp’n. (Warburg).

Fruit No. Int. O;-(%): Resp’n.
(PRettenkofer). In Air. lia ILS, Oe. 1 OVS Oh,
7-0 200 233 223 190
2 6:2 196 221 215 190

It is clear from the above table that, when the cut tissues were surrounded with
a gas mixture containing a percentage of oxygen approximately equal to that present
in the internal atmosphere prior to cutting, the calculated respiration rate of the whole
fruit was very close to the observed value prior to cutting. This indicates (1) absence
of prolonged stimulation of respiration due to cutting, and (2) pronounced effect of
increase in oxygen concentration within certain limits on the respiration rate. The
second point is in accordance with the results obtained in previous years when whole
apples were held in-an atmosphere of pure oxygen (Hackney, 1944).

When the skin and flesh were returned to air after being surrounded with the
6% oxygen mixture the calculated respiration rate of the whole fruit returned to the
air level. Substitution of 100% oxygen for air did not result in any further increase
in respiration rate.

The change brought about in respiration by decreasing the oxygen supply from
11% to 6% was greater than that brought about by decreasing it from 21% to 11%.
In the two fruits used, the effects of changes in oxygen concentration were apparently
greater at lower oxygen concentrations than at higher oxygen concentrations. Another
point of interest is that these effects were much more pronounced on skin tissue than
on flesh tissue. This may have been due to the respiration rate per unit weight being
so much higher in the skin than in the flesh.

-The experiment described above indicated the possibility of using the cut tissue
technique as a means of investigating the respiration of apple tissue under conditions
where oxygen supply is not limited by the presence of the skin.

SUMMARY.

The technique for the determination of the respiration rate of sliced apple tissue
has been described and discussed in detail.

Respiration rates were determined for tissues from different regions of the Granny
Smith apple. There was no significant difference in respiration rate between flesh
tissue slices from various parts of the apple, but the respiration rate of the skin per
unit weight was many times higher than that of the flesh. This was shown to have
been probably due to the greater number of cells per unit weight in the skin than in
the flesh.

There was a strong positive correlation between the respiration rate of the skin
and that of the flesh from the same apple.

There was a strong positive correlation between the respiration rate of the whole
apple and that of flesh tissue cut from it.

BY FRANCES M. V. HACKNEY. 345

Values for the total respiration rates of whole apples, calculated from the respiration
rates of skin and flesh tissue, were greater than the values obtained directly with the
uncut apples. This increase in total respiration rate after cutting was shown to have
been probably due to the increasing of the oxygen supply to the tissues consequent
upon cutting.

ACKNOWLEDGEMENTS.

The writer wishes to express her thanks to Professor E. Ashby and to Acting
Professor J. McLuckie, Department of Botany, University of Sydney, for their helpful
advice and for the use of laboratories and facilities; to Professor J. S. Turner,
Department of Botany, University cf Melbourne, and to Dr. R. N. Robertson, Department
of Botany, University of Sydney, for their guidance in the development of the technique
and in the planning of the investigations; to Dr. J. R. Vickery, Dr. S. A. Trout, and
other members of the staff of the Division of Food Preservation, Council for Scientific
and Industrial Research, Homebush, and to Mr. S. M. Sykes, Department of Agriculture,
New South Wales, for their helpful advice and criticism.

REFERENCES.

BENNET-CLAREK, T. A., and BExon, D., 1943.—Water Relations of Plant Cells. ili. The
Respiration of Plasmolysed Tissues. New Phytol., 42 (2): 65-92.

BOSWELL, J. G., and WHITING, G. C., 1938.—A Study of the Polyphenol Oxidase System in
Potato Tubers. Ann. Bot., N.S., 2: 847-863.

Drxon, M., 1943.—Manometric Methods. Cambridge University Press.

HAcKkNeEy, F. M. V., 1943.—Studies in the Metabolism of Apples. li. The Respiratory
Metabolism of Granny Smith Apples of Commercial Maturity after Various Periods of
Cool Storage. Proc. LINN. Soc. N.S.W., 68: 33-47.

——— —, 1944.—Id. iv. Further Studies in the Metabolism of Granny Smith Apples, with
Special Reference to the Importance of Oxygen. Ibid., 69: 91-107. ;

ROBERTSON, R. N., 1941.—Studies in the Metabolism of. Plant Cells. 16 Accumulation of
Chlorides by Plant Cells and its Relation to Respiration. Aust. J. Hup. Biol. Med. Sci.,
19: 265-278.

TETLEY, U., 1930.—A Study of the Anatomic Development of the Apple and Some Observations
on the ‘Pectic Constituents’ of the Cell-Walls. J. Pom. Hort. Sci., 8 (2): 1538-172.

————,, 1931.—The Morphology and Cytology of the Apple Fruit. Ibid., 9 (4): 278-297.

Trout, S. A., et al., 1942.—Studies in the Metabolism of Apples. i. Preliminary Investigations
on Internal Gas Composition and its Relation to Changes in Stored Granny Smith Apples.
Aust. J. Eup. Biol. Med. Sci., 20: 219-231.

TURNER, J. S., 1938a.—The Respiratory Behaviour of Carrot Tissue. i. Materials and Methods.
New Phytol., 37: 232-253.

————, 1938b.—Id. il. The Effect of Sodium Monoiodoacetate on the Respiration. and
Fermentation. Ibid., 37: 289-311.

———, 1940.—Id. iii. The Drift of Respiration and Fermentation in Tissue Slices, with
Notes on the Respiratory Quotient. Aust. J. Exp. Biol. Med. Sci., 18: 273-298.

PLATE VIII.

1945.

N.S.W.,

LINN. Soc.

Proc.

Marsupials.

The Hair Tracts in

Ix.

PLATE

N.S.W., 1945.

Linn. Soc.

Proc.

uminous Plants.

Fixation in Leg

itrogen

T,
4

Proc. Linn. Soc. N.S.W., 1945. PLAT Xx.

Diptera of the Territory of New Guinea.

ee y
a
= ¢ t
we
=
ee
a
; = 2
~ ; ; \
«
- = 7
g
‘
4
r Sele
‘ wire - 2 a
re

Proc. Linn. Soc. N.S.W., 1945. PLATE XI.

Reptiles in the Macleay Museum.

—

f

——

Proc. Linn. Soc. N.S.W., 1945. PLATE XII.

Wings of Tripteroides spp.

—\

>

Proc. Linn. Soc. N.S.W., 1945. PVADT i xcnue

tia

PO ence’

Wings of Tripteroides spp.

Z
‘
{ae

af

ti

“i

THE

PROCEEDINGS

OF THE

PIPSEAN SOCIETY

tela

NEw SouTH WAL ES———_—-— |}

FOR THE YEAR

1945.

eR RSE SA TEST

Parts i-ii (Pages i-wxviii; 1-52).

i

CONTAINING THE PROCEEDINGS OF THE ANNUAL MEETING
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$ Wice-Presidents:
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PROCEEDINGS, LXX, PARTS 1-2, 1945.

- CONTENTS.

Pages

Presidential Address, delivered at the Seventieth Annual General Meeting, 28th ; ‘a

March, 1945, by W. R. Browne, D.Sc. EO Nice Rae al a tn eA ia ele Ca
Elections ..

Balance Sheets for the Year ending 28th February, 1945 .. .. Re eee.) KXVIEXXVili

’ Some Points in the External Morphology of the Pouch Young of the Marsupial,
Thylacinus cynocephalus Harris. By W. Boardman. (Plate i and three
AMP ext APUG. Jiu.) ervey aa fy Go ba eee Ua ea Sales a ac matey eo aoe no 1-8

Stenoporids from the Permian of New South Wales and Tasmania. By Joan
Crockford, M.Sc., Linnean Macleay Fellow of the Society in Palaeontology.
(Plates li-iii and twenty-five Text-figures.) .. 0 .. 0 .: 0 2.004. ee 9-24

The Cranial Nerves of Neoceratodus. By: ob. Leighton Kesteven, D.Se., M.D.
(Hour PERtHe ures ei Par Mae ha alien ence aeh a MNS Unatedes Airc CON ARIere a RM ga et 25-33

Correlation of Some Carboniferous Sections in New South Wales, with Special Bs
Reference to Changes in Facies. By Alan H. Voisey, M.Se. (Plates iv—v.) 34-40

Evidence of an Eustatic Strand-line Movement of 100 to 150 feet on the Coast of |
New South Wales. By Wilson H. Maze, M.Sc. (Four Text-figures and one -
Map.) Sat mins neat Sita Meant Hosein SEU ag SRE aR OI a Al Ne ep RO 41-46 ©

On Australian Dermestidae. Part iv. Notes and the Description of a New
-.Genus and Four New Species. By J. W. T. Armstrong. (Three Text- Een
FIGULES..) cg A ii coal aair sient orn Unicoi Uh ange aa let ars facies ath ne otstar eyo aed De aiaee nate 47-52

(Issued 15th February, 1946.*)

oc ; co

J Vol. LXX. Nos. 319.320, |

Parts 3-4.
THE
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FOR THE YEAR DEC 2-4 F
1945. : WOODS HOLE, |

Parts IiI-IV (Pages 53-178).
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* This issue has been delayed by circumstances over which the Society had no control.

The Linnean Society of New South Wales
LIST OF OFFICERS AND COUNCIL, 1945-46.

President:
Ida A. Brown, D.Sc.

‘Vice-Presidents:
R. H. Anderson, B.Sc.Agr. E. Le G. Troughton, C.M.Z.S., F.R.Z.S.
F. H. Taylor, F.R.E.S., F.Z.S. ; W. R. Browne, D.Sc. ;

Hon. Treasurer: A. B. Walkom, D.Sc.
Secretary: N. S. Noble, D.Sc.Agr., M.Sc., D.C.

Council:

R. H. Anderson, B.Sc.Aegr. G. D. Osborne, D.Sc., Ph.D.
E. C. Andrews, B.A., F.R.S.N.Z. R. N. Robertson, B.Se., Ph.D.
Professor E. Ashby, D.Sce., A.R.C.S., D.4.C., T. C. Roughley, B.Sc., F.R.Z.S.

¥F.L.S. C. A. Sussmileh, F.G.S.
Ida A. Brown, D.Sc. F. H. Taylor, F.R.E.S., F.Z.S.
W. R. Browne, D.Sc. E. Le G. Troughton, C.M.Z.S., F.R.Z.S.
A. N. Colefax, B.Sc. A. B. Walkom, D.Sc.
Lilian Fraser, D.Sc. H. S§. H. Wardlaw, D.Sc., F.A.C.1.

Professor J. Macdonald Holmes, B.Sc. ;
Ph.D., FR.GS., F.RSGS. " _W. L.. Waterhouse, M.C., D.Sc.Agr., D.LG

N. S: Noble, D.Sc.Agr., M.Sc., D.I.C. *A, R. Woodhill, B.Sc.Agr.
Auditor: S. J. Rayment, A.C.A. (Aust.).
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Ne ea hee

DESCRIPTIVE CATALOGUE OF AUSTRALIAN FISHES. By William Macleay, F.L.S. [1881].
A few copies only. Price £1 net.

The TRANSACTIONS OF THE ENTOMOLOGICAL SocIRTY OF Now SouTH WALBS, 2 vols., 8vo.
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(complete in five parts, 1869-73), price 30s. net, or single Parts 7s. 6d. each.j

Girton Ss Ae Te eS

PROCEEDINGS, LXX, PARTS 3-4, 1945.

CONTENTS.

Relation of the Orchid Flora of Australia to that of New Zealand, with the
Description of a New Monotypic Genus for New Zealand. By H. M. Ry
Rupp, B.A., and E. D. Hatch

Geographic Variation in the Lizard Hemiergis decresiensis (Fitzinger). By
Stephen J. Copland, B.Sc. (Plate vi and five Text-figures.) ..

Revision of Australian Lepidoptera. Oecophoridae. xiii. By A. Jefferis
Turner, M.D., F.R.E.S. ..

Studies on Australian Marine Algae. ii. Notes extending the Known ~

Geographical Range of Certain Species. By Valerie May, M.Sc.

A Bryozoan Fauna from the Lake’s Creek Quarry, Rockhampton, Queensland.
By Joan Crockford, M.Sec., Linnean Macleay Fellow of the Society in
Palaeontology. (Twelve Text-figures.) ..

Miscellaneous Notes on Australian Diptera. xi. Evolution of Characters in
the Order: Venation of the Nemestrinidae. By G. H. Hardy. (Two Text-
figures. )

Principal Microspore-types in the Permian Coals of New South Wales. By
J. A. Dulhunty, B.Sc. (Plate vii and three Text-figures.)

Petrological Studies in the Ordovician of New South Wales. ili. The
Composition and Origin of the Upper Ordovician Graptolite-bearing Slates.
By Germaine A. Joplin, B.Sc., Ph.D., Linnean Macleay Fellow of the
Society in Geology. (Four Text-figures.)

On the Family Smarididae (Acarina). By R. V. Southcott, M.B., BS. (Four
Text-figures.) Re SUD ta iu \eend Shem rar a

ot

Pages.

53-61

62-92

93-120

121-124

4

125-134

135-146

147-157

158-172

173-178

\

THE

PROCEEDINGS

OF THE

@ _INNEAN SOCIETY

OF

ee re ee

wating Biologi

| : |
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D
FOR THE YEAR

1945. | WOODS HOLE,

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Parts V-VI (Pages 179-345; xaxix—-axli).

CONTAINING PAPERS READ IN SEPTEMBER-NOVEMBER,
ABSTRACT OF PROCEEDINGS, LIST OF MEMBERS,
AND GENERAL INDEX.

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STEER TEESE PR Le TE OM ETE AE SE OE a A a eS EE OE

* This issue has been delayed by circumstances over which the Society had no control.

The Linnean Society of New South Wales.
LIST OF OFFICERS AND COUNCIL, 1945-46.

President:
Ida A. Brown, D.Sc.

WVice-Presidents:
R. H. Anderson, B.Sc.Agr. . H. Le G. Troughton, C.M.Z. s., F.R.Z.S.
*F. H. Taylor, F.R.E.S., F.Z.S. W. R. Browne, D.Sc.

Hon. Treasurer: A. B. Walkom, D.Sc.

Secretary: N. S. Noble, D.Sc.Aer., M.Sc., D.I.C.

Council:
R. H. Anderson, B.Sc.Agr. G. D. Osborne, D.Se., Ph.D.
E. C. Andrews, B.A., F.R.S.N.Z. R. N. Robertson, B.Sc., Ph.D.
Professor E. Ashby, D.Sc., A.R.C.S., B.2.C., T. C. Roughley, B.Sc., F.R.Z.S.
F.L.S. C. A. Sussmilch, F.G.S.
Ida A. Brown, D.Sc. *F, H. Taylor, F.R.E.S., F.Z.S.

W. R. Browne, D.Sc.

A. N. Colefax, BSc. EH. Le G. Troughton, C.M.Z.S., F. R.Z. =

Lilian Fraser, D.Se. ee

Professor J. Macdonald Holmes, B.Sc., H. S. H. Wardlaw, D.Se., F.A.C.1.
Ph.D., F.R.G.S., E.R.S.G.S. W. Li. Waterhouse, M.C., D.Sc.Ager., PD aC-

N. S. Noble, D.Se.Agr., M.Sc., D.I.C. A. R. Woodhill, B.Se.Agr. zt
Auditor: S. J. Rayment, A.C.A. (Aust.). é
* Died 20th December, 1945.

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David Nutt, 212 Shaftesbury Avenue, London, W.C.2. The stock of copies of First
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SIS cates are 5 0 5 0 6 0 7 6 LOO ee dice OO 9 0 5 0 17 6
1879 .. .- 6 0 6 0 8 0 6 6 DOOD aeons 7 0 7 6 426%) 15270,
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- 1Supplement is. 6d. additional. «Supplement 1s. additional.
2 Supplement 2s. 6d. additional. 5 Supplement 2s. 6d. additional.

3 Supplement 3s. additional. a

Year.| Part 1. | Part 2. | Part 3. | Part 4. | Part 5.

2 loa
1922 .

s. d s. er acraay s. d s. d. 8. d.
2 6 13 6 11 0 13 0 20
1923 PAN} 12 0 16 0 18 6 250
1924 2 0 13 6 12 6 10 0 TeA0
1925 2-0 12 0 8 9 14 6 20
» 1926 20 13 6 9 6 15 0 2 0
1927 2 6 10 6 14 0 12 3 2°-0
art | Part | Part | Part Part | Part

Year : 2 3. 4,
3s. d s. d.| s. d s. d. s. d.| s. d.
1928 |2 0 8 6|8 9/10 0 9 0/32 0
1929 | 2 0 6 3/8 6) 11 0/12 6|;2 0
1930 |} 2 0/]10 0:8 9 8526.2} 163) 3, 2520
1931 |2 0 8 0|7 0 Ca-3 §2-9:)|5220

Year. Parts 1-2. |} Parts 3-4. Parts 5-6.

s. d. 8. d. s. d.
1932 6 6 9 6 8 6
1933 ate 8 0 11 9 8 6
1934 7 6 6 8 6 10 9
1935 ne 42239 105-3 11 3
1936 oe 6 6 eon! 9 9
1937 oe 6 0 93 8 9 =
1938 6 3 11 6 9 9
1939 12°3 12 3 8 9
1940 11 9 10 6 8 0
1941 6 6 9 3 9 6
1942 A 9 0 12 8 6 9
1943 5 6 6 9 8 3
1944 6 6 9 0 8 0
1945 6 0 8 9 | 18 6

INDEX To VoLuMBs I-L oF THE PROCEEDINGS [Issued 15th February, 1929]. Pages 108.
Price 5s.
The MAcCLEAY MEMORIAL VOLUME [Issued 13th October, 1893]. Royal ato, li and 308
pages, with portrait, and forty-two piates. Price £2 2s.

DESCRIPTIVE CATALOGUE OF AUSTRALIAN Fisizs. By William Macleay, F.L.S. [1881].
A few copies only. Price £1 net. ;

The TRANSACTIONS OF THE HNTOMOLOGICAL Society or NEw SouTH WALES, 2 volis., Sve.
[Vol. I (complete in five parts, 1863-66), price 70s. net. Parts 2-5 10s. each; Vel. ii
(complete in five parts, 1869-73), price 30s. net, or single Parts 7s. 6d. each.]

(

PROCEEDINGS, LXX, PARTS 5-6, 1945.

CONTENTS.
: Pages.
The Hair Tracts in Marsupials. Part ii. Description of Species, continued.
By W. Boardman. (Plate viii and forty-three Text-figures.) 179-202
Nitrogen Fixation in Leguminous Plants. vi. Further Observations on the
Effect of Molybdenum on Symbiotic Nitrogen Fixation. By H. lL. Jensen,
Macleay Bacteriologist to the Society. (Plate ix and one Text-figure.) 203-210 -
Contributions to a Knowledge of Australian Culicidae. No. viii. By Frank no
Taylor, F.R.E.S., F.Z.S. (One Text-figure.) 211-212
Notes on Some Fijian Mosses. By William Greenwood .. 213-218
Notes on Australian Mosquitoes (Diptera, Culicidae). Part vi. The Genus
Tripteroides in the Australasian Region. By David J. Lee, B.Sc. (Plates
xii-xiii and sixty-nine Text-figures.) 219-275
Observations on the Morphology and Biology of the Subspecies of Anopheles
punctulatus Dénitz. By A. R. Woodhill. (Three Text-figures and one
Map.) Wheat wale nea aon Fe ae eS Lie ott aioe . 276-287
Notes on New South Wales Orchids: A New Species and Some New Records.
By the Rev. H. M. R. Rupp, B.A. (One Text-figure.) .. 288-290
Catalogue of Reptiles in the Macleay Museum. Parti. Sphenomorphus pardalis
pardalis (Macleay) and Sphenomorphus nigricaudis nigricaudis (Macleay).
By Stephen J. Copland, B.Sc. (Plate xi and seven Text-figures.) .. 291-311
Studies on Trombidiidae (Acarina). Some Observations on the Biology of the
Microtrombidiinae. By R. V. Southcott, M.B., B.S. (Three Text-figures.) 312-316
A Contribution to a Study of the Physiology of Decay in Apples. By Mary
Cash, M.Se. (Ten Text-figures.) 317-327
The Diptera of the Territory of New Guinea. xiii. Family Tabanidae. Part i.
The Genus Chrysops. By Frank H. Taylor, F.R.E.S., F.Z.S. (Plate x and
two TOKCHEUTER)) oe ee ae SN I let uad a scent sty anes : 328-332
f
Studies in the Metabolism of Apples. vi. Preliminary Investigations on the
Respiration of Sliced Apple Tissue. By Frances M. V. Hackney, M.Sc.,
Linnean Macleay Fellow of the Society in Plant Physiology. (Three Text-
figures.) Mite aer emma ene Arann cra er 333-345
Abstract of Proceedings .. .. Fea Ter ra A, Se pean Rea ONION OBR ARS LTE Ane XxXixX—XXxii
} os ¥
: \ )
List of Members ©... 0... Be ee i ies 2 EXT ORRRVEL
Lista Plates 2 Os XXXVii.
List of New Genera and Species

XXXVI

Genoral THGEK he iN ea ea RS aE Aaa Sees ees cet ier aie Saas Xxxix—xli

ry

BL/WHOIL LIBRARY

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