THE
PROCEEDINGS
OF THE
LiriMEnn Society
OF
New South Wales
FOR THE YRAR
1918
Vol. XLIII.
WITH NINETY-THREE PLATES,
and 112 Text-figures.
SYDNEY :
PRINTED AND PUBLISHED FOR THE SOCIETY
BY
W. A. PEPPERDAV ik CO., 119a PITT STREET
AND
SOLD BY THE SOCIETY
1918-19.
\V. A. PEPPKkDAY AND CO.,
GRNERAI. PRINTKRS,
119a PITT STREET, SYDNEY.
CONTENTS OF PROCEEDINGS, 1918.
PART I. (No. 1G9).
(Issued Srd July, 1918).
pa(;ks
Presidential Address delivered at the Forty-third Annual Meet-
ing, March 27th, 1918, Vjy Professor H. G. Chai'MAX, M.D.,
B.S., President 1-30
The Geology of the Lower Mesozoic Rocks of Queensland, with
special Reference to their Distribution and Fossil Flora, and
their Correlation with the Lower Mesozoic Rocks of other
parts of Australia. By A. B. Walkom, D.Sc, Assistant
Lecturer in Geology, University of Queensland. (Plates
i.-ii., and six Text-figures) ... ... ... ... ... 37-11.5
Studies in Australian Neuroptera. No. 5. The Structure of the
Cubitus in the Wings of the MyrmeJtont\d<^. By R. J.
TiLLYAKD, ]SLA., D.Sc, F.L.S., F.E.S., Linnean Macleay
Fellow of the Society in Zoology. (With three Text-figures) 116-122
On the Affinities of two interesting Fossil Insects from the Upper
Carboniferous of Commentry, France. By R. J. Tilia'ak]>,
M.A., D.Sc, F.L.S., F.E.S., Linnean Macleay Fellow of the
Society in Zoology. (With three Text-figures) 123-134:
Tha Origin of Yolk in the Ova of an Fiudoparasitic Copepod.
By H. Leighton Kestevex, D.Sc, M.B., Ch.M. (Plate iii.) 13«;-Ul
Contributions to our Knowledge of Soil-Fertility. No. xvi. The
Search for Toxin-Pnjducers. By R. Greki-Smitii, D.Sc,
Macleay Bacteriologist to the Society ... ... ... ... 142-190
On certain Shoot-bearing Tumours of Eucalypts and Angophoras;
and their modifying influence on the natural Growth-habit of
the Plants. By J. J. Fletcher and C. T. Musson. (Plates
iv.-xxvi.) 191-233
Hon. Treasurer's Financial Statement and Balance Sheet ... 30-34
Elections and Announcements ... ... ... ... ... 135,234
Notes and Exhibits 235-236
27801
IV. CONTENTS.
PART IL (No. 170).
(Issued loth Septemhtr, lOtS). pages
Oil the Seasonal Distribution of some Queensland Species of
Arrella Ehrenbeij,' [Rhizopoda]. By C. D. Gilijes, M.8e.,
Biolof,'y Department, University of Queensland. (Four Text-
fi^aires) 287-246
On the Leaf-Anatomy of Sovrola crassi/ofia [Goodeniace.*;], with
special reference to the Epidermal Secretion. By Marjorie
Isabel Collins, B.Sc, Demonstrator in Botany, Universit}^
of Adelaide. (Plates xxvii.-xxviii.; and six Text-figures) ... 247-259
A Fossil Insect-wing from the roof of the Coal-seam in the Syd-
ney Harbour Colliery. By R. J. Till yard, M.A., D.Sc,
F.L.S., F.E.S., Linnean Macleay Fellow of the Society in
Zoology. (One Text-figure) 260-264
The Panorpoid Complex. A Study of the Phylogeny of the
Holometabolous Insects, with special Reference to the Sub-
classes Panorpoidea and Neiirapteroidea. [Introduction].
By R. J. Tillyakd, M.A., D.Sc, F.L.S., F.E.S., Linnean
Macleay Fellow of the Society in Zoology 26.)-284
The Panorpoid Complex. Part i. The Wing-Coupling Apparatus,
with special Reference to the Lepldopttm. By R. J. Till-
YARD, M.A., D.Sc, F.L.S.. F.E.S., Linnean Macleay Fellow
of the Society in Zoology. (Plates xxix.-xxx.; and .sixteen
Text-figures) 2S6-819
The Geology and Petrology of the Great Serpentine Belt of New
South Wales. Part vii. The Geology of the Loom V)e rah Dis-
trict and a portion of the Goonoo Goonoo Estate. B3' W.
N. Benson, B.A., D.Sc, F.G.S,, Profes.sor of Geology and
Mineralogy in the University of Otago, N.Z., late Linnean
Macleay Fellow of the Society in (Geology. (Plates xxxi.-
xxxviii. ; and four Text-figures) 320-360. 363-384
With two Pahwntological Appendices by F. Chapman, F.R.M.S.,
A.L.S., Paheontologist to the National Museum, Melbourne.
(Plates xxxix.-xlii.) 3Sr)-3y4
Studies in Australian Mecoptera. No. ii. The Wing-Venation of
Chorista amtralis Klug. By R. J. Tillyakd, M.A., l^.Sc,
F,L.S., F.E.S., Linnean Macleay Fellow of the Society in
Zoology. (Plate xliii,, and four Text-figures) 395-408
The "Springing" of Tins of Preserved Fruit. By W, W,
LEsTRANGE and R. Greio-Smith 409-414
Elections and Announcements 285,362,416
Notes and Exhibits 361,415,416
CONTENTS. V.
PART III. (No. 171).
(IsmuhI ISth Decemher, IfilSJ. pa(;es
Mesozoic Insects of Queeiislaiul. No. 8. Odonata and Protodonntn.
^^y R. J. TiLLYAKD, M.A , D,,Sc., F.L.8., F.E.8., Linnean
Macleay Fellow of the Society in Zoology. (Plates xliv.-xlv.,
and Text-tiguies 11-16) 417-436
The Carboniferous Trilobites of Australia. By JoHX MiT( hell,
late Principal, Teclinical College at Newcastle, N.S.W,
(Plates xlvi,-liii.) 437-494
New and rare Fieshwater Alga^ By G. I. Playfaik, late Science
Research Scholar of the University of Sydne\'. (Plates liv.-
Iviii., and eleven Text-tigures) ... ... ... ... ... 497-'")43
The Mosses of North Queensland. By Dr. V. F. Bkotiiekus and
the Rev. W. Walter Watts o44-o67
Mesozoic In.sects of Queensland. No. 4. Htmiptera HeJeroptera :
the Faniilj' Dun.sta7iiid<e. With a Note on the Origin of the
Heteroptera. B3' R. J. Tij.lyakd, M.A. , D.Sc, F. L.S. ,
F. E.S., Linnean Macleay Fellow of the Society in Zoolog}'.
(Plate lix., and Text-tigures 17-22) ... ... ... ... 368-592
The (Teolog3' and Petrology of the Great Serpentine Belt of New
South Wales. Part viii. Tlie Extension of the (rreat Serpen-
tine Belt from the Nundle District to the Coast. By W, N.
Benson, B.A., D.Sc, F.G.S., Professor of Geology and Min-
eralogy in the Univei'sity of Otago, N.Z., late Linnean Mac-
leay Fellow of the Societj' in Geology. (One Text-figure) ... o93-599
The Steins of Climbing Plants. By John Shirley, D.Sc, and
C. A. Lamrert. 'Plates Ix.-lxvi.) 6(KJ-609
The Relation between the Fat-Content and the Electrical Con-
ductivity of Milk. By H. S. Halcro Wardlaw, D.Sc,
Linnean Maclea3' Fellow of the Society in Physiology ... 613-62o
The Panorpoid Complex. Part 2. The Wing-Tricliiatiijn and its
Relationship to the General Scheme of Venation. By R. J.
Ti].LYARD, M.A., D.Sc, F.L.S., F.E.S., Linnean Macleay
Fellow of the Society in Zoology. (Plates Ixvii.-lxix., and
Text-figures 17-34) 626-657
A Study c)f the External Breathing-Apparatus of the Larvie of
.some Muscoid Flies. By John L. Froguatt, B.Sc (Plate
Ixx.) 658-667
VI. CONTENTS.
PART III. (Continmd). pages
Notes on Australian Sawflies (Tenthredimdre). By Walter W.
Fro(u;att, F;L.S., (Government Entomologist 668-672
Elections and Announcements ... ... ... ... ...495,610
Notes and Exhibits 495,610
PART IV. (No. 172).
(Ismed 26th March, 1910). pages
Notes on the Native Flora of New South Wales. Part x. The
Federal Capital Territory. By R. H. Cambage, F.L.S.
(Plates Ixxi.-lxxiv.) 673-711
Descriptions of new Species of Australian Cohoplera. Part xiv.
By Arthur M. Lea, F.E.S 715-746
On a new Species or Form of Eucalyptus. By Cuthbert Hall,
M.D., Ch.M. (Plate Ixxv.) 747-749
Studies in Australian Nenroptera. No. 6. The Family Pst/chop-
sidfJd, with Descriptions of new Genera and Species. By R.
J, TiLLYARD, M.A., D.Sc, F.L.S., F.E.S., Linnean Macleay
Fellow of the Society in Zoology. (Plates Ixxvi.-lxxviii. ;
and four Text-figures) 750-786
Studies in Australian Xeuroptera. No. 7. The Life-history of
Psychopsis e/ei/ans (Guerin). By R. J. Till yard, M.A.,
D.Sc, F.L.S., F.E.S., Linnean Macleay Fellow of the Society
in Zoology. (Plate Ixxix., and twelve Text-figures) 787-818
Australian Mef/afoptera or Alder-Flies, with Descriptions of new
Genera and Species, By R. J. Tillyard, M.A. , D.Sc,
F.L.S., F.E.S., Linnean Macleay Fellow of the Soeiety in
Zoology. (Three Text-figures) 819-825
Contributions to a knowledge of Australian (Ju/icid'V. No. iv.
By Frank H. Taylor, F.E.S. (Plates Ixxx.-lxxxiii.) ... 826-843
Note on the Temperature of Echidna acnleata. By H. S. Halcro
Wardlaw, D.Sc, Linnean Macleay Fellow of the Society in
Physiology. (Two Text-figures) 844-849
The Occurrence of Methyl Ltevo-Inositol in an Australian Poison-
ous Plant. By James M. Petrie, D.Sc, F.I.C, Linnean
Macleay Fellow of the Society in Biochemistry. (Two Text-
figures) ... ... ... ... ••• ... ••• ••• 850-867
CONTENTS. Vll.
PART IV. (Continued). pages
An Account of some Observations upon the Life-histoiy of Plwma
citricarpa Mq A\\i. : tlie Cause of the "Black Spot" Disease
in Citrus Fruit in New South Wales. Bj' G. P. Daknei.l-
Smith, D.Sc, F.I.C. (Plates Ixxxiv.-xc.) 868-882
The Occurrence of an inverted Hynieniuni in A<jaricui< canijieMtris,
ByG. P. Darnell-Smith, D.Sc, F.I.C. (Plates xci.-xciii.) 883-887
Notes and Exhibits 713-714
Donations and Exchanges 888-902
Title-page i.
Contents ... ... ... ... ... ... ... ... iij.
Corrigenda ... ... ... ... ... ... ... ... vii.
List of New Generic and other Names ... ... ... ... vii.
List of Plate?... ... ... ... ... ... ... ... viii.
Index i.-xxix.
CORRIGENDA.
Page 11)1, line 31— ;/b/- length, read lengthen.
Page 231, line 2.3— ^/br Museum, read Herbarium.
Page 235, line 19 — for ('ornicidatu-'^, read corniculata.
Page 315, line 17 — for forewing, read frenulum.
Page 377, line 36— for <S'. partita, read S. partitnm.
Page 415, line 32 — for /anrina, read laurifolia.
Page 669, line 7 — for nor(e-any/iie, read nova-anglica.
Page 702, line 21 — for A. riparins, read .S'. riparins.
Page 827, line 4 — for Anophile-^, read Anopheles.
Page 829, line 23 — after length, add 3 mm.
Page 829, line 33— /'or 1914 and (1915) read 1913 and (1914).
Page 832, line 33— after Skuse, add Taylor.
Page 843, last line— ;/a/- sp.n., read Taylor.
LIST OF NEW SUBORDINAL, TRIBAL, FAMILY, AND GENERIC
NAMES PROPOSED IN THIS VOLUME (1918).
PAOE
Aeroplana (Protodonata) ... 426
Aeroplanid.*: (Protodonata)... 425
Aekoplaxoptera (Pro-
todonata) ... ... ... 425
Arisocepha/ns (Coleoptera) ... 732
Austrosia/is (Megaloptera) ... 821
Creoleon (Neuropterai ... ... 436
C R E o L E o N T I X I ( Neu roptera ) 436
Dicti/eiwnemii^ (Coleoptera) ... 728
Dunstaniop8i>< (Hemiptera) ... 584
El caiiops i s \OvthoptGva,) ... 262
Megapsychops (Neuroptera) ... 771
page
Mesopano7'podes (Mecoptera) ... 435
Mesopanokpodid.^ (Mecopter
Xematodinu.s (Coleoptera)
Notopsychops (Neuroptera)
Paradunstania (Hemiptera)
Ferissoph I eh ia ( Odonat a )
Pterobryidium (Muaci)...
Stenosialis (Megaloptera)
Tasmomlpingus (Coleoptera)
Tessellaria (Algie)
TriassoleMes { Odonata ) . . .
Triassolestin .q^. (Odonata)
) 436
. 730
. 759
. 585
. 422
. 558
. 823
743
. 508
. 418
. 418
Vlll. CONTENTS
LIST OF PLATES
Plate i. — Map of Queensland, showing the distribution of Lower Mesozoie
Rocks.
Plate ii. — (xeological Sketch-Map of S.E. Queensland, showing the areas
occupied by the Lower Mesozoie Rocks.
Plate iii. — Origin of yolk in the ova of an endoparasitic Copepod.
Plates iv.-xxvi. — Shootbearing Tumours of Eucalypts and Angophoras,
Plates xxvii.-xxviii. — Leaf-anatomy of Safrolacrasi^ifolia.
Plate xxix. — Jugo-frenate Wing-coupling Apparatus {Microptery(ji(lif).
Plate XXX. — Jugate {Hepialklre) and Frenate {Sphhigidie) Wing-coupling
Apparatus.
Plate xxxi. — Topographical Map of the Loomberah District.
Plate xxxii. — Geological Map of the Loomberah District,
Plate xxxiii. — Pillow-lavas exposed in the cutting b^^ the Loomberah
Bridge.
Plates xxxiv.-xxxviii. — Fossils from the Loomberah Limestone.
Plates xxxix.-xli. — Devonian Foraminifera from the Tamworth District,
N.S.W.
Plate xlii. — CJuetetes steUiformis, sp.nov.
Plate xliii.— Pupal Wing-tracheation of Chorista auf^tralis: Klug.
Plate xliv. — Aeroplana mirahi/i.^, n.g. et sp. Principal preserved por-
tions of the wings; { x 7).
Plate xlv, — Aiiroplana mirahUiM, n.g. et sp. Restoration of fore- and
hindwings; ( x 4).
Plates xlvi.-liii. ^Carboniferous Trilobites of Australia.
Plates liv.-lviii. — New and rare Freshwater Algte.
Plate lix. — Restoration of right hemelytron (forewing) of Dnnstania
pv/ehra Till.
Plates Ix.-lxvi. — Sections of Stems of Climbing Plants.
Plate Ixvii. — Archiptanorpa ma(jmfica Till.
Plate Ixviii. — Prototheora petrosema Meyr. ; Rhyphu'^ hreris Walk.
Plate Ixix. — Chorista austral is Klug; GaUiphoru rillosa Desv.
Plate Ixx. — Spiracles of larvie of Muscoid Flies.
Plate Ixxi. — Casuarina Luehmanni R. T. Baker.
Plate Ixxii. — Grevilha juniperina R.Br.
Plate Ixxiii. — Banksia marginata Cav.
Plate Ixxi v. — Swampy plain between Gudgen by and Cotter River, at about
4,500 feet level.
Plate Ixxv. — Eucalyptus Mai'sdeni, f. vel sp.n.
Plate Ixxvi. — Meyapsychops illidgeJ (Froggatt).
Plate Ixxvii. — Meyapsychojjs ilJklytl (Froggatt), Psychopsis gracilis, n.sp.,
Psychopsel/a gallardi, n.g. et sp.
Plate Ixxviii. — Psychopsis gracilis, n.sp., Psychopsel/a gallardi, n.g. et sp.
Plate Ixxix. — Psychopsis elegans ((4uer. ).
Plates Ixxx.-lxxxiii. — Australian Cuiicida^.
Plates Ixxxiv.-xc. — Phoma citricarpa McAlp.
Plates xci.-xciii. — Agaricus campestria.
PROCEEDINGS
OF THE
LINNEAN SOCIETY
OF
NIi:^\^ SOUTH W^ALES.
WEDNESDAY, MARCH 27th, 1918.
The Forty-third Annual General Meeting, and the Ordinary
Monthl}^ Meeting, were held in the Linnean Hall, Tthaca Road,
Elizabeth Bay, on Wednesday evening, March 27th, 1918.
ANNUAL GENERAL MEETING.
Pi-ofessor H. G. Chapman, M.D., R.8., Pi-esident, in the Chair.
The Minutes of the preceding Animal General Meeting (March
28th, 1917) were read and confirmed.
The President delivered the Arnuial Address.
PRESIDENTL\L ADDRESS.
Ladies and Gentlemen,
This is the forty-third occasion on which the President lias
delivered an annual address to the Members of the Society. The
progress of this Linnean Society, aiming at advancing the study
of Natural Science in this newly-popidated continent, is also
shown by your election of a graduate of an Australian University
to the presidential chair. Such a selection bears witness to the
increasing influence and preponderating numbers of the scientists
trained in Australia in the personnel of the membership. This
is the first occasion on which such an honour lias been conferred
on a student of our Universities by the naturalists of our Society,
I appreciate the honour that you have done me. I think, how-
ever, that I should be ungrateful if I did not render some tribute
to those who have given me of the best of their intellect and
of their assistance. I feel that T owe my educational develop.
2 PRESIDENTS ADDRESS.
ment to two men. One has ceased to be a Member of this Society
and to continue his work in AustraHa, but, in a larger sphere, he
still directs the labours and lends his kindly hand to encourage
the early efforts to learn of Nature of those of another land. I
refer to Charles James Martin. The other lives among us and
sets us the example of an earnest student of the animals, plants,
and rocks ai'ound us. 1 pa}^ my homage to Charles Hedley.
You will pardon me if I spend a few minutes in pointing out
what I owe to my chief teachers. The Australians are said to
lack reverence and gratitude. Perhaps their critics are, some-
times, astray. It is rather more than twenty-one years since I
saw for the first time C. J. Martin. I was a student of medicine
at the Medical School in Melbourne, and had reached my third
year. We students had heard, towards the end of our second
year, that the venerable Professor of Physiology, Dr. Halford,
had been granted leave of absence, and that a lecturer had been
appointed in his place. Early in the next year, I went, in the
compan}^ of a fellow student, to find out if we could not start our
year's reading before lectures commenced. The Long Vacation
seemed too lengthy an holiday. We were young in many ways.
We thought that in books lay the knowledge of natural pheno-
mena. We had not learnt that our work in the laboratory was
anything more than an attempt to assist our memories by visual-
ising what we committed to our minds. I recollect that we
went to the Old Medical School through the Grecian porch. We
passed through the bare hall with its tablet in commemoration
of Professor Kirkland, and into the asphalted court. On the
right was the Department of Physiology. It consisted of a theatre
into which opened the Professor's library and a small ante-room.
Beyond stretched a commodious laboratory with benches equipped
for chemical work, with gas, with water, and with shelves
for bottles. A preparation-room in which the Professor's assist-
ant made ready the microscopical sections and chemical solutions
with which we worked, and two empty rooms completed the suite
of rooms. As we walked to the door, I little thought that I was
to spend the greater part of five out of the next six years within
those walls. In answer to our knock, there appeared a tall, fair
PRESIDENTS ADDRESS. S
man in singlet and pants, a file in one hand and a piece of brass
in the other. We asked for Professor Martin, and were curtly
told to say what we wanted. Slowly it dawned on our minds
that we were addressing a new type of teacher. Those we had
formerly known were dignified and distant. We had never
thought of them, even in the heat of summer, unbending so far
as to remove their coats. Many of them never spoke to us unless
covered by their academic I'obes, and we were taught to prepare
for any interview with them by donning our academic attire. A
new era in the Melbourne School of Medicine had already com-
menced. I would not have you think that we students did not
benefit from the instruction of those with whom we had only a
formal association. The}" contributed to our education, but
neither in the same way nor to the same extent as Dr. Martin,
When we two had stated our errand and sought advice as to
what books we should read, we were made not to value ourselves
too highl}" for our desire to start work early by the kindly j^et
satirical praise that we received. We were also informed that
we could not do better than give some weeks to a renewed study
of the microscopical specimens that we had mounted in the pre-
ceding year. As to the books, we might take our choice. We
were to study Physiology and the functions of the living organ-
ism, not any description of them in a book. V\e went away
with little understanding, and somewhat perplexed.
One fact remained in my mind. I had to examine once more
and to draw again the pieces of animal tissue prepared for micro-
scopical study. I devoted the rest of the vacation to that labour,
and I have kept to this day my sketches and those slides. It
was several years before I realised the value of those few minutes'
counsel. Scientific study not only includes the reading of books
and pamphlets about natural objects and happenings, but de-
mands the investigation of the things and of their behaviours.
The books, papers, and monographs are the records of what other
students have noted on these matters. I wonder often if we
make this position plain in advocating more attention to scientific
subjects. We suggest that children should be taught science in
schools. We mean that the children should be given the objects
4 PRESIDEXTS ADDRESS.
to handle and regard so as to become aware of what is known
of them. Do we make the parent and the schoolteacher
understand that education in science is the examination of things
and not of words or descriptions? I fear that many of us who
have some control over education in this State by means of ex-
aminations and the institution of courses of instruction, some-
times forget at what we aim. We demand too large a store of
knowledge. We force the child to learn of Nature by books.
We do not remember how slowly we acquired our own acquaint-
ance with the world of matter about us. We yield to the critic
w^ho tells us that we set too low a standard, and that we do not
ask the child to know enough. We set out examinations with-
out sufficient regard for the hours needed to experiment and to
observe the facts necessary for an answer. I have tried often to
persuade examiners and boards of examiners to give more weight
to practical examinations. I like to ask students to record their
observations on simple objects. M}^ fellow-examiners have said
to me that the questions are so easy. They are easy to the
student w^ho has learnt to stud}" things, and they mislead the
student who does not know of things but only about words.
Such students do not recognise what they see. Life, however, is
concerned with the persons and objects around us, not with the
descriptions of them that literary men may pen.
I have told you that I used my microscope. To those hours
of observation I ow e the interest that Dr. Martin took in my
future education. Some weeks after lectures had begun, I went,
at the close of the instruction, to ask a question about the act of
breathing on high mountains. The conversation passed to
personal topics. I mentioned that I had seen certain appear-
ances in some sections of the lungs. I was invited to come into
the laboratory to arrange the microscopical objects that Dr.
Martin had brought with him from Sydney. Each day I spent
an hour or so in looking at the specimens and putting them in
the different drawers of the cabinets. Now and then, my teacher
came to hear what I was doing or to invite me to look at some
experiment that he was performing in the laboratory. I saw for
the first time a working laboratory engaged in the observation
PRESIDENTS ADDRESS. 0
of physiological plienoineiia. I noted that experiments were
arranged to ascertain how these phenomena were brought about.
I noticed how^ these experiments gave answers, at times precise,
at other times inconclusive, as to the characters of the events
examined. T was led to think of these experiments, to ask
questions myself, to form my own answers from the sequence of
the events of any experiment, and, indeed, to inquire of natural
occurrences how they hajDpened.
I trust that I do not weary you with these reminiscences, but
T think that it would be advantageous to the people of this
C(jmmonwealth if they knew more of the work of that laboratory.
I believe that the citizens of this continent would have derived
more benefit if Dr. Martin had possessed better opportunities of
making others aware of the meaning of scientific education.
Charles James Martin had that personal magnetism which gave
him command over others. He did not preach to us on his
doctrines, but he set us an example in his own attitude to the
science that he taught. During the first year, I was the only
undergraduate that worked daily in the laboratory at hours
beyond those prescribed by the curriculum. Other older men
came to do research at his inspiration. Dr. (later Professor) T.
Cherry, J. F. Wilkinson, A. B. Colquhoun, A. C. Halford, A.
Finch Noyes, and J. W. Barrett (now Sir James) were amongst
the earliest workers. A little later, E. H. Embley commenced
his investigation upon antesthetics, destined to become one of the
principal studies of the laboratory during Dr. Martin's director-
ship. Other undergraduates, among whom were R. A. O'Brien,
now head of the Wellcome Research Laboratories in London, S.
W^. Patterson, M. Lidwill, and, lastly, G. C. Mathieson, endowed
with genius of highest order, who might have influenced the world
for generations, was regarded as one of the brightest stai's of the
rising generation of British scientists, but gave his life at Gallipoli
while acting as a regimental medical ofHcer, came into the labora-
tory in the next few years. All too soon. Dr. Martin's work
among us came to an end. He held no Professorship. An offer
from Belfast had been declined, but he accepted with reluctance
the post of Director of the Li-ster Institute of Preventive Medi-
6 PRESIDENT S ADDRESS.
cine ill London. This post is one of much importance, for the
Institute is tlie largest medical research laboratory in the British
Empire. Dr. Martin did not wish to leave Australia. His heart
was with his students. His work lay in training them to continue
what he had begun. His influence was growing in Melbourne.
The years of sowing, at times disheartening, were almost over,
and the days of fruition and reaping were at hand. An attempt
to obtain for him a chair in the University failed, and one of
the most inspiring scientific teachers that we have had in Aus-
tralia left our shores.
The influence that Dr. Martin wielded, was due to his personal
relations with those around him. He worked with his students
at the same tasks. He asked all to take an interest in his
researches. When graduates or students started their own re-
searches, he came each day to see what was going on. He made
the necessary instruments with his own hands. He manipulated
the apparatus or took the notes. He drew the conclusions or
criticised the inferences. He expected discussion at every step.
It is well, perhaps, that I remind you that he fitted up the work-
shop, setting up tlie benches with his own hands. He repaired
the gas engine that supplied the power. He made the apparatus
before H. J. Grayson became the mechanic. He was carpenter,
fitter and turner, glass worker and electrician for the laboratory,
and for the rest of us. He was the chief under whom we all
delighted to serve. I am afraid that we shall never be able to
let you know how much we stand in his debt.
I would like to add just a few words about his methods of
teaching science. He did not use diagrams or models to any
extent. If he referred to the structure of a muscle-fibre, he
threw on the screen the image cast by a specimen of muscle
through a projecting microscope. The students thus formed
their own mental pictures of natural objects instead of accepting
the interpretation that some other person had embodied in a
diagram or model. Each lesson was a demonstration rather than
an exposition. Experiment followed experiment throughout each
lecture. These experiments were selected to answer the queries
which rose in tlie mind as the result of the previous experiment.
PRESIDENTS ADDRESS. 7
Great skill was manifested in choosing the experiments and in
arranging their order. The sequence of the experiments was
sufficient instruction to most students. The results were so
obvious and the interpretation so easy that most of those who
saw the demonstration needed no prompting as to the inferences
and conclusions. Dr. Martin possessed little oratorical skill.
He was not a successful lecturer. His expositions were by no
means good accounts of all that is known on a subject. It was
necessary to work with him to obtain his instruction. The
student who attended classes with a note-book in which he
intended to write all he meant to learn on a subject found the
lectures most unsatisfactory. The lesson was not prepared for
assimilation. The student found that he had to work out much
for himself. If the student did what was asked of him, he had,
at the end of the course, a training in the science of the func-
tions of the living organisms.
From Charles James Martin I learnt of the inductive and de-
ductive methods of analysis. I observed how exj)eriments were
employed. I saw how to select an experiment so that some
precise inference could be drawn from it. I learnt of physics and
chemistry and of their use in explaining the behaviour of living-
cells, tissues, organs, and animals. To Charles Hedley I am
indebted for a different awakening. I was a teacher of some
years' standing in the University of Sydney before I became
acquainted with Mr. Hedley. He was a neighbour, and our
common interest in this Society led me to cultivate his friend-
ship. I had attended classes in zoology and botany at the Uni-
versity. During my holidays I had wandered through the bush
or meandered over the rocky shore or beach. Many plants and
animals attracted my notice, but I found my instruction had left
me ignorant of what they were and how to learn of them. As
I became acquainted with Mr. Hedley, I formed the habit of
taking walks with him, and I was soon the pupil of an accom-
plished naturalist. The trees, shrubs and herbs, rocks, birds,
and insects had a message. I was led to look at them and see
what they revealed. Until this time, science had appeared to
me to need incessant analysis. I now saw the science of simple
8 FKK!SII)KXT*8 ADDRESS.
observation. Tlie plants seemed to gi\ e me special interest, and
I started to collect those around my home, to notice their con-
struction, and to learn their names from their descriptions. T
was told for the first time that much could be learnt from noting
what plants occurred together, or how different plants favoured
tlifierent localities and surroundings. I need not tell you much
as to ]io\v Mr, Hedley made stone, animal, and green plant of such
interest that I had to look at them. Few of you knew Charles
James Martin, but all of you know Charles Hedley. There
will be no one in this room who has not had some words from
him on natural history, who has not had his attention turned to
some (jbject of interest, and who has not been led by his inspira-
tion to look again at some natural object. No naturalist has
done more for those of us of the younger generation. I think
some of the older naturalists do not realise how ditHcult it is for
the Australian to obtain information about the things around
him. There is an almost complete absence of simple books about
our plants or animals. E^'en the elementary text-books used in
schools are those of European authors, and deal with European
forms. It seems to me that there is here an opportunity which
should not be missed by those among us capable of supplying
this information. AVe need cheap books dealing with the differ-
ent forms of life of our bush, beaches, and waters. 8uch books
as ha\ e been prepared cover a wide range. They deal with the
birds, fishes, animals, or insects of Australia. I believe there
would be a ready sale for cheap books dealing with the fauna and
flora of small districts. When I started to examine the flower-
ing plants about my home, T read once again Hooker's Botany,
and then used Moore's Flora of New South Wales. I will not
say how many hours I spent in discovering the name of a plant.
No beginner will find it easy if he has to use Moore's Flora.
The classification of plants depends on characters which are not
readily ascertained. After several weeks, I named Bauera
7'ubioides, and was materially aided by the presence of eight
petals. Observ^ation seems easy to those who have not observed.
I cannot i-emember how often it became necessary to revise my
description of the characters of a plant. I used to write them
PRESIDENT S ADDKESS. 9
out as I recognised them in tlie })]knt. Wlien my account was
completed, I started to find the family, the genus, and the species.
If T found the correct genus, the rest was comparatively simple.
How I wished for a book with some figures or diagrams of the
fioral characters of the principal families of plants found in our
bush! From these laboui-s, I learned to appreciate the value of
precise observation. T have mentioned plants, Ijut our rocky
foreshores are alive with animal life, and no one but a specialist
can learn of them. Butterflies and moths attract many. We
have a useful book about butterflies, and we would all welcome
others treating of other kinds of insects. May I hope that the
day is not very distant when ten or twenty of these useful books
will have been written.
On this occasion our th<nights may fittingly turn to our absent
soldier-members. In addition to those mentioned at the last two
Annual Meetings, Mr. E. Griffiths, B.Sc, of the Department of
Agriculture, Mr. Tilley, of the Geolc^gical Department of the
University of Sydney, and Mr. R. B. Oliver, of Auckland, New
Zealand, have departed for the front during the year, or are pre-
paring to do so. Not only have we not had the pleasure of
N\elcoming any of them .back during the year, but we haxe heard,
with sorrow, of the death of Dene B. Fry, one of our promising
junior Members.
Four Ordinary Members lune been elected during tlie Session,
two resignations have l)een received, and three Members ha\e
been lost by death.
Dene Barrett Fry, killed in action in France, on April Uth,
1917, aged 23, was the first of our Soldier-members to fall. He
was a rising young biologist of great promise, elected a Member
in 191. "3; and his loss is greatly to be dej^lored. His training-
began at the Australian Museum, as a cadet, in 1908, where he
i-emained until 1914. When the war broke out, he was a student
at the University, and a demonstrator in zoology; but he gave
up his University work in order to enlist, joining the Army
Medical Corps in May, 1915, After two voyages in a hospital-
ship, he transferred to the Infantry, qualifying for the i^ost of
lieutenant. But as there was no vacancy available, he left for
10 president's address.
the front with reinforcements, as sergeant. After some time
spent at Sahsbury Plains, his regiment was sent to France, where
he took part in several engagements. His last contribution to
science, "Description of Aphantophvyne^d, new Batrachian Genus
from New Guinea," and his only one to our Proceedings, ap-
peared in the volume for 1916. Ten other papers dealing with
reptiles or amphibia wei-e published in the Records of the Aus-
tralian Museum (1912-13), in the Memoirs of the Queensland
Museum (1913), in the Records of the Australian Museum (1913-
14), and in the Proceedings of the Royal Society of Queensland
(1915).
Dr. Thomas P. Lucas, one of our Senior Members, elected in
1861, died in Brisbane on November 15th, 1917, in his 75th
year. He came out to Australia in 1876; and after practising
his profession in Melbourne for ten years, he removed to the
warmer climate of Brisbane, where he continued to practise up
to the time of his last illness. He was especially interested in
Lepidoptera, an enthusiastic collector, and the possessor of a very
fine collection.
Alfred John North, another Senior Member, elected in 1886,
died on May 6th, 1917, aged 61. He came to Sydney from
Melbourne in 1886, and became associated with the Australian
Museum, as x\ssistant in the Zoological Department, to under-
take the preparation of a "Descriptive Catalogue of the Nests
and Eggs of Birds found breeding in Australia and Tasmania"
(Catalogue No. 12) which was published in 1889. In 1891 he
was appointed Assistant in Ornithology. He was the author of
numerous papers published in the Records of the Australian
Museum, and other Journals, including the Society's Proceedings.
But his most important contribution to ornithology was Special
Catalogue No. 1 of the Australian Museum, being a second and
much enlarged, well-illustrated edition of Catalogue No, 12,
entitled "Nests and Eggs of Birds found breeding in Australia
and Tasmania," in four quarto volumes (1901-14). Mr. North
was a keen observer of bird-life as well as a voluminous writer;
and he frequently exhibited specimens of interest at our Meet-
ings over a long period.
president's addr'ess. 11
The year's work of the Society's research-statf may be sum-
marised as follows: — Dr. R. Greig-Smith, Macleay Bacteriologist
to the Society, contributed two papers during the Session, which
have already appeared in Parts i.-ii. of the Proceedings for 1917.
One, the fifteenth of his series of contributions to a knowledge
of soil-fertility, dealt especially with the action of certain micro-
organisms upon the number of bacteria in the soil; and the other
was descriptive of an improvement in the technique of the single
cell cultivation of yeast. He also completed the sixteenth
of the series already alluded to, which is concerned with the
search for toxin-producers. Certain soil-bacteria, moulds, and
amcebi«, all reasonably supposed to be capable of furnishing sub-
stances of a toxic nature, have been grown in various media and
under varying conditions, and, in all cases, the signs of toxicity,
which become manifest, can be attributed to an alteration in
in the reaction of the media. The effect of reaction is quite of
a different order from the evidence of toxic action obtained in
former researches.
Dr. Petrie, Linnean Macleay Fellow of the Society in Bio-
chemistry, contributed three papers, which are contained in Part
i. of last year's Proceedings. One of them was a continuation of
his investigations on the occurrence of hydrocyanic acid in plants
(Part iii.), and furnished a record of some new cyanogenetic
plants. The other two treated, in an exhaustive manner, of the
chemistry of the Duboisias, and the alkaloids of Duboisia Leich-
hardtii. Additional subjects which occupied his attention were
certain cpiantitative problems of cyanogenetic plants, the de-
termination of the amount of hydrocyanic acid capable of being
obtained from these plants by fermentation, and the isolation of
the glucoside of Ileterode^tdroii, one of our most fatal stock-
poisons.
Mr. E. F. Kallmann, Linnean Macleay Fellow in Zoology, con-
ti'ibuted a paper descriptive of the genera Echinaxia and Rhah-
dvsignid, which was published in Part ii. of last year's Proceed-
ings. He also had under consideration a paper embodying
a reform in the classification of the Deamacidonidce, a rather
troublesome family to the systematist. At the end of the month,
12 president's address.
Ml'. Halliiianii gives up his Fellowship to engage in other work.
We wish him all success in his new sphere.
Mr. K. J. Tillyard, Linnean Macleay Fellow in Zoology, con-
tributed eleven papers during the Session, of which seven have
already appeared in Parts i.-iii. of the Proceedings for 1917; two
others will be found in the forthcoming Part iv. of the same
volume: and the remaining two will be read at an early Meeting.
Three of these papers dealt with the Odonata, four with fossil
insects, one each with the Orders Jvepidoi)te]'a, Planipennia, and
Mecoptera respectively, and one with the neuropteroid fauna
of Lord Howe and Norfolk Islands. The researches upon
the breathing-apparatus of the larva* of Odonata were com-
pleted ; the work begun upon Australian fossil insects made
considerable progress ; and a start was made with the study
of other neglected Orders. A considerable amount of work was
carried on in the investigation of the Orders comprising the
"Panorpid Complex," namely, the Mecoptera, Planipennia,
Megaloptera, Trichoptera, Lepidoptera, and Diptera, with a
\iew to the elucidation of their phylogeny, and inter-relation-
ships. In making a careful study of the Jugate Lepidoptera,
in this connection, ISh: Tillyard recently made the important
discovery, that five genera of the Family Micropterygida' have a
wing-coupling apparatus not of the jugate type found in llejna-
Hdce, as has hitherto been supposed, but of a more primitive,
jugo-frenate type, closely resembling that found in the Plani-
pennia and Mecoptera. On the hindwing, near the base of the
costa, there is a strongly developed frenulum of from two to six
l)ristles, which becomes engaged, during Hight, in the sinus
formed between the base of the dorsum of the forewing and the
so-called jugum; this latter is bent under the forewing, with its
apex pointing outwards and forwards, and acts as a retinaculum
for the frenulum, and not in any way as a jugum or "yoke" for
the costa of the hindwing, as it does in Ilepia/ida.
Dr. H. S. Halcro Wardlaw, Linnean Macleay Fellow in Physi-
ology, completed the first part of an elaborate study of the
variability of cows' milk, which will appear in the forthcoming-
Part of last year's Proceedings. This dealt with the variations
pfnvStdent's address. 13
and properties of samples of afternoon-milk from 109 normal
cows, kept in connection with a Government Institution solely
for the supply of tlie inmates, and not for profit. It is of interest
to note that about 40 per cent, of the samples contained a per-
centage of solids not fat below 8' 5 — the standard for the per-
ceentage of solids not fat in all the States of the Commonwealth,
and in England. Dr. Wardlaw has in contemplation the con-
tinuation of this important investigation, the extension and con-
tinuation of some ^^■ork of a similar character upon human milk
carried out by him in 1914, and to subject to a more detailed
examination certain questions suggested by his work on the
chemistry of respiration.
In response to the Council's invitation for applications for
Fellowships, 1918-19, only three applications were received; and
I have now the pleasure of making the first public announcement
of the Council's re-appointment of Dr. J. M. Petrie, Mr. R. J.
Tillyard, and Dr. H. S. H, Wardlaw to Linnean Macleay Fellow-
ships for a period of one year from 1st proximo: and, on behalf
of the Society, of w ishing for them a fruitful research-period.
The concluding Part of the forty-second volume of the Pro-
ceedings is nearly ready for issue. The complete volume (926pp.,
50 Plates, and 196 text-figures) will contain thirty-three papers,
two of which have been divided for convenience on account of
their length. Six papers exceed 40pp. (43-82 pp. in length); and
sixteen of them were contributed by members of the Society's
research-staft'.
The war's interference with our exchange-relations w^ith
Societies outside the Commonwealth, continues in an aggravated
form. We have heard from fewer Societies, and there has been
a marked falling off in our receipts during the year. Even
Societies in the United Kingdom are finding it advisable to sus-
pend their despatches for the present, in order to avoid the risk
of losses. The total number of exchanges received during the
Session 1916-17 amounts to 846 additions to the library, received
from 136 Societies oi* Institutions, ttc, and five private donors,
as compared with 1285 (1913-14), 1028, 1243, for the three pre-
ceding Sessions, The Society is much indebted to the Public
14 president's address.
Library of New South Wales, for a special donation of some
partial sets of Anthropological publications, comprising 34 com-
plete vols., 37 incomplete vols., 39 Bulletins, and 11 Reports,
issued by twelve Societies, not previously contained in the
Society's Library.
If I have spent some time on personal topics, I liope that you
will pardon me. They will serve to turn your attention to two
different kinds of scientific knowledge. The educationists of this
State have laid stress on the analytical sciences, mathematics,
physics, and chemistry, and have, perhaps unwittingly, under-
valued the biological sciences. The student in our Universities
is forced to learn the analytical sciences, and is handicapped if he
wishes to study zoology, botanj^, and geology. Some have advo-
cated extending this handicap to the teaching of these sciences
in schools. I am still a young man, and I remember well the
frequent expression of the wish that the student entering the
University should have had no previous instruction in science.
A liberal education at school with no science was to be the
prelude to specialism consequent on learning a little chemistry or
physics. The people of this State owe to Mr. Peter Board,
C.M.G., Director of Education, the adoption of that great prin-
ciple of toleration, of suiting the subjects to the inclination of
the child. Instead of trying to teach all children the same
subjects, the child may select a certain proportion of the subjects
to be studied. Lately an attempt has been made to influence
the child's selection by giving different marks to different subjects
at the examinations for leaving certificates. As free education
at the University is granted on the results of this examination,
it is obvious that those desiring better education will only learn
subjects carrying the highest numbers of marks. What we seek
in this State at the present time is the opportunity to study the
natural objects that interest us, without hindrance. When we
plead for freedom in the choice of subjects in education, we
desire that those in schools may be taught those subjects which
attract their attention and hold their minds. We protest
against the suggestion that any study of natural objects bespeaks
a worship of materialism destined to wreck the progress of our
president's address. 15
evolution. A brighter day for science has already dawned
within our University. The unfair influence exercised upon the
boy or girl leaving school and entering the University, b}^ the
scanty recognition granted to any study of scientific objects, has
been done away with. It is seen that it is possible in Australia
to train students well in science without forcing them to go to
Europe and become bound by European traditions. The force
of these traditions is seen in our University with its mediaival
architecture. The scientific departments of British Universities
are housed in badly lighted rooms because they have had to
accept the buildings handed to them from the Dark Ages. In
this country, it is ridiculous to build a laboratory for micro-
scopical study, in which many students have to work, and fit
it with narrow, arched Gothic windows because our forefathers
revered such building. In Great Britain tlie modern laboratory
is built to facilitate the investigation that is to take place within
it. Here we are told that artificial lighting is most suitable for
microscopical study, but are left to get what light we can from
long, narrow windows twenty to forty feet away from the mirrors
of the microscopes.
May I remind you what a small company we are here in New
South Wales. We number among us practically all those dwell-
ing in this State who are engaged in increasing our knowledge
of the living objects of New South Wales. Surely we should aid
each other in learning of our flora and fauna. Some seem to
consider that our work consists in writing and publishing the
results of their own inquiries. The papers embodying these
observations and investigations appear in our Proceedings. They
may be read in the printed parts. There is another and quicker
method of imparting information. When a Member describes
to us his work, his personality adds interest to his description,
and we easily become acquainted with his results. A few well
chosen remarks on an exhibit will cause every one to examine
what is shown to them. We would all be pleased if every Member
attending our meetings would bring some object of interest to
himself and explain that object and its interest to his -fellow
Members. In this way we can ail acquire a better acquaintance
16 president's address.
with natural history. Discussion leads to clarity in our ideas,
and assists in formulating our scientific conceptions. During
last year, we had some discussion on the characters of hybrids.
The discussion was brief, but it helped us to understand better
how we were to recognise hybrids in the plants around us We
should not be always waiting for those in Europe to lay down
the principles on which we are to work. These are problems
which can be studied more readily in Australia than elsewhere.
The separation of Australia from the other great land-masses and
its isolation for long epochs have affected the characters of our
animals and plants. I hope that our Council will promote dis-
cussions of the general questions to the stud}" of which contri-
butions are made in the monographs of our Members. Two
such discussions might well be arranged each year.
If I have spent some time in making more precise my concep-
tion of the essential f«^atures of scientific work, I have been
desirous of ensuring that there is no misunderstanding upon this
point. We are now in the fourth year of the insensate war
which is devastating more and more of Europe. We recognise
that we must utilise our resources t)f men and material in the
most economical and advantageous manner if we are to emerge
victorious from the struggle. We have learnt that it is our duty
to give the best of our manhood in the prime of life for the
service of our country, not counting what the sacrifice ma}"
involve personally. AVe give our savings to provide for the cost,
which shows still a tendency to increase to a greater daily ex-
penditure. We endeav^our to do what we can in personal service
to assist in defeating our enemies. We have heard much of our
duty to our country, but we ha\e heard very little of the duty of
the Government of this Commonwealth to use men, money, and
training to its full extent. I suppose that no one asserts that
this war has revealed the ability of tlie Government to place
each man and woman whose services have been put at its dis-
posal in the position in which the training and experience of the
individual can be fully employed. I have looked along this table
from month to month and thought how few of our Members are
utilised in ways in which their special ti-aining can be of service
president's address. 17
to the community. Surel}^ it cannot be true that the years of
observation and experiment have made our members useless to
the Department of Defence. I am aware that the habit of
casting ridicule upon Dr. Dry-as-dust existed in the past. I
know that it is still common to hear jokes at the expense of the
naturalist, but I thought that those raised to authority were
acquainted with the fact that science can play a part in every
phase of work in Australia in connection with the War. Some have
affirmed that the recognition of the importance of Science will
be one of the permanent benefits conferred on our nation by the
war. It is perhaps difficult to imagine any benefit given us b}^
the war, which has caused such unspeakable misery and sorrow,
yet we can understand the statement. Whether there is any
greater use of Science made in Europe, I cannot say from personal
knowledge, but-I can remind you that there are few evidences in
this country of a much greater employment of those possessed of
scientific training. It is true that the Federal Government is
using the services o'ratis of a small number of scientists who
devote the time that is not taken up with other duties to some
investigation. The Council of Science and Industry supplies some
money towards the cost of this research work. This is a step in
the right direction, but that is all. When we sa}^ that greater
use should be made of Science, we do not mean that some par-
ticular method should be employed, but that every problem
should be studied by the means adequate to give the solution.
In connection vvith the war, we find much evidence of the failure
to use scientific methods. The whole number of those who have
spent time and trouble in acquiring knowledge of Nature is a
very small fraction of the population. Even this small number
has little to do. It has been said that it looks as if our
authorities regarded scientific training as a disqualification rather
than as a qualification. I do not suppose that there is any desire
not to use the experience and learning of scientists, but I do
think that the majority of our community are quite unaware of
the scope and character of scientific knowledge. The scientist
cannot replace the practical man who is managing a business or
dealing with some particular problem, but he can supply informa-
2
18 president's address.
tion, and be can make observations to provide facts on wbicb tbe
practical man can work. A scientist does not make, as a rule,
wbat is known as a good business man. I think it unlikely that
he ever will, but I think a study of the uses made of Science in
the last century will teach any one who will examine the question
that information obtained by scientific study has been of in-
calculable value to the community. Our Government does not
especially need advice and direction from scientific persons, and
there are so many ready to offer this. It needs scientific workers
who will obtain accurate information by making studies on the
problems of the hour. Each scientific worker can only do a little.
It takes a long time to make precise observations and to conduct
experiments. Let me take an illustration. There are millions
of bags of wheat stored in Australia. Insects, fungi, and other
forms of animal and plant life are destroying that wheat. If
every zoologist now^ in Australia worked for a century upon that
wheat, there could be no thorough examination of the whole of
it. To examine the contents of even a million bags would require
a very great time. To determine where an insect or fungus
came from, how its grow^th was favoured by various circum-
stances, wdiat is the life-history, and at what stage in its devel-
opment can the destructive agent be destroyed so that it does
not reach the wheat, would take many months. I do not want
you to think that I exaggerate. May I remind you that it
took many men and many years to obtain some information
about the mosquito in its relation to malaria, that to-day we
are not in a position to prevent those of our soldiers who go to
New Guinea and elsewhere from acquiring the disease by the
bite of the mosquito, and that it has taken several years to obtain
even a little knowledge of the different mosquitoes scattered over
a small portion of Australia^ There is an opportunity for every
zoologist in Australia with some acquaintance with entomology,
and for every botanist with some knowledge of fungi, to be of
service in saving some of that wheat. Is there no way in which
the Defence Department can be informed of this simple fact? Is
it better that many of these trained zoologists or botanists should
be doing none of that work for which they are specially fitted in
/ president's address. 19
relation with the war. It surely cannot be true that we have
such a vast surplus of food that it is unnecessary to take tliese
steps to preserve this wheat against loss. Further, it cannot be
true that we know so much of the insects and fungi in this wheat
that further knowledge is unnecessary. I believe that the De-
fence Department has already at its disposal the services of those
who can advise it what to do when the knowledge of what insects
and fungi are present has been obtained. Ma)^ I invite all the
Members of this Society to assist in making these facts known?
I have often thought that the scientific experts in the State
and Federal services do not make the authorities, under whojn
they work, aware of what a small amount of scientific investiga-
tion can be done by one person. We have a State Entomologist
who sets us an example of industry and devotion in the way in
which his whole life is dedicated to the study of insect life. I
am, however, amazed at the range over which his advice is sought.
He is expected to report and advise on matters in a few weeks
when months of research would probably only serve to reveal
the nature of the problem to be attacked. Even if we had
twenty entomologists as learned as Mr. Froggatt in the ways of
insects, we would be aware that all the twenty naturalists could
go on working for many years without exhausting the possibilities
of research in insect life. Does not the work done by all the
entomologists of the world in the century that has passed onl}^
serve to show us what remains to be done? Are we not all like
Newton, when he remarked that he was as an infant playing
with a few grains of sand with the whole ocean before him yet
to be explored?
The community has little appreciation for the laljours of those
seeking the advancement of knowledge of Natural Phenomena.
The man who could supply a pound of tea to every person on
this globe of ours would gain an immense fortune. Lister, who
did something to diminish the suffering of every living person,
and who was, perhaps, better rewarded than most other scientific
men for his discoveries, had much less reward than any really
successful man of business. Have not scientists some right
to ask more encouragement from their fellows? Most of us
20 president's address.
have made our scientific observations in our leisure time. Even
those of us employed in our Universities soon learn from
those under whom we work that teaching and not scientific
inv'estigation is what the University demands. We learn that
it gives greater satisfaction if no scientific research is done, since
the absence of scientific investigation shows that there cannot
have been any leisure time in which it could be done. It is only
a few years ago that the Senate of the University of Sydney
was informed by the President of the Professorial Board that
only those with leisure from teaching duties did any research
work. The attitude of official discouragement should be re-
moved. It seems to me desirable that young graduates should
be promoted more often w^hen they do research work. Some
have seen those who have never made a single original observa-
tion placed above others who have spent three or four years in
research after graduation. We often find the attitude that it
were better that a problem be not studied at all than that we
should study it. I once suggested to the late President of the
Board of Health that it would be a fortunate day when every
town in New South Wales was the home of one biochemist making
investigations in respect to the people's food. To my surprise,
his answer expressed the hope that not a single biochemist
would meddle with the affairs of the Department of Public
Health. The opinion is held by many holding positions in the
educational and professional services that it is better that
scientific research should not be done at all than it should be
done by Australian graduates or by those trained in this country.
Many of the Members of this Societ}^ will have had experience
of the contempt for their capacity, training, and public utilitj-
exhibited by some holding high official stations in our University,
our Department of Public Instruction, our Civil Service, and our
learned professions. Scientific research is little I'egarded. Few
attempts are made by the official bodies connected with scientific
work to assist the progress of investigation. The research
student finds that it often takes longer to fight for the material
with which he wishes to work, in order that he may have the
opportunity to conduct his research, than to carry out the actual
pI{p:sident's address. 21
Avork uf investigation. The research student learns tliat his in-
vestigation is regarded as a personal matter, and that the time
taken in its performance is considered misspent, since some of
the experiments may have been done in the hours for which he
is presumably paid by the University or State. Australian
graduates are considered by some of the official heads in our
Universities as required only for the "teaching of the drill-work."
The graduate who does research work rather than this drudgery
need expect no promotion in some of the Departments of our
University. Surely this is a poor policy in our educational
development. Ability to do research work is not commonly
found. The desire to carry it on continuously is even less fre-
quently present. The power of directing investigation in others
to a successful conclusion is one of the rarest accomplishments.
These facts are well known to the members of the Senate of our
Universities, to the heads of our Departments of Health, Mines,
Agriculture, and Education, to the members of the State Com-
mittees and the Executive Committee of the Federal Advisory
Council of Science and Industry. Why is it so difficult to pass
from opinion to practice in these matters? Why is there not
a better attempt to ensure that every scientist in ISTew^ South
Wales who shows that he can train students to become more
efficient investigators, has as many students as possible brought
under his influence? Not infrequently after years of labour in
building up a laboratory and a team of workers, the investigator
is driven to cast aside what has been done, to provide himself with
an income sufficient for his needs.
The naturalist is not only interested with the structure of
dead animals and dried plants, but he spends his time in learning
of the behaviour of living animals and plants. Living animals
and plants seem endowed with attributes not found among inani-
mate objects. Life is associated with a definite form of con-
struction. If we cut a thin slice of a rock and recognise that
the rock is built up of inorganic material arranged to show a
body-wall and nucleus, such as are found in vegetable or animal
cells, we know that we are looking at the petrified remains of
what was once alive. Not only so, but if we find any substance
22 PRESIDENTS ADDRESS.
composed of certain chemical compounds, as proteins, carbo-
hydrates, or fats, we are aware that this material has been part
of the animal or vegetable body. Life is thus associated with a
particular architecture built with characteristic chemical sub-
stances. AVhile the qualities and properties of living things ap-
pear so different from those of lifeless matter, the biochemist soon
discovers that all living things are constructed from lifeless sub-
stance. As an animal or plant grows, it adds to its bulk material
from the inanimate world. All the carbonaceous material in the
protoplasm of the cells of all living objects contains carbon once
diffused through the atmosphere as carbon dioxide. For a brief
period this carbon is found in some animal or plant. Perhaps it
in the green leaf of a growing wheat-plant. A few weeks later,
it lies in the golden grain. The farmer harvests that grain and
sends it to the miller. The same carbon is part of a particular
sack of flour, delivered to a baker. It is portion of some loaf of
bread eaten by John Smith. It is the carbonaceous constituent
of «iluco8e in the blood of John Smith. It is combusted in some
part of John Smith's body, and it is dissolved as carbon dioxide
in his blood. It diffuses through the epithelium of his lungs. It
goes out again into the atmosphere in the expired breath of the
individual. Some day it will be taken up by another plant, and
so on in an endless cycle. It is difficult to suppose that this
carbon becomes endowed with new qualities while it lies in the
living tissues <jf the animal or plant. It is necessary to seek some
other explanation of the characteristic properties of living things.
It has long been obvious that animals and plants make the
chemical constituents of their bodies from raw materials by dif-
ferent processes from those used in the chemical laboratory or in
the factory for the manufacture of similar substances. Heat and
pressure, so freely used in the synthesis of bodies akin to those
found in living things, play no part in the manufacture of starch,
fat, or protein in a plant. We have a dim idea that those in-
tangible materials, known as ferments, are chiefly concerned with
vital synthesis. In fact, the presence of ferments and the capacity
to produce ferments appears to be one of the most distinctive
chemical characters of living matter. Ferments bring about
president's address. 23
decoDipositions and incite combinations under the conditions
favourable to the continued life of animals and plants. We sow
a wheat-seed, presumed to contain minute amounts of a number
of these ferments, and from the fully grown wheat-plant we can
obtain a greater degree of ferment-activity. The powers of these
ferments do not differ from those possessed by lifeless chemical
substances under certain conditions. Most chemical substances
appear capable of bringing about decompositions or combinations
in other bodies in the same way and under analogous circum-
stances as do the ferments from living tissues. When acting in
this manner, these substances are called catalysts. Chemists do
not, however, suppose that platinum black inciting the union of
hydi-ogen with oxygen contains some unknown ferment. The
sulphuric acid which brings about the decomposition of cane
sugar into glucose and hevulose is presumed to be chemically pure
sulphuric acid. Chemists are beginning to recognise that cata-
lysts are not a group of substances possessed of these special
qualities, but that many, if not all, chemical substances can act
as catalysts under particular conditions. Evidence is accumu-
lating to show that it is on the arrangement of the particles,
molecules, or whatever we may call the minute masses of the
substance, that the capacity to act as a catalyst dej^ends. The
physical state of a substance rather than the kind of matter of
which it is formed, confers catalytic activity. There appears to
be no reason to infer that the catalytic action of the extracellular
and endocellular ferments found in animal and vegetable tissues
has a different basis. I have always been impressed by the fact
that the powei- of an extract of an animal or vegetable extract
to act as a ferment was abolished by those agencies which pro-
foundly alter the colloidal state of proteins and lipoids. The
attempts to identify the unorganised ferments as a special class
of chemical substances have invariably ended in failure. May
we not look for better progress along other lines of investigation?
The study of the position of the particles composing inorganic
crystals has given us an insight into the arrangement of the
atoms or ultimate particles in the molecule. The crystals of
24 presidp:nt's addrp:ss.
proteins and fats have not yt^t been examined in the same way.
May we not hope that these studies may be ext(ended in con-
nection wdth fermentative activity?
The functions of living matter have been the object of that
branch of science in which I have been a student. The aim of
the physiologist, is to explain the behaviour of living things in
terms of the mechanical, physical, chemical, and psychical sciences.
It is believed that progress is at present possible if an endeavour
is made to reduce biological phenomena to physical and chemical
problems. Living things are built from inanimate matter, and
the attributes of living things are the expressions of the pro-
perties of matter. I am aware that such a belief is condemned
as materialistic. I must confess, however, that I see no diffi-
culty in supposing that, as heat, light, magnetism, and electricity
are qualities of the arrangement of matter, the psychical qualities
of the higher forms of living things are likewise material.
Heat apart from matter does not exist. Why should we suppose
that memory and thought exist apart from some matter in the
living organism ? Whoever will become acquainted witli the
history of physiological knowledge will find that progress is
represented by the explanation of a greater number of the func-
tions of animals and plants in terms derived from the study of
physics and chemistry. On such a conception, there is no sucli
thing as inanimate matter, though there may be lifeless objects.
With these ideas in your mind, I invite your attention to the
problem of variation as exemplified in the different functions of
each kind of animal and plant. For many centuries, naturalists
had believed that each kind of animal or plant was endowed
with unchanging characters. The fixity of species was con-
sidered axiomatic by zoologists and botanists. Darwin and his
followers have demonstrated to us the progressive evolution
from one form to another in animal and plant. Every animal
and plant showed a certain degree of variation in its characters.
Some of these characters favoured the living organism in its
struggle for existence, others rendered it less able to cope with
the conditions of environment. Some organisms survived longer
and had a better chance of reproducing their kind, others died
president's address. 25
early and bad less opportunity for transmitting their peculiar
qualities to their descendants. It is well known that the off-
spring of any organism possess, in general, similar external
characters and biochemical properties to their parents. The
mechanism by which the resemblances and variations were
brought about received little study in the last centur}'. Indeed
it was held by most biologists that environment had little in-
fluence on the qualities of the different generations of any race,
however much it might affect the individual in a particular
generation. It was denied emphatically by the Darwinian evo-
lutionists that the conditions of the environment could produce
new characters in the descendants of any individual. While
naturalists, in general, held these views, there were notable dis-
sentients from these doctrines. Physicians, who have many
occasions for observation in this field, have in general believed
that some acquired characters can be transmitted to the offspring.
The progress of knowledge is showing us that the question of
the inheritance of acquired characteis no longer interests this
generation in the form in which it attracted the previous genera-
tion A wider acquaintance with the qualities of a living thing
has led us to have a more comprehensive idea of inheritance.
Differences that seemed insurmountable to older generations are
no longer differences to us. We can produce an artificial immu-
nity to some particular chemical substance, and we find that this
immunity can be transmitted permanently to the race. We can
cause an animal or plant to acquire new characters, which seem
to become fixed characters in succeeding generations. We find
that the characters of animals and plants are much more labile
than had been supposed, and that it is a comparatively easy
matter to engraft some new qualities upon them. There are
limits to what can be done, but we have not yet fathomed what
can be accomplished. It may be of interest to you to consider
how some of these changes can be produced.
We may take as an example the control of some of the quali
ties of micro-organisms which are known to have such intimate
relations with the existence of more differentiated animals and
plants. The behaviour of yeasts in connection with the conver-
26 president's address.
sion of glucose into alcohol and carbonic acid gas, the so-called
alcoholic fermentation, will serve for this purpose. The ferment-
ation of sweet liquids has attracted some attention from every
generation of natuial philosophers. The slow accumulation of
real knowledge is illustrated by the fact that we have no record
of any observation, current in our descriptions of the process,
before the end of the seventeenth century. Until that date, it
was thought that the spirit of the liquid made its escape in the
heaving struggle of the bubbling liquid. Van Helmont identi-
fied the gas vinorum escaping from the fermenting liquid with
the gas sylvestre found in woods and caves or freed from marble
by acids. Becher observed that only sweet liquids underwent
this change. Leeuwenhoek, indeed, saw the countless myriads
of particles with his microscope, but did not connect their
presence with fermentation. Another century rolled by before
Lavoisier made use of the balance, and discovered that sugar is
split into approximately equal parts of alcohol and carbon
dioxide. Cagniard Latour, using a better microscope, noted in
1831 that the yeast particles multiplied by budding, and that
they were living plants. He suggested that the growth of these
plants occasioned fermentation. Theodore ISchwann came to
the conclusion that fermentation was brought about by the
entrance of living micro-organisms into the fermenting liquid.
Liebig denied the relation of the yeast-plants to the process of
fermentation. He drew a humorous picture of the yeast cell
equipped with its still and condenser, furnishing alcohol and
blowing off gas. He asked whether such scenes were really
visible through the microscope. Later he accepted the yeast-
cell as a living plant, but affirmed that its presence had nothing
to do with the occasioning of fermentation. Pasteur showed
that the sugar was split in the body of the yeast-plant, and
attributed this decomposition to the vital powers of the living
cell. Buchner demonstrated that an unorganised ferment could
be separated from crushed yeast-cells, and that this lifeless body
decomposed glucose into alcohol and carbon dioxide. It has
taken more than two hundred and fifty years to gain this infor-
mation. To those of you who know little of fermentation, our
president's address. 27
knowledge seems very complete. The agent for decomposition
is made in the body of the yeast-plant, in which it remains. Tlie
sugar enters the yeast-plant and is split into roughly equal
weights of alcohol and carbonic acid gas. To those who have a
more extended acquaintance with these objects, it is obvious
that much more has to be learnt. We are not aware how the
sugar is split into alcohol and carbon dioxide. The arrange-
ment of the carbon, hydrogen, and oxygen in glucose does not
make it easy to imagine a decomposition into alcohol and carbon
dioxide. Further, we have some knowledge that, without other
agencies, such as the presence of salts of phosphoric acid, no
decomposition of sugar can be accomplished by the ferment.
Time will undoubtedly reveal other aspects of this phenomenon
at present entirely unsuspected.
The process of fermentation has not only a scientific aspect.
It forms the basis of some manufacturing industries of great
antiquity. The preparation of wines, beers, and spirits is an
application of this fermentation. Yeast is also used commer-
cially in the manufacture of wheaten bread. In making bread,
bakers mix Hour, water, and a little salt with a small quantity
of a specially prepared yeast at 28*3°C. The dough thus formed
is kept warm, when it swells to three times its original volume.
At the present time, doughs stand about nine hours in the
troughs to ripen. With a greater amonnt of yeast, doughs ripen
more quickly, and the time of waiting for the distended dough
can be diminished. It is also found that a quantity of one yeast
may be much more efficient in liberating gas in the dough than
the same weight of another yeast. The yeast obtained by tlie
brewers of beer in such large quantities, will not ripen a dough
in the time required by a baker, even when used in large quan-
tities. A yeast has to possess special characters if it is to bring
about alcoholic fermentation rapidly in a baker's dough. At
one time it was supposed that some strains or races of yeast
were capable of multiplying freely in a baker's dough, while
others could not breed quickly under these conditions. It was
supposed that it was necessary to search for these yeasts in
nature and cultivate them. The capacity of fermenting doughs
28 president's addkess.
at some suitable speed was considered a fixed cliaracter of each
race of yeast-cells. It was thought that the cultivation of the
progeny of an individual possessed of this quality would produce
a race with the desired properties. In practice, bakers found
that it was impossible to continue the cultivation of a particular
strain of yeast, as the fermenting power appeared to undergo
deterioration. Ever\^ few weeks, a baker started a new stock of
yeast. Failure in maintaining suitable yeast by continued culti-
vation was ascribed to contamination of the stock by unsuitable
yeasts. When I started my study on yeasts for the Advisory
Council, I thought that my task was to isolate suitable races of
yeast-Cells and then to cultivate them under conditions which
would prevent contamination with uiisuitable yeasts. T hoped,
indeed, to isolate the strains that I needed by physiological
means rather than by bacteriological separation and the routine
examination of the progeny of hundreds of individuals, but I
did not imagine that I was going to find that any race of yeast
could be taught in a few days to ferment glucose quickly in a
baker's dough. Without going into details, I may say that I
found that any yeast of the species Saccharomyces cerevisics,
growing under certain conditions in a baker's wort, caused rapid
evolution of carbon dioxide in a dough. These conditions, as
far as I could define them, were a temperature less than that of
dough, aeration with a plentiful supply of air, the presence of
certain extractives from flour, the presence of certain substances
from hops detrimental to the growth of acid-forming bacteria
and a particular concentration of the sugar and other products
of malted starch. These conditions were so favourable to the
growth of yeast that in the bakehouse of the Technical College,
Sydney, Mr. F. Elliott was able to keep one strain of yeast for
making bread for almost six months. This yeast was so active
that two and a half ounces of the moist yeast ripened four
hundred and ten pounds of dough in six hours. If any of the
conditions in the wort were altered, the yeast rapidly became
quite unsuited for making bread.
The manner in which the yeast is selected may be conceived
in two different ways. Each yeast-cell buds off' daughter-cells
presidejst's address. 29
Some of these may possess a greater capacity for fermenting
glucose in a dough than others. The conditions may favour a
more speedy multiplication of these desirable cells. In this %vay
the resulting yeast would be made up of a larger number of cells
capable of inducing fermentation in the dough. On the other
hand, we can suppose that the yeast-cells all become so altered by
their environment that they can ferment the glucose in the dough
more rapidly. It is not possible to explain with certainty just
what happens in this process. By cultivation from a single cell,
yeast suitable for a baker can be obtained. It is therefore clear
that the desirable yeast may be selected from the progeny of a
yeast not endowed with the required qualities. A similar
selection is made in the dough. This explains the rapid ripening
of doughs made with "sponges." The elucidation of what
happens is made clearer by study upon other organisms.
When the Bacillus coli communis is grown in a nutrient liquid
containing glucose, both acid and gas are produced by the fer-
mentation of the glucose. The gas consists of carbon dioxide
and hydrogen, and the acids which are liberated, comprise lactic,
acetic, succinic, and formic acids. It has been found by W. J.
Penfold that strains af Bacillus coli coinmtvuis which do not pro-
duce gas from glucose, can be isolated by cultivation in the
presence of sodium mono-chloracetate. These races differ from
the typical parent-stock in their fermentative character on
glucose, and no individual member of the selected race appears
to retain the action of the typical parent micro-organism. The
presence or absence of the quality of liberating gas enables us to
recognise that, in the production of this character, every indi-
vidual of the selected race exhibits the altered function. If a
number of selected races are examined, it is, however, noted that
some show a complete absence of the gas-forming quality, while
others still retain more or less activity in this direction. The
careful studies of Egerton Gray have enabled us to learn some-
thing of the enzymatic changes which accompany this selection.
The estimation of the amount of the different products of the
decomposition of glucose supplies evidence to demonstrate that
lactic acid is formed in greater amount, and acetic acid, formic
acid, and alcohol in less amount, bv the selected strains than bv
30 president's address.
the typical micro-organism. Evidence has also been obtained
to show that the failure to ferment glucose is associated with tlie
absence (or diminution) of the reducing ferments in the cell. In
this way, the intermediate substance from which formic acid,
acetic acid, and alcohol is formed, is not decomposed. The
selection by the sodium mono-chloracetate can thus be explained
on a rational chemical basis, since those microbes endowed with
reducing qualities would liberate hydrochloric acid, and thus
occasion their extinction. Only microbes in which the reductase
is absent or diminished, would survive.
The study of these micro-organisms has made it evident that
the characters of these plants are determined by the conditions
of the environment. The question remains as to whether such
a conception can be applied to higher forms. Have soil and
climate any influence on the structure and functions of highei-
plants? There is much to suggest that the forms of plants are
dependent on the interaction with the environment. Before
this problem can be considered, it is necessary to obtain a better
knowledge of the plants which occur together in any district.
Owing to the small aboriginal population and their low state of
civilization, there has been little interference with the flora of
this Continent. Studies can be made upon areas in which plants
have grown together with little, if any, human selection. These
areas are, however, being settled by the white population.
Before these areas have been destroyed, it behoves our naturalists
to make their survey. In this vast continent, we can travel
from Cape Otway, in the south, to Cape York, in the north,
without losing sight of one or other species of Eucalyptus. It
is possible to go from Sydney, in the east, to the western shores
of Australia without missing anywhere the familiar Eucalyptus.
Is it not possible to determine what characters have enabled
these trees to extend over such a vast area"? May I suggest to
you that the members of our Linnean Society consider the sys-
tematic study of the distribution of our plants and the factors
w^hich determine their association. Such a labour would be a
worthy memorial of the work of this generation, and would
remain a testimony of our appreciation of the benefaction of our
revered founder. Sir William Macleav.
PRESIDENTS ADDRESS.
31
Mr. J. H. Campbell, Hon. Treasurer, presented the balance
sheets for the year 1917, duly signed by the Auditor, Mr. F. H.
Rayment, F.C.P.A., Incorporated Accountant, and he moved that
they be received and adopted, which was carried unanimously.
The income accounts are summarised in the following table: —
Heads of Expenditure,
General, Bacteriology.
Fellowships.
Total.
Administration
Maintenance
Publication
Research (Salaries) ...
Capital Accounts
£
586
39
303
654
£
5
63
too
100
£
221
1,525
£
591
102
524
1,925
754
Totals
1,582
568
1,746
3,896
Income (all sources) . . .
1,245 567
2,328
4,140
No nominations of other Candidates having been received, the
President declared the following elections for the Current Session
to be duly made: — President: Professor H. G. Chapman, M.I).,
B.8. — Members of Council (to fill six vacancies): Messrs. R. T.
Baker, F.L.S., W. W. Froggatt, F.L.S., Alex. G. Hamilton, C.
Hedley, F.L.S., Thomas Steel, F.L.S., and G. A. Waterhouse,
B.Sc, B.E.— Auditor: Mr. F. H. Rayment, F.C.P.A.
On the motion of Dr. J. B. Cleland, a very cordial vote of
thanks to the President, was carried by acclamation.
32
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II
36
ORDINARY MONTHLY MEETING.
March 27th, 1918.
Professor H. G. Chapman, M.D., B.S., President, in the Chair.
The Donations and Exchanges received since the previous
Monthly Meeting(28th November, 1917), amounting to 16 Vols.,
155 Parts or Nos., 27 Bulletins, 8 Reports, and 7 Pamphlets, re-
ceived from 75 Societies, Institutions, etc., and two private
donors, were laid upon the table
37
THE GEOLOGY OF THE LOWER MESOZOIC ROCKS
OF QUEENSLAND,
With special referp:nce to their Distribution and Fossil
Flora, and their Correlation with the Lower Mesozoic
Rocks of other parts of Australia.
By a. B. Walkom, D.Sc, Assistant Lecturer in Geology,
THE University of Queensland.
(Plates i.-ii., and six Text-figures).
Contents. pac^e
Introduction, AND .Scope OF Paper
Acknowledgments
Previous Literature
The Lower Mesozoic Rocks OF Queensland
(a) General
. (h) Geological Succession
(c) Lithological Characters ...
W Coals
f^e^ Extent and Distribution...
(i,) The Ipswich Series...
(ii.) The Bundamba Series
(iii.) The Walloon Series
(f) Artesian Water ...
(g) Folding Movements
The Flora of the Lower Mesozoic Rocks of Queensland
(a) The Flora of the Ipswich Series
(^/>; The Flora of the Walloon Series
(c ) Comparison of the Floras of the Ipswich and Walloon Series
(d) The Age of the Floras
Correlation of the Queensland Lower Mesozoic Rocks with
other Australian Lower Mesozoic Strata
(a) New South Wales
(h) Victoria
(c) South Australia ...
( d) Western Australia
(e ) Tasmania ...
88
41
42
45
45
47
48
50
52
52
58
54
58
60
(58
ti4
71
77
81
82
82
89
90
91
93
38 GEOLOGY OP LOWER MESOZOlC feOCKS OF QUEENSLAND,
CfJ Summary ... ... ... ... ... ... ... ... 9o
Geological History ... ... ... ... ... 96
Pal.*:ogeographical Considerations ... ... 102
SlTTVIMARY 11-2
Explanation or Pl.\tes ... ... 115
Introduction, and Scope of the Paper.
In Eastern xA.ustralia and Tasmania there is no large develop-
ment of rocks of marine origin, representing deposition during
the period between the close of the Palaeozoic era and the
beginning of the Cretaceous period. During this interval, with
a single exception, the sediments deposited in this region were
of origin other than marine; the organic remains included in
these rocks consist for the most part of plants, but, in addition,
fish* and insects! are abundant on some horizons, phyllopods
are represented by Ef<theria, and pelecypods by Ufiio and Unio-
itella. The exception mentioned above is the upper portion of
the Wianamatta Stage in New South Wales; the top of this
stage is formed by a thickness of 100 feet of a calcareous rock,
which contains a fauna of Ostracoda and Foraminifera. This
fanna has been described by Chapman, whose conclusion regard-
ing the species is as follows : '• These undoubtedly represent a
brackish or estuarine fauna, having a curious intermingling of
Hhsetic and Lower Jurassic types, with others more properly
referable to the Upper Pakeozoic of Europe. ■':|:
The term "Lower Mesozoic "' will be used throughout this
work for these rocks: the name "Trias- Jura ' has been generally
used, but it is not a suitable one, and we have not been, up to
the present, in a position to assign either a Triassic or Jurassic
age definitely to all of them. The name Lower Mesozoic may
be used generally to include the whole of these rocks, and to
refer to all the strata in Eastern Australia which were deposited
between the close of the Palaeozoic era and the be^innini; of the
Cretaceous period.
"' Woodward, Mem. Geol. Surv. N. S. Wales, Palseontology, Nos.4, 9, 10.
t Etheridge and Olhff, ihid., Palaeontology, No. 7; Tillyard, Queensland
Geol. 8uiv., Publication No.2o3.
J Records Geol. Surv. N. S. Wales, viii., p.33o, 1909.
BY A. B. WALKOM.
3&
The cofrelation of these beds is necessariU^ based on the fossil
plants, since they are the only remains which are of widespread
distribution. In the past, fossil plants have formed a ver}-
uncertain means for the accurate correlation of strata. This has
been due, particularly in the case of Australia, to imperfect
study of the material available. This statement is not made to
detract from the value of the work done by the earlier Australian
geologists; we owe much to their efforts, for they did an immense
amount of work under conditions much less favourable than
those under which we work at the present time. The vastly
improved conditions existing at the present day must always be
borne in mind, when the work of the older geologists is under
consideration; this point cannot be too strongly emphasised, for
there is very often a tendency to forget or overlook it.
The author has just completed an examination of the fossil
floraof the Queensland Lower Mesozoic strata,* and the present
paper is concerned chiefly with a discussion of the results of
that examination.
Briefly stated, the object of this contribution to Australian
Geology is to attempt to place the correlation of the Lower
Mesozoic rocks of Australia on a sound basis. The paper
aims at :
(a) Dealing in a fairly comprehensive manner with the general
characters and distribution of the Lower Mesozoic rocks of
Queensland.
(6) A critical discussion of the relations of the flora of the^se
rocks.
(c) A comparison and correlation of the Lower Mesozoic I'ocks
of Queensland with other Lower Mesozoic rocks of Australia,
paying particular attention to the relations of the fossil floras of
the various occurrences to one another.
(d) The determination of the position of the Australian Lower
Mesozoic rocks in the Geological Record. And
(e) A discussion of the palaeogeography of the Australasian
region during Lower Mesozoic time.
"' Queensland Geol. .Suiv., Publications No.s.252, 2.)7, '2,yj.
40 GEOLOGY OF LOWER MESOZOIC ROCKS OF QUEENSLAND,
It maybe of advantage hereto summarise the present position
of our knowledge of the Horas of Australian Lower Mesozoic
rocks.
Rocks of this age are developed in all the Australian States,
but only in Western Austi-alia are they of marine origin. The
state of our knowledge of the fossil tloras cannot be considered
satisfactory. In Western Australia, the number of plants in
these rocks is very small, but the fact, that they are associated
with strata containing marine fossils, is of great importance as
giving some indication regarding their exact position in the
Geological Record. Jn South Australia, a few fossil plants have
been described from the Lower Mesozoic rocks at Leigh's Creek,
and PhyJlopteris Feistmanteli has also been recorded from Ooroo-
wilanie Swamp, 100 miles north of Leigh's Creek; this latter
record may be from the Cretaceous rocks overlying the Rolling
Downs Series. In Tasmania, fossil plants are abundant, and a
large number have been described and figured from Lower
Mesozoic rocks. Unfortunately, some of the determinations
are open to doubt, and many of the figures and description are
too imperfect to be of value in a comparison of the Tasmanian
Mesozoic flora with other Australian Mesozoic floras In Vic-
toria, Lower Mesozoic rocks occur in three areas (South Gipps-
land. Cape Otway, and Wannon areas). A collection of fossil
plants from these strata has been described by Professor Seward,
and more recent additions have been determined by Mr. F.
Chapman. This flora has been determined as of Jurassic age,
and it is sufficiently well-known to enable reliable comparison to
be made with other floras. In New South A\ ales, fossil plants
are abundant in Lower Mesozoic rocks, but no comprehensive
examination of the whole flora has been undertaken. Lists of
the floras of the different Series, prepared by Mr. W. S. Dun,
were published in Carne's memoir on the Western Coalfield of
New South Males, in ly08. Ihese have been supplemented,
from time to time, by descriptions of additional specimens, by
Mr. Dun. From these lists, it is possible to obtain a fairly good
idea of the flora of the Lower Mesozoic rocks in New^ South
Whales. In Queensland, the author has just completed an ex-
BY A. B WALKOM. 41
amination and revision of tlie floras of the Ipswich and Walloon
Series, the results of which have been published by the Queens-
land Geological Survey.
The correlation of the Lower Mesozoic rocks of Australia
(based mainly on their fossil floras) and their exact position in
the Geological Record have been the subject of much discussion,
and different views regarding their correlation have been put
forward from time to time. In a summary of the literature
dealing with the Queensland Lower Mesozoic rocks,* I liave
already briefly summarised the majority of these views, and
there is no need to discuss them in further detail here. It is
sufficient to state that no finality has been reached on this point,
and, in many cases, conclusions have been based on insufficient
evidence.
As a result of my examination of the Queensland fossil flora,
together with a consideration of the evidence of the fossil faunas,
and, in addition, general considerations of the stiata containing
the fossils, certain conclusions regarding the correlation of the
various series have been arrived at (see p. 95) which differ from
any previously put forward.
Acknowledgments.
It is almost impossible to express the extent of my indebted-
ness to Mr. B. Dunstan, Chief Government Geologist of Queens-
land, in connection with the whole of my geological and palteon-
tological work in Queensland. He has placed at my disposal
every facility for examining the large collection of Mesozoic
plants in the possession of the Geological Survey; in addition,
he has made available to me a very considerable amount of
unpublished information, and permitted me to use such for the
purpose of making this paper as complete as possible. In the
section of this paper dealing with the extent and distribution
of the Lower Mesozoic rocks, much field-information is published
for the first time, and this is based almost entirely on informa-
tion supplied by Mr. Dunstan. I would like, therefore, to make
special acknowledgment of my indebtedness to Mr. Dunstan,
* Queensland Geol. Surv., Publication No. 252, p. 6.
42 GEOLOGY OF LOWEK MESOZOIC kOCKS OP QUEENSLAND,
and to place on record my appreciation of the generous way in
which he has lent me all possible assistance.
To other officers of the Geological Survey, I am indebted for
their willingness to assist me with their knowledge of these
strata in the field, especially Mr. W. E. Cameron, who has made
such a detailed study of the Ipswich Series in the Ipswich Coal-
field, and who has given me access to the information which he
has obtained for the preparation of his third report on this
field.
For assistance in connection with the geology of the Lower
Mesozoic rocks in Western Australia, South Australia, and Tas-
mania, I have to thank Messrs. A. Gibb Maitland, L. K. Ward,
and W. H. Twelvetrees, respectively.
To Professor David, I owe my introduction to the broader
problems of Australian Geology, and the resulting desire to assist
in the solution of some of the problems of Australian strati-
graphy. I have been inspired to carry out the palseo-botanical
portion of my work as thoroughly as possible, by the kindly
encouragement of Professor Seward.
I am indebted to the authorities of the University of Sydney
for permission to publish this paper.
Previous Literature.
Most of the papers dealing dii-ectly with the subject of the
Lower Mesozoic rocks of Queensland, or their flora, have already
been briefly summarised in an earlier papei', but there are a few
additional papers to which reference may be made here.
^Ir. C Hedley, in his Presidential Address to Section D of
the Australasian Association for tl)e Advancement of Science
in 1909, published two maps of the Queensland region in Meso
zoic time, one reproduced after Keumayr, showing the distribu-
tion of land and water in Triassic time; and the other original,
showing the same at the close of the Mesozoic Kra.
Dr. H. I. Jensen, in a paper entitled '-The Building of Eastern
Australia,"* has many references to the Lower Mesozoic rocks.
The paper partakes rather of the nature of a summary of views
"" Proc. Roy. Soc. Queensland, xxiii., Ft. 2, 1912, p.U9.
GV A. M. WALKOM. 43
the author stating in his introduction that the object of the
paper "is not to offer the reader any really new material, but
rather to present the knowledge we alieady possess in a concise
form "' JS^evertheless, many interesting points are raised,
one or two of which bear more particularly on the problem of
the Lower Mesozoic, and these will be referred to later. A
series of maps accompanies the paper, showing suggested dis-
tribution of land and sea in the Australian region during the
various geological periods.
Mr. E. C. Saint-Smith has carried out extensive observations
on the Lower Mesozoic rocks in the Koma District, and presented
a summary of his results* to the Second Interstate Conference
on Artesian Water. These observations cover a very large area
of Lower Mesozoic rocks, and fossil plants were found at various
localities. The present author had the pleasure of accompanying
Mr. Saint-Smith on a reconnaisance-trip over part of the area
between Yeulba, Goongarry (Hornet Bank Station), and Roma,
and can confirm Mr. Saint-Smith's remarks regarding the strata
of that area. 'J'he greater part of the country traversed consists
of outcrops of sandstones, with, here and tliere, shales and coal-
seams, belonging to the equivalents of the Walloon Series.
Professor David, in the Federal Handlx^ok for the Meeting of
the British Association for the Advancement of Science in Aus-
tralia in 1914, refers briefly to the Lower Mesozoic rocks of
Queensland, which he classes as Jurassic.
Mr. R. J. Tillyard has described a number of fossil insects,
from Lower Mesozoic rocks in Queensland and New South
Wales. t The majority of the insects were obtained from
Denmark Hill, Ipswich, in strata belonging to the Ipswich
Series, and others from St. Peter's, near Sydney, N.S.W., from
the Wianamatta Stage of the Hawkesbury Series. Twenty-two
species were described from the Ipswich Series, and six species
from the Wianamatta Beds.
The insects appear to be of considerable interest from the
* Rept. Second Interstate Conference on Artesian Water, Brisbane, 1914,
p. 19.
i Queensland (ietjl. Sur\ ., Publication Xu.2r)3.
44 GEOLO(iY OF LUWRli MBSOZOIC ROCKS OF gUEKNSLAXD,
point of view of the phylogeny of the Iiisecta, but the results,
up to the present, do not indicate that they will be of material
value in the determination of horizons, or in fixing the position
of the strata in the Geological Record.
In Appendix B to the " School Geography of Queensland," by
G. Harrap, published in 1916, Mr. Dunstan gives a new classi-
fication of the geological formations of Queensland In this
classification, the Lower Mesozoic rocks are divided into (?)Tri-
assic and Jurassic, the Ipswich Series and Bundamba Series
being classed as (?)Triassic, and the Walloon Series as Jurassic.
No definite evidence is given for the separation. Included also
in the Jurassic are the trachytes of the Glasshouse Mts., which
all recent work goes to show are of Cainozoic age.
Professor Schuchert, in a paper recently issued, entitled "The
Problem of Continental Fracturing and Diastrophism in Ocean-
ica,"* gives a series of palseogeographic maps of Oceania; these
include one showing the distribution of land and water in the
Triassic period.
Mr. E. C. Andrews, in a recent paper entitled " Notes on the
Structural Relations of Australasia, New Guinea, and New Zea-
land,"! makes many statements which invite criticism, some of
them coming within the scope of this paper. He considers the
growth of Australia, New Guinea, New Caledonia, and New
Zealand as independent units. The question suggests itself —
Are not these portions of the one continental mass, which have
.become separated as a result of folding-movements'? In discuss-
ing the 'J'rias-Jura, he infers two basins of deposition in New
South Wales and Queensland, viz.: the Hawkesbury basin, and
a northern basin, separated by high land-barriers, and with
sedimentation taking place simultaneously in the two basins.
There are many papers dealing with the volcanic rocks of
south-eastern Queensland, in which passing reference is made to
the Lower Mesozoic rocks occurring in the same area as the
volcanic rocks. It is unnecessary to mention these in detail
* Amei\ Journ. Sci., xlii., 1916, p. 91.
t Journ. Geol., xxiv. , 1916, p. 751.
BY A. B. WALKOM. 45
here: such papers include the works of Jensen, Wearne and
Woohiough, and Richards, in addition to the publications of the
Queensland Geological Survey.
The Lower Mesozoic Rocks of Queensland.
(a) General. — Until quite recently, the Lower Mesozoic rocks
of Queensland have been officially designated "Trias-Jura."
This name was first used in 1892, it being suggested that the
Burrum and Ipswich Formations represented a period of time
extending from the base of the Trias to the top of the Oolite.*
Prior to that date, in 1888, regarding the Ipswich and Burrum
Formations, Jack saysf "The Burrum Coalfield is plainly on a
higher horizon than the *Bowen River field [Permo-Carbonifer-
ous]. It contains a fossil flora in which many plants are common
to the Mesozoic Ipswich Formation, and also, it is said, Glossop-
terin with a very meagre fauna, most of it peculiar to the coal-
field." "Probably to call it Triassic would not be very far
from the mark, in al least a homotaxial sense." In the same
paper, speaking of the Ipswich coalfield, he says " The coalfield
contains an abundant fossil flora of a strongly Jurassic facies,
and is probably the equivalent of the Clarence River beds of
New South Wales."
It appears, then, that the recording of Glossopteris in the
Burrum Formation was originally responsible for its being
regarded as older than the Ipswich Formation. There is, how-
ever, no authentic record of the presence of Glossopteris in the
Burrum Series.
When the two formations were united, in 1892, as the Trias-
Jura, the Burrum Formation was regarded as Lower, and the
Ipswich Formation as Upper Trias-Jura.+ Ten species of plants
were described by Etheridge from the Burrum Formation, and
thirty-one from the Ipswich. The two formations were not
known in contact in the field, so there was no stratigraphic
evidence as to their relation to one another, and this had to
* Geology and Palaeontology of Queensland, 1892, p. 312.
t Report Aust. Assoc. Adv. Sci., i., 1889, p. 196.
ij: Geology and Palieontulogy of Queensland, 1S92, p. 312.
46 GEOLOGY OF LOWER MESOZOIC ROCKS OF QUEENSLAND,
be determined from the contained fossil plants. The fossils,
which were available from the Burrum Formation at the time,
were, unfortunately, fragmentary, and were not a representative
collection. Further collections have been obtained from time to
time since, and a cursory examination of the material now avail-
able in the Geological Survey collections shows distinct differ-
ences from the floras of the Ipswich and Walloon Series. It is
intended that an examination of the Burrum flora will be under-
taken after the completion of the present paper.
In 1907, Mr. Cameron, in discussing the age of the Ipswich
Formation, says* " The evidence for considering the Burrum
Beds as belonging to an earlier period of the Trias- Jura is not
conclusive. The two formations have long been considered as
identical in age by the Geological Survey, and the recent obser-
vations of Mr. Jensen lend confirmation to that conclusion.''
Our present knowledge shows that a large part of what was,
at that time (1907), regarded as part of the Burrum Formation,
is actually a continuation of the upper series of what was con-
sidered then as the Ipswich Formation. Dr. Jensen had
recorded the fact that these were continuous in the field in the
neighbourhood of Point Arkwright.f
The question of the age and extent of the Burrum Formation
has, however, been established beyond doubt by the obser\ations
in the field of Mr. Dunstan. As a result of these observations,
it is now certain that the Burrum Series, in the Maryborough-
Howard district, overlies, with apparent stratigraphic conform-
ity, | rocks of marine origin, whose contained fossils indicate a
Cretaceous age, probably equivalent to the Rolling Downs Forma-
tion of Western Queensland. JV'Ir. Dunstan has also shown that
the strata to the south and south-west of Maryborough, origin-
ally mapped as part of the Burrum Formation, dip towards the
north-east beneath the marine Cretaceous rocks, and are equiva-
lent to the Walloon Series. For these equivalents of the
* Queensland Geol. Surv., Publication No. 204, pp.12, 13.
t Proc. Linn. Soc. N. 8. Wales, 1906, xxxi., pp. 74-75.
t Ann. Rept. Dept. Mines, Queensland, 1911 (1912), p. 195; Queensland
Govt. Mining Jouin., xiii.(1912), p. 641,
BY A. B. WALKOM. 47
Walloon Series in this area, Mr. Dunstan has proposed the
name " Tiaro Series."*
Of the two formations which were united to form the so-called
Trias-Jura System, then, one has been removed into the Creta-
ceous System, and the remaining one (the Ipswich Formation;
is that which was originally regarded as the probable equivalent
of the Jurassic System.
The Ipswich Formation was divided into three Series by Mr.
Cameron, viz.: Ipswich Series, Bundamba Series, and Walloon
Series. The use of the terms Ipswich Formation and Ipswich
Series (part of the Ipswich Formation) has probably led to a
certain amount of confusion, but the removal of the Burrum
Formation to the Cretaceous System renders the use of the name
Ipswich Formation superfluous, and it would be an advantage
to abandon it.
(6) Geological Succession. — The Lower Mesozoic rocks of
Queensland consist of three Series, viz.: —
3. Walloon Series (Upper).
2. Bundamba Series (Middle).
1. Ipswich Series (Lower j.
The complete succession is found only in South-CHstern Queens-
land, where a considerable amount of detailed geological work
has been carried out, chiefly because both Ipswich and Walloon
Series ccmtain workable coal-seams.
Ihe Ipswich district has been examined in detail bv Mr.
Cameron, who has published two reports on the area,t and is
preparing a third, incorporating the results of recent develop-
ments in the district.
South-east Moreton has been the subject of investigation by
Mr. E. O. Marks, late of the Queensland Geological Survey,
whose results are embodied in a publication entitled "The Coal-
Measures of South-east Moreton. '| The examination of the
continuation of the Coal-Measures, from the area examined by
* See Rept. Second Interstate Conference on Artesian Water, Brisbane,
IftU, p.7.
t Queensland Geol. Surv., Publications 147, 204.
X Queensland (ieol. Suiv.. Puljlication 22o.
48 GEOLOGY OF LOWER MESOZOIC ROCKS OF QUEENSLAND,
Mr. Marks towards the New South Wales border, has been com-
menced by Mr. J. H. Reid, of the Geological Survey.
Other areas of Lower Mesozoic rocks in Queensland have been
studied in detail by other officers of the Geological Survey, the
more important ones being : the Stan well Coal-Measures, ex-
amined by Mr. Dunstan;* the Laura Coalfield, west of Cooktown,
by Mr. Ball;! the Lower Mesozoic rocks in the Roma district,
by Mr. Saint-Smith. Unfortunately, the full results of Mr.
Saint-Smith's work have not been published, but a summary was
communicated to the Second Interstate Conference on Artesian
Water. I For local details regarding these areas, reference must
be made to these publications.
(c) Litholoyical Chavdcters. — The lithological characters of the
Lower Mesozoic rocks in Queensland do not call for very special
remark. For the greater part, they consist of sandstones and
shales in varying proportions, with which are associated occa-
sional conglomerates, grits, and also coal-seams.
The Ipswich Series consists, for the most part, of light-coloured
shales and sandstones, with associated conglomerates, and four-
teen or fifteen workable coal-seams. There are coarse conglom-
erates developed near the base of the series, indicating vigorous
erosion in the early portion of the period.
The Brisbane Tuff, which is near the base of the Ipswich
Series in the Brisbane area, is a rock probably of volcanic origin
(though there is no indication of its source), resulting from the
deposition of volcanic ash of acid nature over a long, narrow area
in the vicinity of Brisbane. The general direction of the out-
crop of this belt of tuff is N.30°W. As far as known, there are
no volcanic rocks of undoubted Lower Mesozoic age in South
eastern Queensland, with the exception of a series of andesites
south of Maryborough, which are apparently interbedded in the
Tiaro Series. Marks, Andrews, and Wearne and Woolnough
have, at times, advocated a Lower Mesozoic (Trias-Jura) age for
* Queensland Geol. Surv., Publication 131.
t Queensland Geol. Surv., Publication 222, p. 5.
X Report Second Interstate Conference on Artesian Water, Brisbane,
1914, p.l9.
BY A. B. WALKOM.
49
some of the South-east Queensland volcanic rocks^ hut l)i-.
Richards has recently examined all their evidence* and shown
that, in each case, there is no doubt that the volcanic rocks are
of Cainozoic age. Since then, Mr. Dunstan lias placed the
volcanic rocks of the Glasshouse Mountains in the (?)Walloon
Series;! the only field-evidence regarding the age of these rocks
is that tliey are intrusive through a series of sandstones, which
Mr. Dunstan believed, at the time, to belong to the Ipswicli
Series. On the evidence of the relationship between these vol-
canic rocks and other volcanic rocks of South-eastein Queensland,
their age certainly seems to be Cainozoic. Some of the beds of
the Ipswich Series at Denmark Hill, Ipswicli, are regarded by
Mr. Dunstan as tufi'aceous in character. |
The Bundamba Series, which succeeds the Ipswich Series,
consists, for the most part, of massive sandstones. With the
exception of one coal-seam about 1 foot, 6 inches thick, known
as the West Moreton seam, they have proved unproductive of
coal; they are also practically barren of fossils, the only record
being of silicified wood. The basal portion of the Bundamba
Series lies about seventy feet above the top seam (Aberdare
seam) in the Ipswich Series, and consists of coarse grits and
conglomerates.
The Walloon Series, lithologically, is of somewhat similar
nature to the Ipswich Series. In consists mainly of compara-
tively soft sandstones, with which are interbedded fine-grained
shales and coal-seams. Conglomerates are developed, but are of
limited occurrence. The soft sandstones and shales weather
away rapidly, and, consequently, in many places, outcrops are few
and inconspicuous.
In the Stewart's Creek (Stanwell) district, the Series includes
a hard, white, siliceous tuff in which abundant fossils are pre-
served.
A remarkable feature in the Walloon Series is the verv wide-
* For summary of evidence, see Proc. Roy. Soc. Queensland, xxvii. (2)
1916, p. 126.
t Harrap's School Ueographj' of Qvieensland, Appendix B, p 16
X Queensland Geol. Suiv., Publication No.2o3, p.5
I ! B
R Y
If' #
50 GEOLOGY OF LOWER MESOZOIC ROCKS OF QUEENSLAND,
spread distribution of a fine-grained, ferruginous sandstone.
This rock is very characteristic, and is developed on one or
perhaps more horizons. It has been observed from the following
localities : Beaudesert, Kalbar(late Engelsburg), near Warwick,
near Toowoomba, three miles north of Texas, a number of
localities in the Wallumbilla-Roma district, and Stewart's
Creek, Rockhampton. All these occurrences are in the Walloon
Series or its equivalents. This rock, in nearly every case,
contains fossil plants, the genus Otozamites being particularly
characteristic. It is also to be noted that, in the Jurassic
rocks of Western Australia, Otozamites occurs in a fine-grained,
ferruginous sandstone.
The widespread distribution of this rock, apparently on a few
horizons, leads us to believe that it gives indication of some
special conditions of deposition. It would, however, be useless
to attempt to account adequately for it in the present state of ,
our knowledge of the conditions under which these beds were
laid down; most of the areas from which it has been obtained
have not yet been studied in detail geologically.
{d) Coals. — The development of a number of coal-seams of
workable quality and thickness is an important feature of the
Lower Mesozoic rocks of Queensland. Coal is obtained from
both the Ipswich and Walloon Series, and the character of the
coal from the two Series is generally distinct. This distinction
has been expressed both by Mr. Cameron and Mr. Marks. The
former, speaking of the Walloon coals, says* "the coals hitherto
found show characteristic conchoidal fracture in the hand-speci-
men, burn readily with a long, luminous flame, and give off a
much larger proportion of volatile hydrocarbons when heated in
a closed vessel than do the brittle, bituminous coals of the
Ipswich Beds."
On the same subject, Marks saysf "The Walloon Beds are
characterised by coals which, like those of the Darling Downs,
are of the nature of a cannel coal — highly gaseous, hard, and
* Queensland Geol, Sufv., Publication No.204, p. 16.
t Queensland Geol. Surv., Publication No. 225, p. 9.
BY A. B. WALKOM.
51
breaking with a concboidal fracture, in contrast to the more
brittle bituminous coal of the Jpswich Beds."
The Ipswich coals are steam-coals, and are suitable for heatino-
and coke-making; the Walloon coals are essentiall}^ gas-coals. A
table of typical analyses of the coals from various areas has been
published by Mr. Dunstan,* from which we may quote the typical
Ipswich and Walloon coals for comparison.
Ipswich Coal.
Walloon Coal.
Mean %.
Range.
Mean %
Range.
Moisture
Volatile hj'drocarbons
Fixed carbon
Ash
1-5
27 0
58-5
140
2 0-0 -7
33-21
72-50
21-4
P.-O
39-0
44-0
11 0
8-4
40-23
48-29
25-6
The most notable point of contrast between the two is in the
relation of volatile hydrocarbons to fixed carbon; in the Ipswich
coals, the latter is very much in excess of the former, while, in
the Walloon coals, the two are of about the same value. In
some cases, Walloon coals show a much higher percentage of
fixed carbon than of volatile hydrocarbons, and the analysis is
then indistinguishable from that of Ipswich coals; in these cases,
however, the field-relations of the strata usually supply the ex-
planation for this irregularity, by the presence of intrusive rocks
not far away, and resultant alteration of the coal.
In normal cases, then, there is a more or less marked distinc-
tion between the coals of the Ipswich and Walloon Series in
Queensland, and this distinction is of some practical value in
helping to distinguish between the two Series. The Walloon
coals are distinctive, and can frequently be recogni.'^ed M'ith a
reasonable amount of certainty as belonging to that Series. The
Ipswich coals, however, though they are distinct from the Wall-
oon coals, are very similar to the coals of the Burrum Series, of
Cretaceous age, and their age cannot be identified by the nature
of the coal.
Queensland Geol Surv. , l^ublication No.239, p. 23.
52 GEOLOGY OF LOWER MESOZOIC ROCKS OF QUEENSLAND,
(e) Extent and Distrihiition.—ii.) 7'he Ipswich Snnes. — The
Ipswich Series is of comparatively limited extent, and has a
thickness, in the type-district, of about 2,000 to 2,500 feet, as
estimated by Mr. W. E. Cameron. Its best development is in
the Ipswich district, where the strata have been studied in detail
by Mr. Cameron.*
North-west of the town of Ipswich, the north-western end of
the Ipswich Series is hidden by Cainozoic rocks. ^Mapping in
this portion of the area has not been carried out in detail, but
apparently the Ipswich Series cuts out between the Bundamba
Series and Brisbane Schists (as shown in Plate ii.), and it is not
known to outcrop further in this direction. It is, of course, pos-
sible that this series extends some distance north under the
Walloon Series, but one might expect, in this case, to find some
indication of its presence by outcrops between the outcrops of
the Walloon Series and the older rocks to the east.
From Ipswich, the Ipswich Series extends in a general easterly
direction to Oxley, where it disappears beneath overlying Caino-
zoic rocks, as shown on the most recent maps prepared b\' iMr.
Cameron. It reappears along a line running approximately
N.30°W.-S.30''E through Brisbane, and is succeeded to the east
by a line of schists of Pala?ozoic age. This belt of Palaeozoic
rock is not very wide here, and, on the eastern side of it, the
Ipswich Series reappears. Between Mt. Cotton and Mt. Petrie,
the two belts of Ipswich Series are in direct connection, as also
are they between Mt. Petrie and White's Hill
Still going to the east, the Ipswich Series again disappears
under the overlying Bundamba sandstone along a line running
in a N.30°W. direction through Hemmant, and reappears on the
other side of a syncline at various points on the coast south of
the Brisbane River. The axis of this syncline is in a direction
N.30°W.-S.30°E.
The whole of the Ipswich Series so far described skirts the
southern extremity of an extensive occurrence of the schists
known as the Brisbane Schists, whose age is uncertain, and can
only be stated definitely as Pre-Mesozoic.
* Queensland Geol. Surv., Publications Nos.147, 204,
BY A. B. WALKOM. 53
From the Brisbane River, extending north as far as Mary-
borough, there is a narrow, coastal belt of Lower Mesozoic rocks.
These are, here and there, intruded or overlain by volcanic rocks
of Cainozoic age, with which we are not concerned here. The
Mesozoic rocks of this coastal belt are gently folded into anti-
clines and synclines, the general direction of dip being north-
easterly or south-westerly.
This coastal belt averages about 15 to 20 miles in width. At
its southern end, it consists of rocks of the Ipswich Series, repre-
senting a continuation of the syncline just mentioned south of
the Brisbane Biver. The Bundamba sandstone in this synclinal
area does not extend much north of the Brisbane Fviver, and
appears to be entirely surrounded on its northern end by the
Ipswich beries. The north-eastern side of this syncline, pro-
duced northwards, forms the south-western arm of an anticline
whose axis passes approximately through the region of the
Glasshouse Mountains, in a direction N.SO^W., and whose north-
eastern arm dips away towards the north east under the sand-
stones of the 'J'oorbul Point-Landsborough district, which pro-
bably represent the Bundamba Series.
This is the last that is seen of the outcn^p of the Ipswich
.Series, and their extent under the Bundamba Series to the east
and north east cannot be determined.
The distribution of this Series is indicated generally in Plate ii.
(ii.) The Bundamba Series. — The Bundamba Series comprises
a development of massive sandstones in which there are practi-
cally no fossils, the only ones so far recorded being examples of
fossil wood. Mr. Cameron estimates the thickness of the Series
at between 3,000 and 5,000 feet.
This Series overlies the Ipswich Series conformably. It ex-
tends, roughly, from the Ipswich District eastwards to a little
beyond the railway-line between South Brisbane and Kingston,
and in a general south-easterly direction past Canungra to the
New fcouth Wales border. It is impossible, in this area, to
define accurately the limits of the formation, but its existence is
beyond doubt. Mr. Dunstan has recently observed it in iSew
South Wales, not far from Mt. \N arnini{.
54 GEOLOGY OF LOWER MESOZOlC ROCKS OF QUEENSLAND,
The Series also occupies a syncline, with an axis in a direction
approximately N.30°W.-S.30°E., between Hemmant and Wyn-
num, extending southwards to about the latitude of Redland
Bay, and northwards only just beyond the Brisbane River.
Further north, there is a parallel belt of the Bundaniba Sand-
stone extending from Toorbul Point in a direction approximately
N.SO'VV. through the Landsborough district. The presence of
the Bundamba sandstones has been indicated at Toorbul Point
by Mr. Cameron.* At this point, a bore (the Bribie View Bore)
in Portion 28, Parish of Toorbul, passed through 508 feet of
massive sandstones, with one or two seams of hard, black, sandy
shales and, near the bottom, pebbly conglomerates; no coal-seams
were observed in these sandstones. Mr. Cameron correlated these
sandstones with the Bundamba grits and conglomerates occurring
at Logan Village. The sandstones of the Landsborough district
have been correlated with the Bundamba Series by Mr. Dunstan. f
It can thus be recognised, that there is a belt of the Bundamba
Series running through this area, and its position can be defined
in a general way, but the exact determination of its boundaries
would involve a careful geological survey of the whole area, and
it is not certain even then that the boundaries could be accu-
rately defined.
This belt disappears towards the north-east under the southern
extension of the Tiaro Series, which is equivalent to the Walloon
Series. The distribution of the Series is indicated in Plate ii.
The Helidon sandstones may be the equivalents of the Bun-
damba sandstones, as suggested by Mr. Dunstan,! but we do not
know sufficient detail of the geology of the Helidon district, at
present, to be sure of their position.
(iii.) The Walloon Series. — The rocks belonging to this Series
cover a vastly wider area than either the Ipswich or Bundamba
Series. Their distribution is indicated in Plates i. and ii. In
South-eastern Queensland, they outcrop over most of the country
between a line drawn from just west of Esk to Toowoomba,
thence along the eastern foot of the Main Bange to the New
* Ann. Rept. Dept. Mines, Queensland, 1908, p. 172.
t Queensland Geol. Surv., Publication No 252, p.4.
BY A. B. WALKOxM. 55
South Wales border, and a line drawn roughly from Ipswich to
('anungra, and thence to the New South Wales border. In a
good deal of this area, they are overlain by volcanic rocks of
Cainozoic age, but there is no doubt that they are practically
continuous through the whole area. In addition, there is a belt,
live to fifteen miles wide, from Esk up the valley of the Brisbane
River nearly to Cooyar Creek, but this belt is not a continuous
outcrop; there is also an extension from Esk in a north -easterly
direction past Mt. Brisbane.
Rocks belonging to the Walloon Series also outcrop verv ex-
tensively to the west of the Main Divide. There is probably a
continuous belt from the New South Wales border near Kil-
larney, running through Warwick to Dalby and Chinchilla, then
swinging round to an east and west direction parallel to the
railway-line past Roma. The exact extent of the belt past this
point is not absolutely proven, but there seems little doubt that
it swings round in a general north-north-westerly direction, and
extends right to the Gulf of Carpentaria towards Cape York.
Mr. Dunstan* has placed, in the W^alloon Series, sandstones and
shales in this belt at the following localities: the Upper Maranoa
River, the Upper Dawson River, Jericho, the Upper Flinders
River, Croydon, and probably the heads of the rivers flowing west-
erly across the Cape York peninsula to the Gulf of Carpentaria.
This belt dips to the west and south-west, and underlies, with
apparent conformity, the marine Cretaceous rocks of the Rollins*-
Downs Series. In the east, the Walloon Series is represented
by a number of occurrences scattered over a very wide area. To
the south and south-west of Maryborough is the Tiaro Series
(equivalent to the Walloon Series); other occurrences include
those at Callide Creek, Westwood-Wycarbah-Stanwell district
Waterpark, Brovinia, a small area about 10-15 miles west of
Mundubbera, a small area west of the Burnett River in latitude
25° 45'S., a small area west of Barambah Creek in latitude 26°S
at Mondure on Barambah Creek, and the Laura Coalfield, Cook-
town district. The exact positions of these areas are shown on
* Harrap's Geography of Queensland, 1916, p. 166.
56 GEOLOGY OF LOWER MESOZOIC ROCKS OF QUEENSLAND,
Plates 3, 5, and 9 of the Queensland Mineral Index. On Plate
1 1 of the same work, a small patch is shown as belonging to the
Walloon Series, just west of Anakie. Mr. Dunstan, who in-
vestigated the Anakie district some years ago, kindly informed
me tliat tliere is no representative of the AYalloon Series in that
district, and that the colouring of this area as Walloon was due
to an error in lithographing. Another area, to M'hich attention
should be called, is that of the Styx River, marked on Plate 3
of the Mineral Index as belonging to the Ipswicli Series. Mr.
Dunstan now believes this to be of Cretaceous age, and has
recently obtained, from the Styx River Coal -Measures, a frag-
ment which appears to be a dicotyledonous leaf. This is an
interesting find, and the greatest age, which can reasonably be
assigned to these Measures in view of it, is Cretaceous.
In the case of most of the isolated occurrences of the Walloon
Series just mentioned, there is sufficient evidence to render it
almost certain that the determination of' the age is correct. The
evidence is here summarised : —
In the Westwood-Wycarbah-Stanwell district, the fossil Hora
is distinctly of the Walloon type {infra^ Table vii.). At Water-
park, the coal is of Walloon type, but there appears to be no
record of fossils. The areas in the vicinities of Brovinia, Mun-
dubbera, Burnett River, Barambah Creek, and Mondure are all
residuals, occupying rather higher ground, and Mr. Dunstan is
of opinion that they undoubtedly represent outliers of the exten-
sive outcrop a little to the south-west. In the Laura coalfield,
the strata consist of arenaceous beds, with occasional thin beds
of shale and coal-seams. Mr. Ball records the finding of the
following fossils : Fhyllotheca(i), Tte'itioj)teris, Alethopteris, Bra-
chyphyllunn^ and Taxitesi^.). The presence of BrachyphyUiim a,i[\(\
TaxUes(l) is sufficient to indicate a Walloon age. In the remain-
ing area, Callide Creek, there is no definite indication of age;
the fossils, Thinnfeldia odonlopteroides [ = T. FeislmaiUeli], and
Tceniojjteriti sp., have been recorded, but they are not sufficient
to determine the age. In view, however, of the wide distribu-
tion of strata of V\ alloon age, the Callide Creek beds are re-
garded as probably of similar age.
BY A. B. WALKOM. 5<
It is thus seen that the Walloon Series extends uninterriipt
ediy over a great part of Eastern Queensland. Though occurring
iu comparatively small, isolated areas at the present time, I
believe that these occurrences represent the remnants of a de-
position which probably covered the greater pait of Queensland.
This will be discussed later.
Further, that the western belt, extending past Toowoomba
and Warwick, probably extends a great deal further west than
is shown on the map, is indicated by the known occurrences of
Lower Mesozoic rocks to the south-west of Dalby, and west of
Warwick.
Towards the south, this belt divides into two; the eastern
portion extends past Killarney into New South Wales, and is
continuous with the western extent of the Clarence Series; and
the western portion continues along the western margin of New
England, and dips away to the west under the marine Cretaceous.
In New South Wales, this latter belt is known as the Artesian
Series.
It must be noted that the Walloon Series, in the Toowoomba-
Warwick District and in South-eastern Queensland, have pro-
bably been separated by heavy faulting along the Main Range,
but were formerly continuous; and it is quite natural, therefore,
that this Series should, in these two areas, be continuous with
parts of the Clarence Series of New South Wales.
That the Walloon Series continues for great distances west-
ward under the Cretaceous rocks, is proved by the Artesian
bores of Queensland and New South Wales. It is fairly well
established now that the rocks from which the Artesian water
(or rather that part of it which is of meteoric origin) is obtained,
are part of the so-called Trias-J ura System, and not of the Cre-
taceous System. This fact is of great value in determining the
extent to which the Lower Mesozoic rocks continue under the
Cretaceous.
Small Hows of water have undoubtedly been obtained from
the Cretaceous rocks, but the rocks from which the large flows
of water have been obtained are of Lower Mesozoic age. Ex-
amination of the bore-records, then, will show the extent of the
5<S GEOLOGY OF LOWER MESOZOlC ROCKS OF QUEENSLAND,
Lower Mesozoic rocks, both in Western Queensland and in New
South Wales. At some places, there is a very great thickness
of Cretaceous and possibly Cainozoic strata overlying the Lower
Mesozoic rocks; e.g., in South Australia, the bore at Goyder's
Lagoon struck water bearing strata at 4, 700 feet, and the Patcha-
warra bore was abandoned at a depth of 5,458 feet, being still
in the Cretaceous rocks.
Palaeontological proof of the age of the water-bearing strata
in the bores is not often forthcoming, but, in a number of cases
in New South Wales, records have been made, e.g., (a) in the Bul-
yeroi bore, 60 miles W. by S. of Moree, Lower Cretaceous rocks
with marine fossils were passed through down to 520 feet, and
then shales, sandstones, and coal-seams with fossil plants: (b) in
the Wallon bore, 20 miles N. by W. of Moree, Lower Cretaceous
rocks with marine fossils were encountered dow^n to 1,500 feet;
at 1,630 feet, fragments of Tceuiopteris spatnlata [7'. Baiiiti'eei]
were obtained, and water was struck at 2,330 feet; (c) in the
Coonamble bore, both T(vniopteris sjmtulata [7'. Daiiitreei\ and
Thlniifeldia odoiitopteroides were obtained. These are not all of
the recorded occurrences.
The Walloon Series (or its equivalents) is considerably thicker
than the Ipswich or Bundamba Series. In AVestern Queens-
land (Roma District), the map and sections prepared by Messrs.
Saint-Smith and Thom show a width of outcrop of about 60
miles, and dips of the order of 3 or 4 degrees; the dips are small,
but, if the average dip be only 2 degrees, the thickness repre-
sented is about 11,000 feet. En the Maryborough District, the
thickness of the Tiaro Series has been estimated at 12,000 feet
by Messrs. Blake and Bryan.
if) Artesian IFa^er. —A general consideration of the Lower
Mesozoic rocks of Queensland would not be complete without
some reference to the question of artesian water. This question
has been the subject of considerable controversy between the
exponents of the two theories as to the origin of the water,
known respectively as the '^ Meteoric" Theory, and the "Plu-
tonic" Theory. Suffice it to state, that there seems now to be
no reasonable doubt that a large portion of the water is of
BY A. B. WALKOM. 5^
meteoric origin, and that the belt of Lower Mesozoic rocks ex-
tending, probably without interruption, from near Dubbo in New
South Wales, to the Cape York Peninsula, forms the intake-beds
of the Great Australian Artesian Basin. Probably also a small
percentage of the water is of plutonic origin. It is also now
generally agreed among Australian geologists that, in the Great
Australian Artesian Basin, the rocks, from which all the large
flows of water are obtained, are of Lower Mesozoic age, under-
lying the Marine Cretaceous (Rolling Downs) Series.
An examination of the bore-records shows conclusively that
the sandstones of the Walloon Series in Queensland, and the
Artesian Series in New South Wales, extend continuously in a
westerly and south-westerly direction into the north-eastern por-
tion of South Australia. The depths at which the sandstone is
found vary, but, in genei al, the deepest occurrences are in the
north-eastern corner of South Australia, not far from the Queens-
land border.
It must be noted here, that, near the border between Queens-
land and New South Wales, in the vicinity of Hungerford, there
are patches of granite at the surface, and also that some of the
bores in that vicinity have struck granite at comparatively
shallow depths There is here, then, indication of an island in
Lower Mesozoic time.
In the north-east of South Austialia, where the Lower Meso-
zoic sandstones are at great depths, it seems probable that the
strata above them include Cainozoic, Cretaceous freshwater-beds
(equivalent to the Winton Series of Western Queensland), and
Cretaceous marine beds (Rolling Downs Series). This is in-
ferred from a comparison with conditions in parts of Queensland;
in the great majority of bores, unfortunately, the records kept
are of little value for detailed geological purposes. In the case
of the Patchawarra Ijore in South Australia, which was aban-
doned at 5,458 feet without reaching the water-bearing strata,
it appears that the bore, when abandoned, was still in the Cre-
taceous rocks, and there is nothing to indicate that the Artesian
Series of sandstones does not exist further down. I am in-
debted to Mr. L. Keith AVard, Governnient Geologist of South
60 GEOLOGY OF LOWER MESOZOIC ROCKS OP QUEENSLAND,
Australia, for supplying me with all available information re-
garding this bore. Mr. Ward also remarks that the bores sunk
by the South Australian Government, with tlie exceptions of
Hergott and Marree at the margin of the basin, have failed to
reach bedrock. In other bores in the north-east of South Aus-
tralia, water is obtained from sandstones at great depths, in-
dicating surely the presence of equivalents of the Walloon Series.
e.g., Uoyder's Lagoon Bore, 4,700 feet: Mount Gason Bore, 4,420
feet; and others.
Examples of palteontological proof of the age of the sandstones,
carrying the large supplies of artesian water, are quoted above
{see p.o8). and these show that the presence of artesian water is
an indication of the existence of the Lower Mesozoic Sandstones,
and can be used in the determination of the extent of these
beds.
{y) Foldiny -movements In studying the folding-movements
to which the Lower Mesozoic rocks of Queensland have been
subject, two areas may be considered separately, viz., {a) the
western belt, extending from the Cape York Peninsula to the
New South Wales l)order, and (6) the occurrence in South-eastern
Queensland, (a) The western belt has not been subject to any
considerable movement, and the rocks usually dip gently to the
west and south-west beneath the marine Cretaceous strata; oc-
casional high angles of dip have been observed. (6) In South-
eastern Queensland, a study of the distribution and directions
of dip shows that the Lower Mesozoic rocks have been consider-
ably folded, and that the folding in some cases gives place to
faulting. The folding takes the foi-ni of a series of anticlines
and synclines, whose axes are in a direction approximately N.tJ0°
W.-S.30°E. The distribution of the Lower Mesozoic strata in
S<juth eastern Queensland has never been thoroughly understood,
but the recognition of this series of folds seems to explain the
distribution in a simple and reasonable manner. In places, the
folding gives place to faulting, e.y..^ north of Ipswich, and pro-
bably also near Hemmant. Mr. Ball has also described a fault
near Woodford,* which he believes to represent a continuation
■' Queensland (government Minin^^ Journal, xvii., 1916, p. 169.
BY A. B. WALKOM. 61
of the supposed fault at Heminant. The general directions of
the dip and of the axes of foldino- are shown in Plate ii.
The extent of this folding, which has aftected both the Lower
Mesozoic rocks and the overlying Cretaceous strata, has not been
generally recognised, as may be seen from the following quota-
tions. Dr. Jensen* says, "Our Mesozoic sediments show no
folding of consequence. Generally speaking, they show only
slight dips, and have never been under the influence of tangential
pressure like the Mesozoic sediments of the Alps, Himalayas,
Java, etc "' Dr. Richards,! writing of South-eastern Queensland,
says, "Folding-movements of only a very gentle nature have
taken place since the Palseozoic era."
Such statements are not consistent with the folding which
has been observed to the west of Ipswich, in the neighbourhood
of Hemmant; and in the Maryborough District, particularly on
\\oody Island, where the strata are often nearly vertical.
The nature of the folding is illustrated in two Sections (Text-
fig. 1 ), of which Section A has been prepared from a section drawn
by Mr. Dunstan, and Section B from a combination of sections
by Messrs. Cameron and Marks. Neither Mr. Dunstan's nor
Mr. Cameron's original section has been published, and I am
indebted to them for permission to modify these sections for
use here.
This folding may be quite adequately and reasonably explained
as the result of the adaptation of the earth's crust to a shrinking
nucleus. Such folding would, naturally, be expected along zones
of weakness, and the east coastal area of Australia is undoubt-
edly such a zone. Folding of this nature often passes into fault-
ing, and this may have occurred near Hemmant, where there
seems to be a line of faulting associated with folding, and also
in the Ipswich District, where, Mr. Cameron informs me, the
severe folding just south of Ipswich gives place to a fault further
north.
From the originals of the sections, it was possible to Estimate,
approximately, the amount of shortening of the earth's crust
* Proc. Roy. Soc. Queensland, xxiii., Pt.2, p. 163.
t Proc. Roy. Soc. Queensland, xxvii.,Pt.2, p. 114.
^///^///'
/ /. y
ri^V//
Text-tig. 1 A.
cQ
Text-tig. 1 B.
'I'ext-fig. lA. — Sketch-Section
from near Colton to Frazer
Island, sliowing the folding
of the Jurassic and Creta-
ceous rocks. Vertical scale
exaggerated about 2^ times.
(Modified from Section
drawn by Mr. B. Dunstan).
Text-fig. 1 B. — Diagrammatic
Section from Ipswich to
Manly, showing the folding
of the Lower Mesozoie
Strata. Vertical scale ex-
aggerated. (Modified from
Sections by Messrs, W. E.
Cameron, B.A., and E. 0,
Marks, B.A.).
BY A. B. WALKOM. 63
represented by the folding. Both sections are approximately at
right angles to the axes of folding. In the section between
Ipswich and the mouth of the Brisbane River, a shortening of
the order of 1|^ miles in .34 miles has been effected; and, in the
Maryborough district, a shortening of about 0*8 mile in 34
miles. This shortening is circumferential, and if it be regarded
as the shortening represented at this particular time for the
east-west extent of Australia (a not unreasonable supposition,
since we know of no other folding of the same age across Aus-
tralia), the radial shrinkage represented would be of the order
of 0-05%.
Regarding the age of the folding, it certainly took place after
the Burrum Coal-Measures (probably Lower Cretaceous) were
deposited, for both Marine Cretaceous strata, which are ap-
parently conformable with the Tiaro Series, and the overlying
Burrum Coal-Measures have been subject to this folding force,
and have been folded to an extent closely comparable with the
Lower Mesozoic strata. This is shown by a comparison of
Sections A and B.
This same folding took place before the Cainozoic, since it
has not affected any of the Cainozoic rocks of South eastern
Queensland. Also the disposition of the Cainozoic volcanic
rocks in South-eastern Queensland indicates that the Pre-Caino-
zoic drainage-system was approximately meridional, and this
may be closely connected with the N.30°W. strike of the folds
of the Lower Mesozoic rocks.
In the Ipswich District, there appears to have been a sub-
sequent period of folding. This is comparatively restricted, and
the folds produced by it have an approximately meridional
direction. The Cainozoic deposits have been affected by this
movement, which must, therefore, have been later than the
period of folding already mentioned. I am indebted to Mr.
Cameron for information regarding this latter folding.
Flora of the Lowek Mesozoic Rocks op Queensland.
The lists, presented below, of the Horas of the Ipswich and
Walloon Series have been prepared after a careful examination
64 GEOLOGY OF LOWER MESOZOIC ROCKS OF QUEENSLAND,
of the specimens available in the collections of the Queensland
Geological Survey, the University of Queensland, the Queens-
land Museum, and the " Simmonds " Collection. In addition,
previous records have been examined and verified as far as pos-
sible, and, only when it is believed that they are reliable, have
they been included. In cases where the original specimens have
been lost, the records have been used only when the determina-
tion is not subject to the least doubt. These lists, then, are
regarded by the author as being as complete and reliable as it is
possible to make them, in the light of our present knowledge.
(a) I/>stvir,/i. Seines. — The flora of the Ipswich Series comprises
the following species : —
Equlsetalh:s.
Equisetites rotiferum Tenison- Woods.
Equiseiites sp. (tubers).
Phyllotheca aitstralis Brongniart.
N eocalamiles luerensis (Schimper).
Neocalamites cf. Carrerei Zeiller.
Schizonenra cf. ajricaiia Feistmantel.
Filicales.
Osmundaceffi.
Cladophlebis australis (Morris).
C. Roylei Arber.
Cyatheacete.
Coniopteris delicatula (Shirley).
Dipteridinse.
DictyophyUum 7nvyosuin Lindley and Hutton.
Thinnfeldiea?.
Thinnfeldia Feistinanteli Johnston.
Th. laMci/oIia (Morris).
77/-. odontopteroides (Morris).
Th. acuta VValkom.
Marattiaceai.
Danfeopsi.s Hnyhesi Feistmantel.
Hydropteride8e('?;.
Sagenopteris rhoifolia (PresI).
BY A. B. WALKOM.
Of)
Genera of Ferns and Plant.?: incert.^ sedis.
Spheiiopteris hi.r.uiiosa Shirley.
Sph. superba Shirley.
TcEiiioptaris Tenison-Woodsi Etheridge Jr.
T. Carruthersi Tenisoii-Woods
T. lentriculiforme (Etheridge Jr.).
T. Duiistani Walkom.
T. ivianamattcp, (Feistmantel).
T. crassinfTvis (Feistmantel).
Stenojiferis elouyafa (Carruthers)..
GiNKGOALES.
Ghikgo aittarctica Saporta.
G. digitata ( Brongniart).
G. cf, tmtgnifoHa Fontaine.
Baiera Siiiimondai Shirley.
B. bidens (Tenison- Woods).
/> ipsvicieusis Shirley.
B. ginkgoides Shirley.
('?)Ginkgoales.
Stnchyopitys annularioides Shirley.
aS'. Simmojidsi Shirley.
CYCADOPHYrA,
Bennettitales.
Beiinettit-s ( Williamsonia) sp.
CyCADOPHYTA INCERT.E SEDIS.
Pterophyllum, multilineatum Shirley.
Gymnospernious seeds.
Examination of this list at once shows certain distinctive
features, the most notable being the comparatively large number
of Ginkgoales, the relatively fevv Cycadophyta, and apparent
entire absence of coniferous remains.* The small proportion of
* This statement maj' need subsequent modification. Silicified woods
from the Ipswich Series have, in the past, been described as Ai^aiicari-
oxylon spp. These woods are at present being re-examined by Professor
A. C. Seward, and it is quite possible that they may not be of undoubted
coniferous affinities. At any rate, we know of no remains from the Ipswich
Series, other than these woods, which may, at the present time, be referred
to the Conifers.
66 UEOLOGY OF LOWER ME80Z0IC ROCKS OF QUEENSLAND,
Cycads may ultimately be very greatly increased, if it should be
shown that the genus Tce/)iiopteris belongs to this group. In a
recent publication, Thomas includes species of Tceniopteris with
the Cycadophyta, as a result of his examination of the epidermis
of a number of recent and fossil Cycads.* None of the Queens-
land Lower Mesozoic examples of Tceniopteris obtained have
been preserved as carbonaceous films, and it is at present impos-
sible to state the nature of the structure of the epidermis in
them.
The list of species in this Series may be summarised, and the
figures expressed in percentages, as in the following Table : —
Tablk
i.
1
2
8
4
No. of
%
o/
o/
Species.
^
/o
Equisetales
15
15
15
Filicales
10
80
1
\
1
89
Filieales incertc-e sedis
8
9
57
Twniopteris
6
18
1
24
Cycadophyta
2
G
6
(linkgoales
7
21
21
21
Total ...
88
—
•
Such tables as these must be used with a good deal of caution,
as their indiscriminate use may lead to quite incorrect and even
absurd results; but careful use, with a full realisation of their
value and their drawbacks, may yield interesting and, to some
extent, reliable results. The use of such numerical methods has
not come into very general use, but their preparation and use
have been illustrated recently by VV ieland.f
In the above Table (Table i.) of the Ipswich flora, the species
of Stachyopitys, and gymnospermous seeds have not been used,
since they, in all probability, represent seeds or reproductive
organs of plants already represented in the list by sterile fronds.
The species of Tceniopteris are placed separately in the Table,
* Thomas, Q.J.G.S., Ixix.; p. 223.
t Anier. Journ. Sci., xxxvi. (1918), pp.268-278.
B. WALKOM.
67
since their position is uncertain. Tn column 3, the percentages
are given, including Tc^niojAeris with the ferns; in column 4,
this genus is included with the cycads. This latter result gives
the flora a not abnormal character in the proportion of ferns and
cycads, and indicates a general similarity to some lihsetic floras;
there are, however, certain points which may indicate a some-
what greater age, e.y., the high percentage of Equisetales, and
the rather smaller percentage of Cycads, than is usual in Rhsetic
and Jurassic floras.
For convenience of reference and comparison, the following
Table is quoted from Wieland : — *
Table ii. (after Wieland).
Elements of typical Rluetk-Oolitic Ft one.
^
-2
S
_d
% i
,'o
X
a;
oT a;
90
S
'^
S
■> 'o
-^.2
^i
2 ^
^ r/.
g^
90
S^
■o^
•^-^
i^ ^
o
o
>HhS
pq
K
d;^
hS
Ferns
42
40
37
30
32
18
48
Cycadeans
28
38
43
33 1
34 +
70
33
Conifers
27
12
),6
17
8
(•2)
9
(Tinkgos
4
0
•/
2
Cordaites
•>
9
9
8
2
Eqnisetums ...
2
-1
4
5
2
2
o'o
The flora of the Ipswich Series may now be examined in
greater detail.
Equisetales. — Thki percentage of Equisetales in the Ipswich
flora is unusually high for a Mesozoic flora. As I have already
pointed out,t they show attinities with the Equisetales of Rhsetic
floras. Neocalarnites hoereiisis occurs in the Rhsetic of Sweden,
and Neocal<imites Carrerei in the Rha?tic of Tonkin, and in the
Stormberg flora (Molteno Beds) of South Africa. Phyllotheca
australis is one of the few species which continue from the
Palaeozoic into the Mesozoic in Australia, occurrincr in the Per-
mian (Permo-Carboniferous) of Eastern Australia, and also in
* Amer. Journ. Sei., xxxvi. (1913), p.272.
t Queensland Geol. Surv., Publication No. 252, p. 38,
68 GEOLOGY OF LOWER MESOZOIC ROCK.S OF QUEENSLAND,
the Wianamatta Stage of the Hawkesbury Series of New South
Wales. Schizoiienra africaiia occurs in the Beaufort Series in
South Africa (Permian), and, according to Seward, is related to
iS'. yondwa7ieiLiiis from the Talchir, Damuda, and Panchet Series
(Permian and Lower Triassic) of India. The remaining species,
Equisetites rotiferum, is a widespread type, and very similar
forms are widely distributed in both Rheetic and Jurassic strata.
Filicales.— A majority of the true ferns of the Ipswich Series
indicate a Rhtetic or possibly greater age for the flora. Sterile
Cladophlebis-ironds of the C. anstralis type are of very wide-
spread occurrence in rocks varying in age from Rhsetic to Middle
Jurassic. Cladoph/ebis Roylei is a somewhat older type, occur-
ring in the Raniganj Series (Upper Permian) of India. Dictyo-
phyllnm riigosum is a fairly widespread Jurassic (Lower Oolite)
species, but there is so little difference between it and species of
Dictyophyllum in Rhsetic floras, that it cannot be regarded as a
reliable indicator of age. The species of Thinnfeldia in the
Australian Lower Mesozoic rocks do not appear to be represented
in the Northern Hemisphere. The three species, T. Feistmait-
teli, T. odontopteroides, and T. lancifolia, are present in the
Molteno Beds in South Africa; and T. odoiilojiteroides and T.
lancifolia in the Mesozoic flora in Argentina. Dancr-opsis
Hughesi is a Rhretic or older form, occurring in the Rhsetic Beds
of Tonkin, the Burghersdorp Beds of South Africa (Triassic),
and the Middle Gondwanas of India (Triassic). Sagenopteris
rhoifolia is itself a Rhsetic species, but it is at times difhcult to
distinguish between it and S. Phillipsi, a common Jurassic form.
Ginkgoales. — The Ginkgoales are represented by a compara-
tively large number of species, and, in addition, they are also
relatively abundant as regards numbers of individuals. Ginkgo
antarctica occurs in the Lower Mesozoic rocks in New South
Wales, but there is no record of its exact locality or horizon. G.
digitata is a very variable form met with in both Rhsetic and
Jurassic rocks. G. magnifolia occurs in the Jurassic strata of
Oregon, and also bears some slight resemblance to Baiera storm-
bergensis from the Stormberg flora of South Africa. Baiera
bidens is a very common type, and is similar to species which are
BY A. B. WALKOM. 69
abundant in both Rhsetic and Jurassic rocks. B. ipsviciensis
may be compared with B. multifida, occurring in the Rhsetic of
North America: and B. Simniondsi also occurs in the Wiana-
matta Stage of the Hawkesbury Series of New South Wales.
Stachyopilys annularioides is a very similar type to specimens
referred to as Stachyopilys sp., in the Stormberg flora of South
Africa, and also to Sphenolejns rhcetica from the Pvhsetic of San
Juan (Argentine).
Gycadophyta. — The percentage of species of this group is un-
usually small in the Ipswich Series. Should the genus Tceui-
opteris ultimately prove to be a cycad, however, the percentage
of cycads in the Ipswich flora would be a normal one for eaily
Mesozoic floras. The specimen showing expanded bracts, re-
ferred to Bennettites ( Williamsonia) sp., cannot be regarded as
indicating any special age, as it agrees with similar specimens
from the Mixteca-Alta tiora of Mexico (Rhsetic-Liassic), and also
with B. Carruthersi from the Wealden of England. Fterophyllum
multUineatiim occurs in the Rhajtic of Tonkin, and similar
species are found in the Burghersdorp Beds of South Africa.
Genera incerkB sedis. — The species of Sphenopteris do not offer
any evidence as to the age of the Series. Species of Tceniopteris
are not usually to be regarded as reliable indicators of geological
age. Of the species in the Ipswich Series, some appear to be
confined to Australia, while others of them occur in South Africa
and India. 2\ Tenison- Woodsi and 7\ Carruthersi are found in
the Stormberg flora (Molteno Beds), and the latter also in the
Burghersdorp Beds in South Africa; T. crassiuervis occurs in
the Rajmahal Series (Lias) of India. Stenopteris eloiigata occurs
also in the Stormberg flora of South Africa.
The accompanying Table (Table iii.) shows the relationships
of the species of the Ipswich flora to species in other floras. The
Table is arranged in four columns, column 1 showing the species
with affinity to species older than Rhsetic, column 3 those with
affinity to species in Rhsetic floras, and column 4 those of Jurassic
affinity. The Stormberg flora of South Africa is so similar in
general appearance, that the species common to the two are
grouped separately in column 2.
70 GEOLOGY OF LOW KR MESOZOIC ROCKS OF QUEEXSLAXD,
o
% I
< 2
CO
4)
'i
'■re"
'5
^
"/'
4>
&
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^
_
y<
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yT
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■Z
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c
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^
^
Th
5
.4j
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-
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be
• —
o
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>;
y
•—
,
i
i
:?
i3
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_r
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~j
5
-9
"3
3
E
J^
15
?
^
J
^' I?
^
"Z -t
■~ ^
oc •'^ :;
i^
_:;
r
V. :^
■C:
S <
^ • —
HI
1
2.
c"
5
o* 0
11
"5
1
_r <J^ ^ .i;
^ •= X
n1
"" ^
.^
c
t:
©
S ^
s
bc
S '0
^
i
!?
«}
•^ s c I- ^ -I i
•§ i .^' "i - 1' I i,
^ f ^ .11 5 J t-^
-^ c ■= ^ 2 <i"^ ■>.•
c> ?- c-~ f-" v; ;: '^ "^
.2 S
X ID
o -a
§ 2
'^ —
^ o ^
'^ ,<K ^ :t u
9^v ■- = - ^
C- S ?:■
^ ;r -a
it
^-=3
<SC.S^ «<i;S ;^bc^3
=^ 05 o c:i ^ a.
BY A. B. WALKOM. 71
This Table at oDce shows that the flora of the Ipswicli Series
is of a facies at least as old as Rhaetic. In addition to the species
occurring in Uhietic floras, there are also a number of species
which occur in other parts of the world in rocks older than
Nhsetic. The number of species in the Ipswicli flora, which
represent Jurassic types, is not large, and nearly all of these
are examples of species in which it is often difficult to find satis-
factory distinctions between Rhsetic and Jurassic species. The
only two species of the Ipswich flora, which are characteristic of
Jurassic rocks, are T'ceniopteris crassinervis and Ginkgo magtii-
folia] and, regarding the latter of these, I have a slight doubt
as to the determination in the Ipswich Series
Of twenty-two species in this 'J'able, eighteen occur in Rhtetic
floras, six occur in floras older than Rhsetic, and seven in Jurassic
floras, only two of the latter, however, being characteristic of
Jurassic floras.
The Table, together with the foregoing discussion, shows fairl}^
conclusively that the flora of the Ipswich Series must be regarded
as at least as old as Rhietic, and probably somewhat older.
[h] Walloou Serias. — The flora of the Walloon Series comprises
the following species : —
Equisktales.
Equisetifes rotijerum Tenison-Woods.
Equisetites cf. rajinahalensis Oldham and Morris.
Schizoiieura sp.a Seward.
Schizo'iieura sp.
Filicales.
(1;Osmundaceye.
C'lodophlebis aiistralis (Morris).
C. Roylei Arber.
{?)Matonineie.
Phlebopteris aleihopUroides Etheridge Jr.
Dipteridinse.
Dictyophyllum riigosumil) Lindley and Hutton.
D. Davidi Walkom.
UausmaniLia(\) Buchii (Andrjc).
I GEOLOGV OF LOWER MESOZOIC ROCKS OF QUEEXSLAS-D,
Thiniifeldiese.
Thiniifeldia Feistmaitteli Johnston.
Th. odontopteroides (Morris;.
Th. Imtcifolia (Morris).
(IjHydropteridete.
Sayenopteris 7'hoijolia (Presl).
Genera incehtve sedis.
Spheiioi^teris snperba Shirley.
Stenopteris elongata (Carruthers).
Phyllopteris Feistma/itteli Etheridge Jr.
T(eiiiopteris spatulata McClelland.
T. spatulata var. ynajor Seward.
T. Tenison-Woodsi Etheridge Jr.
1\ Carrttthersi Tenison- Woods.
T. lentricidi/orme (Etheridge Jr.).
T. crassiiiervis (Feistmantel).
GiNKGOALES.
Ginkgo magnifolia Fontaine.
Baiera Simmoiidsi Shirley.
Cycadophyta.
Bennettitales.
Ptilophylluiu ( WUliamsoiiia) pecten (Phillips).
Cycadophyta incert^. sedis.
Pterophylluni ahuorme Etheridge Jr.
P. contiguum Schenk.
P. Nathorsti (Seward).
Pseudoctenic eathiensis (Kichards).
Otozatnites queenslatidi Walkom.
0. obtusus (Lindley and Hutton).
0. Feistmanteli Zigno.
0. Mandelslohi Kurr.
Coniferales.
Araucarites polycarpa (Tenison-Woods)
Br achy phy Hum crassuia Tenison-Woods
Taxites planus Feistmantel.
Genus incert^ sedis.
PhoRuicopsis elongatus{]) (Morris),
BY A. B. WALKOM.
73
The outstanding feature of this flora is the large increase in
the number of cycads. Other noticeable points are the decided
decrease in the number of species of Ginkgoales, and the presence
of a few Conifers.
Arranged in tabular form, and omitting from iht'Tshle Schizo-
neura sp., which may possibly be the external casts of which
S. sp. a is the pith-cast, and also Fhamicopsis elongat2i,s, whose
position is quite unknown, the following result is obtained.
Table iv.
• 1
2
3
4
No of
species.
%
%
%
lOqiiisetales
8
S
8
8
Filicales
10
28
1
/
36
Filicales incertae sediy
3
8
53
Taniiopteris
6
17
}
42
Cycadophyta
9
25
25
(rinkgoales
2
6
6
6
Coniferales
3
8
8
8
Total
SQ
—
This Table shows distinct difierences from that of the Ipswich
flora (Table i., p. 66), and when, as in column 4, the species of
Tce7iiopteris are combined with the Cycads, the percentages agree
quite well with the percentages in Jurassic floras, particularly
with the Lower Oolite Hora of Yorkshire {see Table ii., p,67).
The flora of the Walloon Series may now be discussed in more
detail.
Eqiiisetales.—M.em\)ers of this group are not of uncommon
occurrence in the Walloon Series. Equisetites rohferuni is
almost indistinguishable from the widely spread E. columnaris
of Middle Jurassic floras, and Equisetites cf. rajmahalensis is
closely comparable with the species in the Liassic flora of India.
Schizo7ieurafiij.a is identified with the species described from the
Stormberg Beds of South Africa. Pith-casts of Equisetales,
however, cannot be regarded as of any value in difi'erentiating
between Rhsetic and Jurassic floras.
74 GEOLOGY OF LOWER MESOZOIC ROCKS OF QUEENSLAND,
Filicales. — Tlie ferns, although not particularly numeious,
have the general appearance of a Jurassic tiora. Cladophlebis
australis, as already stated, is a type of frond of widespread
occurrence, but is specially close to the C. denticulata-iy^e, a
widespread Jurassic form. Phlehopteris alethopter aides is, in all
probability, very closely allied to Laccopteris polypodioides from
the European Lower Oolite, though there appears to be a differ-
ence in the venation. DiGtyophyllurti rugosum is a Jurassic
type, but its occurrence in the Walloon Series is doubtful.
Hausmannia Buchii, with which some specimens from the Wall-
oon Series have been compared, occurs in Jurassic floras of both
Liassic and Kimeridgean age. Sage7wpteris rhoij'olia, though
itself a Rhsetic species, is, at times, indistinguishable from the
Jurassic S. Fhillipsi. The species of Thinnfeldia, and Stenopteris
elongata in the Walloon Series are survivals from the Ipswich
flora. The survival of a number of species from the Ipswich
flora to the Walloon flora, and even to later floras, in Queens-
land is to be expected, since there were no violent earth-move-
ments during these times, and no marked changes are known,
which might have been expected to lead to any unusual dying
out of the older flora. It is indeed fortunate, for stratigraphical
study, that there are so many forms in the Ipswich Series, which
apparently have not survived into the AValloon Series.
Ginkgoales. — There has been a remarkable change in the
number of members of this group. In the Ipswich Series, we
have at least seven species represented, while, in the Walloon
Series, there are only so far two. Ginkgo magnifolia is probably
identical with Fontaine's G. IhUtoni var. magnifolia from the
Jurassic of Oregon. The other species, Baiera Sinimondsi, is
another survival from the Ipswich Epoch.
Cycadophyta. — Cycads form the most prominent element of
the Walloon flora, in which they are present to the extent of at
least 25 per cent, of the species, and, perhaps, (if Tceniojderis is
a Cycad) 42 per cent. On the whole, they indicate very dis
tinctly the aspect of a Jurassic flora. Ftilojjhyllum ( Williavi-
soitia) pecten is a very widespread tjpein Jurassic (Oolite) floras.
Up to the present, there is no indication of any Williamsonia-
BY A. fi. WALKOM.
75
flowers associated with these sterile fronds of FtMophyllum
ppxten. Pterophylhim A'athorsli occurs in the Jurassic of Suth-
erland, Scotland, and very similar forms occur in the Jurassic of
Oregon ; Ptenyphyll nni contigiiuin occurs in the Jurassic of
Oregon, and also in the Rhsetic of 'J'onkin. Pseiidoctenis eathi-
eiisis is practically identical with specimens from tlie Jurassic of
Yorkshire and Sutherland. The three species of Otozamites, 0.
ohtusiL.s, 0 Feisimariteli, and 0. Maiidelslohi are species occurring
widely in Jurassic rocks, and 0. Mandehlohi occurs in the
Mixteca-Alta flora of Mexico.
Coniferales. — Remains of Conifers are not abundant in the
Walloon Series. Cones have been described, and referred to the
genus Araucarites, indicating that they are similar in general to
cones of the present-day Araucatia. A few fragments have
been referred to the genus Br achy phy Hum, and some specimens
to Taxites. The specimens referred to Taxites planus show a
very close resemblance to, and are probably identical with, that
species, as described from the Upper Gondwana Beds (Liassic)
on the Madras coast of India. Similar forms to Brnchyphyllum
crassutn occur in the English Jurassic floras.
The accompanying Table (Table v.) shows the flora of the
Walloon Series arranged in two columns, column 1 including
those species comparal)le with species occurring in beds older
than Jurassic, and column 2 those comparable with Jurassic
species.
Tabi.e v.
Flora of ^Valloon (Series, showing affinities of species with regard to age.
Species with affinit}^ to ,
Species older than Jurassic.
Species u-jth affinity to Jurassic
Species.
Equ ixet ite>< rot iftra m
Schizoneara sp. a
(J/ a. doph I eh is a ns( ra/is
C. Roy hi
Th in nfeJdia Ftistma ntel i
Th. odontopteroides
Th. laiicifoUa
Equisedtes rofiferum (M. Jur.)
E. cf. rajmahaltnsus (Liassic)
Cladopldehix ausfra/is (L. Oolite)
Phhhopferis ahthopteroidt.^ (L. Oolite)
Dictyophylhim rwjosum (L. Oolite)
Htoi-^inaiiiiiii (''.) /Jiir/iii {]j\tiHH\c; Kimeridge)
Sagtnopteris rhoifolia {S. PJiiHipsi; Oolite)
76 GEOLoi'iV' OF Lower Mesozoic rocks of Queensland,
Table v. — (continued).
1
Species with affinity to
■
Species with affinity to Jurassic
Species older than Jurassic.
Species.
Sagaiopferis rhoifolia
Phyllopteris FeistmauteU
Sttnopteris e/oiif/afa
Titmiopferii< spatulata (Lias)
Tcvniopterl.^ Tenison- Woodd
T. spatulata var. major
T. Carrufhersl
T. crassmerri.s (Lias)
G inkgo maf/n ifol ia
Ghil-(jo DtfKjnI/o/ia (Jurassic of Oregon)
Baiera Simmoitdxl
Ptilophylhnu [ Williamsonm) pecten (Oolite)
Pterophyllnm ronti'juum
Pterophyllum confi;/iiiim (Jm-dSBiG of Oregon)
Otozamites Mandelslohi
P. Xaihoysti (Kimeridge)
Pseudoctenis eafhiensis (Kimeridge)
Otozamites ohtusus (L. Oolite)
0. Feistmanttfi (L. Oolite)
0. Mandtlsfohi (Lias)
Brachyphyllum crassum
Taxites planus (Lias)
The ages of the species, or of very closely related species, in
cohimn 2 are indicated in brackets after each one.
Examining this Table, we find that, excluding the six species
common to the two lists, there are nine species with affinities to
species older than Jurassic, and fifteen species with Jurassic
affinities. Of the nine species regarded as older than Jurassic,
eight occur also in the Ipswich Series, and are the species which
survived from the Ipswich to the Walloon Epoch; there is only
a single species (and that an equisetaceous pith-cast, Schizo7ieura
sp.a) which could be considered as an argument in favour of an
age older than Jurassic for the Walloon Series; whereas there
are fifteen species characteristic of Jurassic floras, which appear
in the Walloon Series, and were not present in the Ipswich
Series. No other conclusion, then, seems possible, than that the
flora of the Walloon Series indicates that the Series is homo-
taxial with strata of Jurassic age, and the greater number of
species (see Table v.) indicate a lower Jurassic age, Liassic or
Lower Oolite. It is possible that the Walloon Series represents
deposition over a period covering both the Liassic and Lower
Oolite of Europe.
BY A. B. WALKOM. / /
(c) Ccmipariso'n of floras of the Ipsivirh and WaUoon Series. — -
We are now in a position to compare the fioras of the Ipswich
and Walloon Series. As a result of all previous examinations
of these floras, it has been observed that there was no palseonto-
logical distinction between the floras of the two Series; e.g.^ Mr.
Cameron states •* "It does not, however, seem possible to draw
any distinction between the formations from pala?ontological
evidence, most of the fossils as yet found in the Walloon Beds
occurring also in the Ipswich Beds."
The failure to note palseontological distinction between the
two has been due, however, to imperfect collecting, and to the
fact that the material available has not been thoroughly ex
amined since the examination by Mr. Etheridge, prior to the
publication of the " Geology and Palaeontology of Queensland
and New Guinea," in 1892. In 1898, Dr. Shirley examined and
described a number of specimens, but all the Lower Mesozoic
specimens among them were from the Ipswich Series. The same
author, in 1902 recorded a few more specimens from the Ipswich
Series, and some from the Stewart's Creek Beds (of Walloon age).
At the time of these publications, however, there was no subdivi-
sion of the Ipswich Formation into Ipswich, Bundamba, and Wal-
loon Series, so it is not remarkable that no distinction of horizons
was noted in the upper and lower portions of the Formation.
A glance at the following Tables (Tables vi. and vii.) at once
shows that there are points in which the two Series can be dis-
tinctly separated, and that there are a number of species char-
acteristic of each. These species, characteristic of only one
Series, are the more important ones from a stratigraphical point
of view, and the finding of some of them may, in the future, be
used for fixing the age of the strata in which they occur.
<^)ueensland Ueol. Surv., Publication No.204, p. 16.
78 GE0L0r4Y OF LOWER MESOZOIC ROCKS OF QUEENSLAND,
Table vi.
Ipswich Series.
Walloon Series.
No
of species.
No.
of species.
Equisetales
;■)
3
Filicales
10
10
Cienera incertte sedis —
a, probably Filicales
^
3
//, Titniopteris ...
6
6
Cycadophyta
9
Ginkgoales
/
2
Coniferales
83
3
36
Table vii.
Table sliowing distribution in Queensland of the Species of Fos.sil Plants
in the Lower Mesozoic roeks.
i
Walloon Series.
T.
^
u
s
X
2
«
= !
ijB
1
ij
02
=1
%
5
1
^ ■
i
IS
o
.5
= 1
i
&
xiS
3i
H
c; 1
Equisetales.
Equisetitei^ rot i/h-um
X
X
X
(?)
E. cf. rajmahalensis
X
PhyUotlieca austral i^
X
•
Xeoca/amifes lufrensis
X
X. cf. Carrerei
X
Schizoneura cf. africana
X
Schizoneura sp. a
X
X
Filicales.
CladophltbU aiisfra/is
X
X
X
{•?)
X
X
X
(?)
C. Roylei
X
X
Coniopteris delicatitla
X
Ph/ebopteris alethopteroides . . .
X
X
Dictyophyllum rugosum
X
.(V)
(?)
D. Davidi
X
i
Hausmannia Biichii{^.)
X
i
TJi hi ii/e/dia Feist mantel i
X
X
i
X
Th. odonfopteroides
X
X
Th. lancifolia
X
X
Th. acuta
X
Daiui'opsis Hiiyhesi
X
Sagenopteris rhoifoUa
X
X
Genera of Ferns and Plants
•
incertae sedis.
Sphenopteris lacunosa
X
I
'
BY A. B. \YALKOM.
79
Table vii. — {continued).
.1
Walloon Series. !
to 1
4i
-
2
' «
5^
^
*>
i
1
it
a
-5 «
^1
6
.2
5*
.S
-j
Ik
c
iJ ~.
75
*
SI
1
E
o
-55
&
2^
It
S
Sphenopferi-'i i^nperJ)a ...
X
(?)
Sfenopferifi eJongafa ...
X
(?)
1
J'hi///opieris Feistmanfe/i
(?)
X
1
Tiifntopteri-y fipafuhifa
X
X
K
X
X
T. ■•^patnlata var. major
X
(?)
X
T. Ttni.<<on-WoocUi '
X
X
X
T. CarrntherM
X
X
T. lenlricu/l/orme
X
(?)
T. Diinstani ...
X
T. icianamatfte
X
T. crasMnern's
X
X
X
Ginkgoales.
(rinh-io a ntarctlca
X
G. di flit at a
X
G. cf. maynifoUa.
X
X
Baiera Simmond^i ...
X
X
B.hide».s
X
/>. ip/.'^ricienM.'s
X
B. i/inkyoides ...
X
(?)Gink»oales.
Staehi/opifi/--! annjilarioide-^ . . .
X
S. Simmond.'<i ...
X
Cycadophj'ta.
Bennettitefi (Widiamsonia) sp.
X
PtUophyUnm ( WiUiamsonla)
ptcten
X
Pferophydnm ahnormc
X
Ft. coiififfiiKm
X
Ft, innltilineatum
X
Ft. Xathorsti
X
F.-<eudocteuis eathieuMS
'
X
Otozamites queenslandi
i
X
0. ohtUSHS
X
0. Fei.^t mantel i
X
X
0. cf. Mandela/ ohi
X
Conifeiales.
,
A raucarite-s poJycarpa
X
Brachyp)hydum crasmm
X
X
: (?)
(?)
Taxitex planus
1
X
X
(?)
Genus ineertse sedis.
!
Fhoenicopsis elongatus
X
(4ymnospermous seeds
X
i
80 GEOLOGY OF LOWER MESOZOIC ROCKS OF QUEENSLAND,
The marked difference between the two floras shows in the
Gyninosperms. In the Equisetales and Filicales, there is little
difference as regards number of species. Only two of the species
of Equisetales, and seven of the Filicales, however, are common
to the two Series; and, of the others, those in the Ipswich Series
are of older type than those in the Walloon Series. When the
Gvmnosperms are examined, however, a marked distinction
between the two Series is to be observed. Twenty-two species
are described from the Lowei* Mesozoic rocks, and only two of
these species are common to the Ipswich and Walloon Series.
These two species are Ginkgo cf. magnifolia and Baiera Sim-
mondsi. Ginkgoales are abundant, both as regards number of
species and number of individuals, in the Ipswich Series, but are
only poorly represented in both respects in tlie Walloon Series.
Cycads are represented very sparsely, as regards number of
species, in the Ipswich Series, whereas they constitute perhaps
the most prominent feature of the Walloon flora, being very wide-
spread in distribution and occurring in large numbers on some
horizons. The species of Otozamites form one of the most char-
acteristic and distinctive features of the Walloon Series. Up
to the present, no Conifers are known from the Ipswich Series,"*^
whilst there are three species representing three genera from
the Walloon Series.
Table viii. shows the species occurring in Queensland, which
are, so far as we know, found only in one Series of the Lower
Mesozoic rocks.
Table viii.
Species known only in the
Ipswich Series.
l^ltylJotlieca australis
Ntocalamiles ha^renAii^
X. cf. Carrerei
Schizoneura cf. a/ricana
Coniopterift delicatula
Thinnfe/dia acuta
JJan(eop,sis Hughesi
!Spli e. nopt 1 1 • /.v lacunosa
Tceniopttris Dunstani
T. wianamattcE
Species known only in the
Walloon Series.
Ph/ehojjferis al ethopterokles
Dictyophyllum Dacidi
Ha usniannia{ V) Buck ii
PhyUopteri.s FeiMmanfeii
Twniopttrix spatn/ata
T. spatidata var. major
FtilophyUum ( Wif/ia/n.-^oiiia) ptcttn
HtcropJiyllnm alnionnc
F. contiguam
F. Nathorsti
* See footnote on p. 65.
BY A. B. WALKOM. (^ 1
Table viii. — {continued)
Species known onlj^ in the
Ipswinh Sexies.
(rinkffo antarctica
a. dujitata
Ha ( era hkhn.'^
/>. </ink!/otf/e-'<
Sfarh f/ojt/fi/'« a n n u/a rioidc-<
S. Sim/nond><i
Bennett itex {Will iaui-^oiiiu) sj). ' Taxitea planiu
Fterophi/lliim nudtilinea/ii/ii \ Ph<eiiiropsi-'< elon(/afii>
(iyninospernious seeds i
Species known only in the
Walloon Sei^ies.
P'^endodenis eathienm.^
Otozam ife-'i qncen-'ilaiidi
O. ol>t)l.s,l.-<
O. Ftisfmaiitell
0. MandeJ.'^loli i
A rancarites pohjcarpa
llrarhy Jill yll urn crax'Oiin
(d) Age of the Queensland Loive^' Mesozoic Rocks. — The evi-
dence of the fossil floras of the Ipswich and Walloon Series is
very strongly in favour of a Triassic Age for the former, and a
Jurassic Age for the latter.
The flora of the Ipswich Series shows marked resemblance to
floras which are regarded as Rhsetic in age in other parts of the
world, particularly those of Tonkin and South Africa. Pvegard-
ing the actual position in the Geological P^ecord of many of the
occurrences classed as Rhsetic from their fossil flora, I have re-
ceived some interesting information from Dr. A. L. du Toit, of
the South African Geological Survey.*
Dr. du Toit has come to the conclusion that some of the so-
called Rhajtic strata would be better placed in the Upper Trias
( Keuper) than in the Rhsetic. He includes among these the
strata in South Africa, Tonkin, and Persia. In the case of South
Africa, he finds this is borne out by an analysis of the vertebrate
fauna of the Red beds and Cave sandstone overlying the Molteno
Beds, which is of Triassic ati&nities. He concludes that the Jiora
of the Molteno Beds is of Keuper Aye and not younger.
The Ipswich flora is undoubtedly very similar to that of the
Molteno Beds (see Table iii., column 2), but the finding of Gloss-
opteris in the latter suggests that it is possibly slightly older
than the Ipswich flora.
* Letter dated 2nd December, 1916.
82 GEOLOGY OF LOAYER MESOZOIC ROCKS OF QUEENSLAND,
From the evidence available, M'e are justified in stating that
the flora of the Ipswich Series indicates distinctly an Upper
Triassic age, possibly Rhsetic, but probably older.
The flora of the Walloon Series is decidedly of a Jurassic type,
and the typical Jurassic forms in the Series seem to consist of
about equal numbers of species characteristic of Liassic and
Lower Oolitic strata. The Walloon Series is of very great thick-
ness (probably of the order of 10,000 feet), and when it has been
studied in greater detail, it seems quite probable that it may
have to be subdivided into a number of stages. For the present,
we may regard the Walloon Series as representing a portion of
the Lower Jurassic System corresponding at least to the Lias
and Lower Oolite.
It is very diflicult to draw any reliable conclusion as to whether
the Bundamba Series should be placed with the Ipswich Series
in the 'I'riassic, or with the Walloon Series in the Jurassic.
There is no fossil evidence which may be taken as a guide.
Both Mr. Cameron and Mr. Dunstan have drawn my attention
to the usual association of the Bundamba Series with the Ipswich
Series, rather than with the Walloon Series; and, on this account,
it is, for the present, tentatively placed in the Triassic System
with the Ipswich Series.
Correlation of the Lower Mesozoic Rocks of Queensland
WITH those of other AREAS IN AUSTRALIA.
In considering the correlation of the Queensland Lower Meso-
zoic rocks with those of the other States, their relations with the
States will be discussed separately in the first place, and then
the results summarised by drawing up a table indicating the
relative positions of the various occurrences.
(a) New South Wales. — In New South Wales, the strata of
Lower Mesozoic age comprise (a) the Hawkesbury Series, includ-
ing the Narrabeen Stage, Hawkesbury Sandstone Stage, and
Wianamatta Stage, to which a Triassic age has generally been
assigned; (b) the Talbragar Beds, the Clarence Series, and the
Artesian Series, which have been regarded as of Trias-Jura age.
BY A. B. AVALKOM. 83
The Hawkesbury Series is divided into tbi-ee Stages, viz. : —
Xarrabeen, Hawkesbury Sandstone, and Wianamatta. in ascend-
ing order. The Narrabeen Stage consists mainly of sandstones
and shales, the Hawkesbury Stage of massive sandstones char-
acterised by current-bedding, and the Wianamatta Stage mostly
of shales.
This Series comprises a basin, and is apparently conformable,
in the central portion of the basin, with the underlying Permian
(Permo-Carboniferous) System; but, in the marginal areas, there
is an overlap amounting to unconformity. The Hawkesbury
Sandstone Stage rests conformably on the Narrabeen Stage,
while the Wianamatta Stage occupies a slightly eroded basin in
the Hawkesbury Sandstone.*
The Talbragar Beds cover only a small area, and occupy a
basin eroded in the Hawkesbury Sandstone; and there is no
doubt of an unconformity, stratigraphical as well as palseonto-
logical, between the Hawkesbury Sandstone and the Talbragar
Beds.
The Clarence Series occurs in the north-east of New South
Wales, between the New England Tableland and the coast; and,
at its northern end, it is continuous with the Walloon Series of
Queensland. Reference to the latest Geological Map of New
South Wales shows that the Clarence Series is practically con-
tinuous with the Walloon Series in the neighbourhood of Mt.
Lindsay, and also that, further west, it is continuous with the
eastern branch of the belt of Walloon Series on the Darling
Downs, extending through Warwick and Killarney to the border
of New South Wales.
The Artesian Series of New South Wales exists on the western
margin of the New EngLind Tableland, and extends along the
eastern margin of the Great Australian Artesian Basin. It is
practically continuous into Queensland with the belt of Lower
Mesozoic rocks extending past Warwick and Toowoomba. It is
seen, then, that the Clarence Series and Artesian Series are each
directly connected with the belt of Walloon Series in the Darling
" See Carne, N.S.W. Handbook, B.A.A.S.. 1914, pp. 601 -607.
84 GEOLOGY OF LOWER MESOZOIC ROCKS OF QUEENSLAND,
Downs, and they must, therefore, be identified with one another
as regards age, on the evidence of their field-occurrence alone.
Lithologically, the Clarence Series is divisible into three divi
sions, the middle one consisting of a series of massive sandstones.
This lithological division led to a consideration of the possibility
of the three stages being the equivalents of the Narrabeen,
Havvkesbury Sandstone, and Wianamatta Stages of the Hawkes-
bury Series. Mr. Carne* has, however, shown, in a convincing
manner, that this is not the case.
Another possibility that must be considered is, that the three
stages of the Clarence Series might be the equivalents of the
Ipswich, Bundamba, and Walloon Series of Queensland. Un-
fortunately, the Clarence Series has not been examined in great
detail geologically, but the fact, that Carne indicates the presence
of 'Tcenioptey^is spatulata \T. Daintreei] in the lower part of the
Series,! is sufficient to render the correlation of any part of the
Clarence Series with the Ipswich Series improbable. It is pos-
sible that the sandstones and conglomerates at the base of the
Clarence Series may be the equivalents of the Bundamba, but,
on the present evidence, I believe that the greater part of the
Clarence Series (if not all of it) is to be correlated with the
Walloon Series of Queensland.
Fossil plants are present in varying abundance in the various
Lower Mesozoic Series of New South Wales. No thorough
examination of the flora of the three Stages has been undertaken,
and, therefore, the compilation of satisfactory lists of the floras
of the different Series is not an easy matter. The following lists
have been drawn up from descriptions and lists published from
time to time, the majority of the determinations of New South
Whales Mesozoic plants having been made by Messrs. R. Ether-
idge Junr., and W. S. Dun. Names included in square brackets in
these lists are concjusious or comparisons suggested by myself to
bring, where possible, the New South Wales determinations into
line with my own work on the Queensland flora.
* Carne, Mem. Geol. Surv. N.S. Wales, Geology, No. 6, 1908, pp.31-4n.
t Carne, op. cit., p. 34.
BY A. B. WALKOM.
85
List of fossil plants recorded from the Lower Mesozoic rocks
of New South Wales : —
i. Hawkesbury Skries.
(a) Narrabeeii Stage.
77iinn/'eldia, u.sp.
Sphenopteris sp.
Oleaiidridium^ TcBuiopteris sp.].
MacrotcBniopteris \_TcBniopieris
sp.].
Tffuiopteris cf. McClellandi.
Ginkgo dilatata var. lata.
Ehipidopsis ginkgoides var.
Suss7Jiih/i{.
Araucarites (cones).
Ih'achyphylluni{%).
(b) Hawkesbury Sandstone vStage.
Equisetiun.
Phyllotheca.
Schizoneura aus trails.
Alethopteris sp.
Cladophlebis cf. Roylei.
Thinnfeldia iiarrabee'n.ens is
T. lancifolia.
T. odontopteroides .
T. oo?o?i^o^7>^e?-oic/(3s( Hawkesbury
type) [ = T . Fe{stniaiiteli\.
Ottelia prcEterita.
Phyllotheca concinna.
P. Hookeri.
Equisetum.
Aletliopteris.
(c]
Phyllotheca Hookeri.
P. australis.
Thinnfeldia odontopteroides [7'.
Feistmanteli and T. odonto-
pteroides].
Thinnfeldia, small vai*. lanci-
folia-type.
Cycadopteris scolopendy'i^ia.
T'hiiinfeldia odontopteroides \7\
Feistmanteli^.
Macrot(E7iiopteris.
M. ^vianamattce.
Oleandridium lentriculiforme.
Wianarnatta Stage.
Macrotmniop)te' is iviaiiamattcE.
Sphenopteris
Peco}>teris(^.) tennifoUa.
Gleichenia ditbia{l).
Alethoptei'is australis [Clado-
phlebis australis]
Baiera 7nultifda[ B. Sim mondsi]
Pterophyllum{]).
Talbragar Skries.
Thinnfeldia odontoptei'oides.
Thin7ifeldia sp.
JVeurojiteridiniii australe.
Sphe^iopteris sp.
'Tceniopte^'is Davntreei [T. spa-
tulata'].
Podozainites lattceolatus.
/\ spathulatus.
P. lotigifolius.
Podozamites sp.
Taxites cf. 2\ plasms.
>^ m i^
86 GEOLOGY OF LOWER MESOZOIC ROCKS OF QUEENSLAND,
iii. Clarence Series.
Alethopte7'is australis [Clado- Tceiiiopteris Daitttreei [7\ spa-
phlebis australis]. tulata\
Thinnfeldia odoufopteroidas. T. Tenison- Woodsi.
Sphenopteris.
iv. Artesian Series.
Tceniopteris Daintreei [7'. spxt- Thinnfeldia odontopteroides.
tulata]. Baiera.
The plant to which special prominence has been given, in
dealing with Australian Lower Mesozoic strata, is TcEuiopleris
spatulata [T. Daintreei], This plant is of very widespread oc-
currence in Eastern Australia, and seems to be a reliable indi-
cator of age. In Queensland, it is found abundantly in the
Walloon Series, but not in the Ipswich Series. In New South
Wales, it is found in the Clarence Series, Artesian Series, and
Talbragar Series, but not in any stage of the Hawkesbury Series.
There is no doubt of the identity of both the Clarence and Arte-
sian Series with the >A alloon Series, for they are continuous in
the tield. The Talbragar Series is unconformable on the Hawkes-
bury Sandstone, and the presence of Toiniopteris spatnlata in the
Talbragar Series indicates the relation of this Series to the
Walloon Series: the occurrence of a Taxites comparable with
7\ planus is additional evidence for assigning the two Series to
the same period.
The next point to be considered is the relation of the rocks of
the Hawkesbury Series to the other Lower Mesozoic rocks.
There is no doubt that the Hawkesbury Sandstone is older than
the Talbragar Series, and, therefore, older than the Walloon
Series. What then is the relation between the Ipswich Series
and the Hawkesbury Series ? In the following Table are in-
cluded the species of plants in the stages of the Hawkesbury
Series which are identical or closely comparable with species
in the Ipswich Series, the allied species in the Ipswich Series
being indicated in square brackets.
BY A. B. WALKOM.
87
Table ix.
Table showing species in the Hawkesbuiy Series identical or closelj^
comparable with species in the Ipswich Series.
Narrabeen Stage.
Thinnfeldia lancifoUa
T. odontopteroides
T. odoitfopteroides (Haw
kesbury type) [— T.
F< isfmauteU]
Cladophhiis cf. Royiti
Hawkesbury Sandstone
Stage.
Thinnfeldia odontopteroides^
[=7^. Feistniavfeli]
Oleaiidridinm [ Tfemopteris\
hntricaliforme
Macrotamiopteris [ T<vniop
teris] fvianamattce
Wianamatta Stage.
Phyllotheca an-sfra/is
Thinnfeldia odontopteroides
[— T. odonfopteroide,s and
T. Fei.stmanttfi]
T. lancifoUa
Macrof<eniopttris [ T<in iop-
teris] nnanamatta^
A lethopteris [ Cladoph Jtbifi]
austral is
Baiera mid t if da {B. Sim-
7no)idsi]
tPterophyJI inn[''.) sp.
It must be admitted that the flora of the Hawkesbury Series
has been but imperfectly determined, but what evidence theie
is, goes to show that the Ipswich flora is much more closely
related to that of the Wianamatta Stage than to either the
Hawkesbury Sandstone Stage or the Narrabeen Stage.
Having considered the evidence of the fossil plants, we may
now briefly deal with the evidence provided by the other fossils
in these Series.
The results so far published, of the examination of fossil
insects* from the Ipswich Series and Wianamatta Stage, do not
give promise of any special value from a stratigraphical point of
view, though, no doubt, they are of interest to the entomologist.
However, the fact that insects are found, more or less abund-
antly, in these two Series, and have not yet been discovered in
the other Series of Lower Mesozoic rocks, with tlie exception
of Cicada (?) loivei in the Talbragar Beds, is a point which must
be taken into consideration when discussing the correlation of
the Series. The insects in the Wianamatta Beds belong to the
same Orders as some of those in the Ipswich Series, aud, in one
case, the same genus is present, but there is no specific identity.
Queensland Geol. Surv., Publication No. 253.
S8 GEOLOGY OP LOWER MESOZOlC ROCKS OF QUEENSLAND,
One species of Estheria has been recorded from the Ipswich
Series {E. mamjaliensis Jones), and one from the Narrabeen
Stage {E. Coghhini Cox). The two species are different, and the
Ipswich species lias been identified by Etheridge* with E. tnan-
r/aliensif< from the Damuda Beds (Permian) of India.
Two species of Unio {U. ijJsviciensis and U. eyi'etisis) have
been described from the Ipswich Series; and two species of Unio
(U. wiaiiamatteiisis and U. Dunstani) together witli two species
of Unionella [U. bowralensis and U. Carnei) from the Wiana-
matta Stage. They are the only pelecypods recorded from the
Lower Mesozoic rocks of New South Wales and Queensland.
Fosvsil fish have been found somewhat abundantly on certain
horizons of the Havvkesbur\^ Series, and in the Talbragar Series:
and have been described by Dr. A. Smith Woodward.!
From the Talbragar Series, seven species were described, all
being new. + Regarding the age of this fish fauna, Woodward
concludes {op. cit., p. 26) " The Talbragar fish-fauna is, therefore,
probably not earlier than the Upper Lias, and may be referable
to the Lower Oolites."
The results of his examination of the fossil fishes from Gosford
and St. Peter's are remarkable. The Gosford horizon is about
the junction of the Narrabeen and Hawkesbury Sandstone
Stages, whereas the St. Peter's horizon is in the Wianamatta
Stage, and is some 900 to 1,000 feet above the Gosford horizon.
Regarding the fish-remains from Gosford, Wood\vard§ concludes:
'' So far as can be determined from the fishes, therefore, the
Hawkesbury beds may be regarded as homotaxial with the
Keuper of Europe, or, at latest, with the Rhsetic; and, on the
whole, the present writer is inclined to adopt the first of these
interpretations."
In the case of the St. Peter's specimens, the fish were collected
in two types of matrix, viz., an indurated shale or claystone, and
a grey mudstone, the former occurring as several bands sepa-
" (ieology and Palaeontology of Queensland, p. 397.
♦■ Mem. Geol. Surv. N. S. Wales, Palaeontology, Nos 4, 9, 10.
t IhicL, Palaeontology, No. 9.
§ Ibid., p. 55.
BY A. b. WALKOM.
89
rated by the latter. Woodward refers to the fish from the
indurated sliale as indicating a distinct Permo-Carboniferous
age, and those from the grey mudstone as Triassic or Rhajtic*
Obviously, great caution must be exercised in using these results
for the purpose of correlation.
(b) Victoria. — The Lower Mesozoic rocks of Victoria comprise
series of shales, felspathic sandstones, and mudstones, with
occasional conglomerates, which are developed in three areas,
viz., South Gippsland, Cape Otway, and Wannon River. Asso-
ciated with these rocks are coal-seams up to nine feet in thick-
ness. Analyses of the coals are of the same general type as
analyses of the Ipswich coals. The flora of some of these beds
has been described by Seward,! whose conclusion regarding their
age is, that they are approximately the same age as the Inferior
Oolite of England, or the Rajmahal Series of India. i Occa-
sional additions to the species described by Seward have been
recorded by Chapman.
The following list comprises the species described by both
Seward and Chapman : —
Marchantites sp.
j^Eqiiisetites wonthaggieiisis
Equisetites sp.
o.Lycopodiles victorice.
Adiantites lindsayoides.
Thianfeldia AlcCoyi.
Thinnfeldia sp.
HhizoynojHeris Etheridgei.
/:)Steiio2)teris eloiujata.
f^Steiiopteris sp.
aCoiiiopleviS hynieiiophylloides o.Gvukgo sp.
var. australica.
Sphenopteris ampla.
Sphenopteri.s sp.
T(Bniopteris Daintreei.
aT. Daiiitrep.i var. major.
[S7\ spaf.uhU'i var. crenafa.
o.Cladophlebis denficAilata var.
australis.
ftThinnfeJdia odontopter aides .
(i.Baiera australis.
a Ik gracilis.
Ginkgo (female flowers?).
(i.Nilssonia sp.
Podozamit'<s Barklyi.
P. ellipticus.
ftp. Kuisto'tii.
P. longij'oliu'i.
ihAraiicarites sp.A.
Mem. Geol. Surv. N. S. Wales, Palaeontology, No. 10, pp.
t Rec. Geol. Surv. Victoria, Vol. i., Pt.3, 1904.
t Op. cif., p. 184.
29.
90 GEOI.OGY OF LOWER MESOZOIC ROCKS OF QUEENSLAND,
Araucarites sp.B, (SPalissT/a australis.
ftConites sp. ftChelrolepis (1) setosiis.
cf. Brachyphyllum sp fi cf. Albertia austi'alis.
ftB. yippslandicum. CarpoJithes sp.A.
aVaxites sp. Carpolithes sp.B.
The species marked /i are recorded by Chapman, in addition
to those described by Seward.
Of the species in this list, those marked with a (10 in
number) are similar to types occurring in the Lower Oolite
of England.
In the Victorian Lower Mesozoic rocks, a tooth and scales of
Geratodus have been found, and also a claw of a dinosaur. One
species of Unio {U. stirlinyi) occurs in these beds.
Although the number of species of plants common to the Vic-
torian and Queensland Lower Mesozoic rocks is small, there is
no doubt that the Victorian flora resembles the Walloon flora
much more than it does the Ipswich flora. A few species, e.y.,
I'ceniopteris Daintreei { = 1\ spaiulata), T, Daintreei var. viajor,
and Cladophlehis deiiticnlata var. a^Lstralis ( = C. australis) are
identical with forms of the Walloon Series; while others, as
Sphenopteris ampla, Baiera yracilis, cf. Brachyphyllum sp., and
Taxites sp., are closely allied to forms found in the Walloon
Seiies. The occurrence of Tceuiopferis spatiilata and the conifers,
Araucarites, Brachyphyllum, and Taxites is sufficient to show
clearly, that this flora is not to be compared with that of the
Ipswich Series. There appears to be no doubt that the Victorian
Mesozoic Coal-Measures are to be correlated with the Walloon
Series in Queensland, and its equivalents in ISew-South Wales.
(c) South Australia. — In South Australia, the strata of Lower
Mesozoic Age comprise the south-western margin of the Great
Australian Artesian Basin, and an outlier of this preserved as a
faulted basin in the neighbourhood of Leigh's Creek. The
Leigh's Creek basin is quite isolated, and surrounded by rocks
of Upper Cambrian Age; the basin is 16 miles long, and has a
maximum breadth of 6 miles; the total thickness of strata in it
exceeds 2,000 feet of shales and carbonaceous shale, with bands
of limestone and sandstone, and a belt of very hydrous coal, 48
feet thick.
BY A. B. WALKOM. 91
Recent developments in Queensland sliow that large areas of
sandstone on the margin of the Artesian Basin, which have pre-
viously been regarded as of Cretaceous Age, must now be con-
sidered as equivalents of the Walloon Series. Future work may
show that the marginal portions of the Artesian Basin in South
Australia represent the same sandstones, though these are all at
present mapped as Cretaceous. Near the margins of the basin,
the water-bearing sandstones obviously approach close to the
surface, as shown from bore-records, and it seems more probable
that they outcrop, than that they are overlapped by the Cre-
taceous.
The fossil plants recoi'ded from the South Australian Lower
Mesozoic rocks are : —
Thiiwfeldia odontopteroides Alefhopteins sp.
[?=7'. Feistmaiiteil]. Equisetum 2 spp.
T media [? T. lancifolia\. Frenelopsif>{l).
MacroUt^niopteris iciaiiamatfxe. AtifJirophyopaisd) sp.ind.
TiHiiiopteris jiuctuans.
Fhyllojderis Feistmanteli occurs at Ooroowilanie Swamp, about
100 miles north of Leigh's Creek, but this may be in tlie Creta-
ceous rocks. This list has been drawn up by Howchin,* maiiily
from determinations by Etheridge. Unio eyretisis also occurs
abundantly in the same rocks.
This list of plants does not provide any very conclusive evi-
dence regarding correlation with other Australian strata. There
seems no reason for not considering the Leigh's Creek basin as
an outlier of the Artesian Series: and, further, the identity of
the Artesian Series with that of New South Wales and the
Walloon Series in Queensland is hardly open to question.
{d) Western A us ti-alia. ^V^ estevn Australia provides the only
Australian example of marine and freshwater fossils of Lower
Mesozoic Age occurring in association with one another. The
exact relation which the plant-beanng beds bear to those with
marine fossils is not clearly stated, but it may be inferred from
a study of Bulletins 36, 3y, and 50 of the Geological Sur^ ey of
Western Australia.
* British Assuc. Adv. of .Science : Handbook of South Australia, 1914, p.22o.
02 GEOLOGY OF LOWER MESOZOIC ROCKS OF QUEENSLAXO,
The marine fossils tix the age of the Lower Mesozoic of Western
Australia as Jurassic, and Etheridge refers those from the
Greenough River to the Oolite.* The Jurassic rocks cover a
large area in the Champion Bay district, near Geraldton, par-
ticularly between the Greenough and Irwin Rivers. According
to Gibb Maitland, they consist of "oolitic limestones, clays, sand-
stones (which are often ferruginous), grits, conglomerates, and
lignites."! Thev are horizontal or gentlv undulating, and their
thickness has been proved to the extent of 2,000 feet, by bores
in the Champion Bay district.
From these rocks, Arber^ has described Otozamites Feist-
mmiteli, and a coniferous fragment from near Minginew: and he
has pointed out the resemblance of these to fossils from lalgai,
Darling Downs, and Rosewood, west of Rockhampton in Queens-
land.
Examples of fossil plants in close association with marine
fossils are noted by Campbell. § Speaking of Mt. Hill.ji he says,
''Fossil stems of a plant, apparently a species of Equisetacese,
and Palm or Otozamites stem-scales and pieces of wood occur in
the sandstones and grits of the north side."
"A limestone band occurs at about 100 feet below the summit
of the hill, and can be well seen on the fence-line, 10 chains
west of the summit. Here, numerous Jurassic fossils occur in
an excellent state of preservation. On the ridge, also, extend-
ing north from the summit, there is a fossiliferous, fine-grained
sandstone."
The results of Dr. Arber's examination of the plants froin
Minginew were apparently available to Mr. Campbell, since he
quotes (p. 58) Arber's conclusions, so it may be presumed that
the plants from Mt. Hill are similar to those from Minginew.
On Plate 5 of the same work, in the vertical section of the Don-
* Geol. Surv. West Australia, Bulletin 36.
t Geol. Surv. West Australia, Bulletin 50, p. 21.
t Unci., Bulletin 36, p. 25.
%Ihid., Bulletin 38.
il Op. cit., p. 60.
BY A. B. WALKOM. 93
gara bore, "fossils" are indicated in shale at 535 feet, and
" plant- remains" at about 1,300 feet.
Although the information is scanty, it niay be concluded that
comparatively flat-bedded rocks of Jurassic Age, and of con-
siderable thickness, cover a large area in Western Australia;
that these beds contain both marine and plant-fossils, and that
the plant-fossils include equisetaceous stems, Otozamitps and
coniferous fragments, and also petrified, coniferous wood.
Where definite evidence for accurate correlation is scant}', as
in the present case, all possible indications must be considered:
and, at the same time, caution must be exercised in drawing
conclusions. In comparing these Western Australian Jurassic
rocks with the Lower Mesozoic strata of Queensland, the follow-
ing points stand out .— {a) In the Western Australian Jurassic,
the only definitely determined plant is Otozamites Feistmcmte/i,
which occurs in a fine-grained, ferruginous sandstone, possibly
of Oolite age; {h) in Queensland, fossil plants are abundant in
the Lower Mesozoic strata, but the genus Otozamites occurs only
in the Walloon Series, and usually in a fine-grained, ferruginous
sandst<jne, which is of very wide occurrence;* (c) the species
Otozamites Feistmanteli occurs both in the Western Australian
Jurassic and in the Queensland Walloon Series.
The indications, then, point to the probability that the AA'all-
oon Series is homotaxial with the Jurassic strata of Western
Australia. If, as the author believes, the development of the
fine-grained, ferruginous sandstone containing Otozamites Feist-
manteli indicates peculiar conditions under which deposition
took place, then there is the further indication that the two
Series are synchronous.
(e) 'Tasmania. — A rich. Lower Mesozoic flora is present in the
strata of that age in Tasmania. This flora, however, has been
untouched for many years, and the existing descriptions and
figures by Johnston are too imperfect to be used for any purpose
of definite correlation. In addition, Mr. Twelvetrees, Govern-
ment Geologist of Tasmania, kindly informed me that the rela-
* See above, p..')*).
94 GEOLOGY OF LOWEK MESOZOIC ROCKS OF QUEENSLAND,
tions of the Lower Mesozoic strata in Tasmania to one another
are not yet definitely settled. No attempt will be made here,
therefore, to correlate any of the Tasmanian Lower Mesozoic
rocks with the other Australian occurrences. One point mav be
noted. There appears to be an absence of Tceniopteris spatulata
in the Tasmanian strata, and this may indicate that they are
older than Jurassic.
Opinions have been expressed regarding the age of Tasmanian
strata. Chapman* suggests that it may be possible to divide
the " Upper Coal-Measures" of Twelvetrees into an older and a
younger series. He concludes, " The Jurassic flora of the Tas-
manian localities, Jerusalem, Fingal, Spring Hill, York Plain,
Hamilton, Richmond, New Farm, Sandfly, Recherche, South
Cape, Longford, etc., appears to contain an assemblage which,
with some few exceptions, as Sageiwj^teris and Phfjenicopsis, is
practically identical with that of the plant-bearing strata of
Jurassic age in South Gippsland, Cape Otway, and Western
Victoria "
Professor Davidf classes them provisionally "as Upper Trias
or Passage Beds into the Jurassic proper,"' whereas \V. S. Duni
regards them as equivalents of the Victorian Jurassic.
Mr. Twelvetrees points out that the evidence of the fossil
flora in Tasmania, as far as it goes, is slightly in favour of a
Rhsetic age. He also considers that some of the Tasmanian
occurrences are the equivalents of the Clarence and Talbragar
Series in New South Wales. §
Lower Mesozoic fossil plants are known from a number of
localities in New Zealand, but, in view of the fact that an ex-
amination of them has recently been carried out by Dr. E. A.
Newell Arber (the results not having been published yet||), it is
deemed inadvisable to discuss them at this stage.
* Records (xeol. Surv. Victoria, iii., Pt.2, 1912, p.223.
t British Assocn. Adv. Science, (4eoIoj^3' of the Commonwealth, p. 277.
t Ibid., p. 297.
^ (jreol. Surv. Tasmania, Bulletin 20, 191o, pp. 14-16.
I! Issued in February, 1918, too late for considei^ation in tiie present
paper,
BY A. B. WALKOM.
95
(f) Summary. - The results of these comparisons of the Queens-
land, Lower Mesozoic rocks with those of the other Australian
States, may be summarised in tabular form thus : —
■II
0) ij^
^ S ^ i ^^.'^
H^ X JS C ^ £
ew Soutl
Wales.
(V
o
.2 .§
Series
Talbraga
Beds
^
W 03
^W
.2^
5 O
96- geology of lower mesozoic rocks of queensland,
Geological History.
In discussing the conditions which existed during Lower
Mesozoic times in Queensland, reference must be made to the
conditions, as far as we know them, towards the close of the preced-
ing Permian ( Perrao-Carboniferous) Period. The latest sediments
of this latter period are the Upper Coal- Measures in both New
South Wales and Queensland. In New South Wales, the New-
castle or Upper Coal-Measures were formed in a large basin. In
an earlier paper,* I have attempted to show, in a general way,
the distribution of land and sea in New South Wales during the
various stages of the Permian (Permo-Carboniferous)t Period,
and have suggested that the successive stages within the period
might have been produced by a tangential force acting towards
the continental mass of Australia in a W. b}^ S. direction.
The close of the Palaeozoic era in Northern New South Wales
and Southern Queensland was accompanied by extensive intru-
sions of granitic masses. In the New England and Stanthorpe-
Warwick districts, these intrusions have resulted in extreme
folding of the Permian (Permo-Carboniferous) stiata, but have
not affected the sediments of the Walloon epoch. The age of
the intrusions is definitely between the upper marine sediments
of the Permian System and the Walloon Series. The Permian
(Permo-Carboniferous) sedimentary strata have been extremely
folded and contorted, and converted into slates resembling, in
general appearance, rocks of greater age than Permian: their
age, however, has been fixed beyond doubt by the occurrence in
them of marine fossils in the Drake district, and near Warwick. |
* Proc. Linn. Soe. N. S. Wales, 1913, xxxviii., pp. 139-145.
t It has recently been suggested bj^ Professor David and Mr. W. S. Dun
(British Assoen. Adv. 8ci., Australia, 1914: Report, p. 379; and Federal
Handbook on Australia, p. 267) that perhaps the term Permo-Carboniferous
in Australia should be replaced by the term Permian. Without going into
reasons here, it maybe stated that the author is entirely in accord with
the suggestion, and, for the purposes of this paper, proposes to write the
name thus : Permian (Permo-Carboniferous).
ij: Andrews, "Report on Drake drokl and Copper Field,'' Geol. Surv.
N. S. Wales, Mineral Resources, No. 12, 1908.
BY A. B. WALKOM. 97
Furtlier north, in the Gynipie district, rocks of similar a,ue are
also folded and metamorphosed to much the same extent as in
the districts just mentioned.
A word of caution must be expressed here regarding the so-
called "Gympie" strata. The results of field work carried out by
the officers of the Queensland Geological Sui vey tend to show
that many rocks have, in the past, been referred to the Gympie
Series without sufficient evidence; these are gradually being
sorted out by the Geological Survey, and put in their proper
position: and they have been found to include representatives of
the Carboniferous and Devonian Systems, in additi<m to true
Permian (Permo-Carboniferous) strata. Care must be exercised,
therefore, when dealing witli rocks classed as Gympie.
In the Gympie district, where undoubtedly Permian (Permo-
Carboniferous) sedimentaries are much altered, there are
granitic, intrusive masses whose age is not directly determin-
able. It seems a reasonable suggestion that they are of similar
age to the New England intrusions, and that they have been
responsible (at least in part) for the folded and altered condition
of the Gympie rocks.
Further north and north-west in Queensland, the Permian
(Permo-Carboniferous) rocks [Bowen River coalfield, Dawson-
AJackenzie coalfield, etc.] appear to be normal, and not to have
suffered any considerable folding as the result of the intrusion of
large plutonic masses.
In the distribution of the Permian (Permo-Carboniferous)
rocks of New South Wales and Queensland, then, there are to
be noted the following important points. Sediments of this age
are known from Southern New South Wales to Northern
Queensland; normally, these sediments are unaltered, and ap-
parently have not been subject to extreme folding-forces; in the
central part, however, (from New England to the Gympie dis-
trict) the intrusion of extensive, granitic masses has resulted in
extreme folding and metamorphism of sediments of this age,
which, as a result, resemble strata of much greater age than the
normal Permian (Permo-Carboniferous; strata of Eastern Aus-
tralia.
98 GEOLOGY OF LOWER MESOZOIC KOCKS OF QUEENSLAND,
These extensive intrusions probably resulted in the elevation
of the areas affected, to a much greater altitude than the neigh-
bouring areas.
The Lower Mesozoic rocks of Eastern Australia appear to have
been deposited in a series of basins, which were not all developed
at the one time. The sediments deposited in these basins are,
with a single exception {supra, p. 38), freshwater accumula-
tions, and this fact tends to show that the basins must have had
some outlet; otherwise, the bodies of water would comparatively
soon have become salt. Regarding deposits formed in basins in
this manner, we may note that Suess quotes Richthofen's dis-
tinction between central and peripheral types of drainage as
illustrated by salt and coal. He says, " Salt with gypsum cor-
responds to a closed drainage system or central position; coal,
when found in extensive freshwater basins, corresponds to the
open outflow, without which no accumulation of water could
maintain for long its original composition.""^ The latter part of
this statement seems to admit of no contradiction, and it applies
to the basins in which the sediments of Lower Mesozoic age in
Eastern Australia were deposited. We shall consider later the
positions of the outlets for the various basins.
The earliest basin developed was that in which the Hawkes-
bury Series of New South Wales was deposited. In the central
portion of this basin there appears to be stratigraphic conformity
between the Upper Coal- Measures, and the Narrabeen Stage of
the Hawkesbury Series. Nearer the margins, however, there is
unconformity. In addition, there is a very distinct palaeonto-
logical break, only a few forms persisting from the lower to the
upper. The unconformity between these two Series is one which
involves a detinite interval of time, but not any considerable
movement. The time-interval must have been sufficiently long
for the Mesozoic flora of the Narrabeen Stage to have almost
completely replaced the typical Palaeozoic flora of the Upper
Coal-Measures
The basin in which the Narrabeen Stage was deposited was
produced by a gradual subsidence, which apparently continued
* Suess, "The Face of the Earth,'" iii., p. 312.
BY A. B. WALKOM. 99
throughout a long period during which the Narrabeen and
Hawkesbury Sandstone Stages were deposited. The area of
deposition increased gradually during this time, the Hawkesbury
Sandstone Stage covering a much greater area than the Narra-
been Stage. During the early stages, volcanic activity appears
to have been in evidence, as indicated by the development of
tufFaceous rocks in the Karrabeen Stage. The thickness of the
sediments deposited in this basin reaches a maximum of about
3,000 feet, made up of a maximum of 2,000 feet of the Narra-
been Stage, and 1,000 feet of the Hawkesbury Sandstone Stage.
The Hawkesbury Sandstone Stage is apparently conformable
with the Narrabeen Stage. The existence of breaks, which do
not show as very marked unconformities, must, however, always
be borne in mind. Writing on this point, Schuchert remarks,*
" The easily seen, marked unconformities are of course
accepted at full face-value; but the many more apparently con-
formable and yet broken contacts, the disconformities, are gener-
ally overlooked, or when seen are generally undervalued. ... In
regard to the breaks, the statement can be made that there are at
least ten disconformities for every known angular unconformity."
These statements may be applied to both the Permian (Permo-
Carboniferous) and Mesozoic Systems of Australia, within which,
apparent conformity appears to be general, but where there are
quite a number of palseontological breaks.
During the time in which the Narrabeen and Hawkesbury
Sandstone Stages were deposited, probably the rest of Australia,
which at that time was of greater extent than to-day, was dry
land.
Between the Hawkesbury Sandstone Stage and the Wiana-
matta Stage there is a distinct break. The Wianamatta beds
have been deposited on the eroded surface of the Hawkesbury
Sandstone, which must, therefore, have been elevated, exposed
to denudation, and again submerged before the succeeding beds
were laid down. In addition, the extent of the Wianamatta
beds was much more limited than the Hawkesbury Sandstone.
* Bulletin Geol. Soc. America, xxvii, 1916, p.497.
100 GEOLOGY OF LOWER MESOZOIC ROCKS OF QUEENSLAND,
Our studies of the fossil floras indicate that the Ipswich Series
in Queensland was laid down at the same time as the Wiana-
matta Stage in New South Wales. The strata of this Series
were laid down over a comparatively restricted area in Southern
Queensland, though it is not possible to obtain an accurate idea
of its exact extent. Parts of the areas bordering the basin in
which the Ipswich Series was deposited were probably very
elevated, e.g., the Stanthorpe-Warwick, and the Gynipie Dis-
tricts, the extent of the elevation having been due to the exten-
sive intrusions of granite at the close of the Palaiozoic era.
It is probable, also, that part of the Lower Mesozoic sediments
of Tasmania was deposited contemporaneously with the Wiana-
matta Stage in New South Wales, and the Ipswich Series in
Queensland.
The Bundamba Series, which follows the Ipswich Series in
Queensland, has no equivalents, so far as we know, in Kew South
Wales ; apparently, sedimentation in the main Hawkesbury
Basin came to a close with the end of the Wianamatta Stage.
Later sediments in that State were laid down to the north and
north west.
In Queensland, the dominant movement from the beginning
of the Ipswich epoch was a subsidence. This movement was not
continuous; there were, at least, breaks represented by the base
of the Bundamba Series and of the Walloon Series, and there
may have been further breaks within the Series. At the time
of these breaks there was probably a cessation of sedimentation,
accompanied by a slight uplift before the downward movement
reasserted itself; the predominance of the downward movement
is proved by the ever-increasing area covered by the Lower
Mesozoic sediments until, towards the close of the Walloon
epoch, the greater part of Queensland was probably covered by
enormous stretches of fresh water. The portions of Queensland
which were not submerged during the Walloon epoch (Jurassic)
include the north-western corner of the State, an area in the
neighbourhood of Hungerford on the New South Wales border,
and probably areas to the east of the present Main Divide, which
had been considerably elevated as a result of movements accom-
BY A. B. WALKOM. lOl
panying the intrusion of extensive granitic masses at the close
of the Palaeozoic era. It is impossible to define these latter
areas at present, but they include the Stanthorpe and Gympie
districts, and probably some of the present coastal portions of
Queensland north of Rockhampton.
The area over which deposition of the Walloon Series took
place extended into New South Wales and South Australia; and
its boundary probably agreed fairly closely with the margin of
the Great Australian Artesian Basin; it is from sandstones in
the Walloon Series and its equivalents, that the great bulk of
the artesian water is obtained, so that, although these are not
known to outcrop along the southern margin of the Artesian
Basin, their presence is known, and they are apparently over-
lapped by the J^ower Cretaceous strata.
It may be remarked here, that it is only comparatively recent
work that has shown the sandstones forming the intake in
Queensland to be of Walloon age. In view of this, it is possible
that outcrops on the southern and western margins of the Basin
may, in the future, be shown to be of the same age, though at
present mapped as Cretaceous.
The Clarence Series represents deposition in the south-eastern
portion of the same basin. The Talbragar beds seem to repre-
sent depo.sition in a small, isolated lake.
In South Australia, the small, faulted basin at Leigh's Creek
probably represents an outlier of the main occurrence further
north.
Contemporaneously with the Walloon Series also, there were
deposited the Jurassic sandstones, etc., of South Gippsland, Cape
Otway, and the Wannon River areas in Victoria, and possibly
also portion of the Lower Mesozoic strata of Tasmania. At the
same time, marine sediments were laid down in Western Aus
tralia, and, associated with these, there are a few plant-remains.
The close of the Jurassic in Australia was not marked by any
violent movement. In Queensland, New South Wales, and
South Australia, the Jurassic rocks are followed by Cretaceous
marine strata (Rolling Downs Series) without any observable
unconformity. Palseontologically, however, there is a very
102 GEOLOGY OF LOWER MESOZOIC ROCKS OF QUEENSLAND,
marked break, and there is also generally a lithological distinc-
tion. The Cretaceous sea represents a transgression from the
north, and covered the greater portion of the Jurassic strata
over what is now the Great Artesian Basin, and, as shown in
Text-fig. 6, this transgression probably divided the Australian
continent in two. The present Main Divide in Queensland was
probably elevated, and became a land-area practically throughout
its whole length in Cretaceous time, and has remained essentially
so to the present day. On the west, the Cretaceous sea covered
the area of the Artesian Basin as already mentioned, while, on
the east, there are Cretaceous marine deposits between Mary-
borough and Bundaberg reaching a thickness of 1,000 feet. It
is impossible to determine the exact extent of these to the east.
The marine Cretaceous, both in western Queensland and in the
Maryborough district, are followed by Cretaceous freshwater
deposits known as the Winton Series and Burrum Series,
respectively.
Lower Mesozoic Pal^eogeography of the Australasian
Region.
Published maps indicating the geography of the Australasian
region durins; Mesozoic times are fevv in number.
Perhaps the oldest is that of Neumayr, which has been con-
tinually quoted and figured by later writers, including Jensen"^
and Hedley.f The latter "| also prepared a map to show the
Queensland coast at the close of the Mesozoic era. Jensen^ has
published a series of maps showing the distribution of land and
sea in the Australian region at various periods, including one in
Triassic time. Recently, Schuchert|| has published a series of
palseogeographic maps of Oceania, including one in Triassic time,
and one in Cretaceous.
The important general feature of these maps is that they show
" Proc. Linn. Soc. N. S. Wales, 1908, xxxiii., p.507.
t Report Aust. Assoc. Adv. Science, xii., 1909, p. 332.
t Op. ciL, p. 333.
§ Proc. Roy. Soc. Queensland, xxiii., Pt.2, 1912.
II Amer. Journ. Science, xlii., 1916, p. 99.
BY A. B. WALKOM. 103
a coast-line not a great distance east of the present position of
the eastern coast of Australia. This coast-line is the western
shore of a long gulf running from the south in a general meri-
dional direction between Australia and New Zealand. Neumayr
called this the Gulf of Queensland. Schuchert's map shows the
extension of this gulf further north to about the latitude of New
Caledonia, when it turns to the east and again joins the ocean.
Evidence as to the existence, or otherwise, of this gulf is scanty.
Hedley* says, "According to Neumayr, a meridional crease in
the earth's crust produced, in Jurassic times, a gulf, which he
called the Gulf of Queensland, whose western shore transgressed
the present east Australian coast." The most important piece
of evidence indicating the existence of this gulf is the presence
of an impoverished fauna of Foraminifera and Ostracoda in the
upper beds of the Wianamatta Stage in New South Wales {supra,
p. 38). This is positive evidence of marine or estuarine condi-
tions in the Wianamatta Basin (probably during late Triassic
time). There is no evidence at all to show that this gulf trans-
gressed the present east coast of Australia during Jurassic time;
it is certain that there are no evidences of Jurassic marine
deposits in Eastern Australia.
The gulf was probably more or less coincident with the present
position of the Thomson Trough; but whether the Thomson
Trough is as old as Lower Mesozoic, is difficult to determine.
The palseogeography of the Australasian region involves a
consideration of the structure of the south-western Pacific resrion.
structural studies of this region have been made by a number of
geologists, including Dana, Suess, Gregory, Marshall, and
Schuchert.
Marshall has pointed out that some of the earlier studies were
based mainly on the geographic distribution of the island-chains,
without much knowledge of structure. He has very rightly con-
tended that conclusions drawn from such distribution mav be
quite erroneous.
Marshall has argued that " the real boundary of the south-
west Pacific passes through New Zealand, Kermadec, Tonga,
* Report Aust. Assoc. Adv. Science, xii., p. 331.
104 GEOLOGY OF LOWER MESOZOIC ROCKS OP QUEENSLAND,
Fiji, New Hebrides, Solomon, and on to the Admiralty Islands."*
This line would then be the eastern boundary of the continental
mass of which Australia, New Caledonia, New Zealand, etc., are
remnants. Within this continental region there are a number
of great deeps, e.g., the two Solomon Island deeps (the northern
one named the Planet Deep, the other unnamed) and an unnamed
deep between the New Hel>rides and New Caledonia. A discus-
sion, which the author had recently with Rev. Father Pigot, of
Piverview College, produced some facts which seem directly to
concern the question of these deeps. The seismometers at the
Riverview College observatory have, within the past few years,
recorded a number of earthquake-shocks, and Father Pigot has
calculated the positions of origin of these shocks. He kindly
informed me that quite a large percentage of the shocks recorded
had their origin along a line from Kermadec, through the deep
between the New Hebrides and New Caledonia, the Planet Deep,
and the Swire Deep (east of the Phillipines). The prevalence of
earthquake- shocks emanating from this line of deeps is probably
connected with faulting-movements, and it may reasonably be
argued that these movements are probably a continuation of
those which produced the deeps, and, therefore, that these deeps
are of very recent origin. If this is so, we have, in these earth-
quake-records, evidence wiiich supports Marshall's contentfon as
to the true margin of the south-west Pacific. Marshall's con-
clusions in this respect seem reasonable; and Text-fig. 2 (p. 105)
indicates the probable limits of the continental mass about the
beginning of Mesozoic time; this map has been drawn-up mainly
from the works of Marshall and Schuchert, already quoted.
This continental mass was connected with Asia about tlie
begitjning of Mesozoic time, and it may have been continuous
witli Gondwanaland. The breaking down of Gondwanaland
commenced in early Mesozoic time, and the permanent enlarge-
ment of the Pacific basin probably commenced about the same
time. There are marine Triassic rocks both in New Caledonia
and New Zealand —evidences of epicontinental seas— but there
is no marine deposit of Triassic age known on the present Aus-
* Report Aust. Assoc. Adv. Science, xiii., 1911, p. 99.
BY A. B. AVALKOM.
105
tialiaij CoMtiiient. The distribution of land and sea during
Triassic time in this region appears to me to be as indicated in
Text-ti^'.2.
Map showinir the ai)pruxiiiiate limits uf the Continental Mass, of which
Australia, New Zealand, New Caledonia. &c., are remnants.
Text-fig.3(p.l07). Speaking of the Triassic marine deposits of New
Caledonia and New Zealand, Marshall"^ expressed the opinion,
• (;eolu>;y of New Zealand, 11)12, p. 18-j.
106 GEOLOGY OF LOWER MESOZOIC ROCKS OF QUEENSLAND,
that it cannot be definitely decided whether they represent
coastal deposits on an eastern or western coast. If the boundary
of the true Pacific basin on the south-west be the line stated, it
seems more probable that they represent deposits along the
eastern coast of a continent.
The evidence for the existence of the gulf, which Neumayr
called the Gulf of Queensland, is not very definite.
The continued deposition of freshwater sediments in the
Hawkesbury basin, indicates a continuance of freshwater condi-
tions in this basin over a long period, and there must have been
some outlet, as noted above (see p. 98). As far as we know, there
is no possibility of an outlet to the north, south, or west, and, in
view of the jDresence of a foraminiferal and ostracodan fauna in
the upper portion of the Wianamatta Stage, it seems probable
that the outlet connected with an arm of the sea, such as the
Gulf of Queensland. The exact extent of the gulf is purely
conjectural.
During the time when the Wianamatta Stage in New South
Wales, and the Ipswich Series in Queensland were being de-
posited, the Gulf of Queensland probably had its maximum ex-
tension in a northerly direction. At this time, there were two
small basins in which the Wianamatta Beds and the Ipswicli
Series respectively were deposited. Both these basins may have
had outlet to the Gulf of Queensland. Text-fig. 4 (p. 109)
indicates a probable distribution of land and water at this
time.
In Jurassic time, there is no clear evidence to show the exist-
ence of the Gulf of Queensland. There are marine Jurassic
rocks in New Caledonia. In New Zealand, in the south of the
South Island, there are freshwater and marine strata of Jurassic
age associated with one another, indicating probably oscillation
of marine and fluviatile deposition in a coastal region. In the
North Island, in the Kawhia-Waikato district, there are marine
Jurassic rocks, and there are also records of fossil plants (Jurassic)
from the same district. This, again, indicates a probable coastal
region.
BY A. B. WALKOM
107
Text-fig. 3.
Map showing the distribution of land and water in the Australasian
Region in early and middle Triassic time. (Solid black represents
the present known extent of sti'ata).
108 GEOLOGY OF LOWER MESOZOIC ROCKS OF QUEENSLAND,
The Jurassic rocks of Victoria and Tasmania are freshwater
deposits, and probably extended some distance east of the present
coast. The Jurassic strata of New^ South Wales and Queensland
extend right to the extreme north of the latter State, and there
are marine Jurassic strata in New Guinea (Strickland River).
This seems to indicate that these Jurassic rocks were laid down
in a large basin, which had some outlet to the north. Another
point which strengthens this, and is against the existence of the
Gulf of Queensland during the Jurassic period, is the enormous
amount of sediment represented by the continuous Jurassic strata
of New South Wales and Queensland. These beds must average
some thousands of feet in thickness over an area of some hundreds
of thousand square miles, and must represent denudation of a
large area. That the drainage to this basin was limited on the
north, west, and south, is easily shown, for, in these directions,
we know the regions of synchronous deposition, and the divides
separating these from the Walloon basin do not allow of a very
extensive area from which the sediments may have been derived.
There is left, then, only extension to the south-east and east to
any very large extent, and, if this was the case, there seems little
possibility of the existence of the Gulf of Queensland.
These considerations have governed the drawing up of Text,
fig. 5 (p. Ill) showing the distribution of land and sea during
Jurassic time.
The Thomson Trough, which lies to the east of Australia, may
have been of comparatively recent origin. The east coast of
Australia has been subject to folding since Lower Cretaceous,
and to considerable faulting during Cainozoic; and it does not
seem improbable that the Thomson Trough is a result of these
movements. Schuchert* concludes his study of continental
fracturing and diastrophism in Oceania thus: "To sum up, we
may say that the bottom of the Pacific Ocean in the region of
greater Australasia seemingly became more and more mobile
with the Lower Carboniferous and especially during the Jurassic
and Cretaceous. During this very long time, the eastern half of
* Amer. Journ. of Science, xlii., 1916, p. 104.
BY A. B. WALKOM.
109
Text-fig. 4.
Map showing the distribution of land and water in the Australasian
Region during Upper Triassic time (?Keuper or Rhaitie). (Solid
black represents the present known extent of strata).
110 GEOLOGY OF LOWER MESOZOIC ROCKS OF QUEENSLAND,
the Australian Continent, a land about 1,800 miles east and
west and 2,200 miles north and south, was folded into a series
of parallel ridges trending north-west and south-east, nearly all
of which went down more and more beneath the level of the sea
to a maximum depth of about four miles and an average depth
of between one and two and a half miles. Small parts of the
ridges still protrude above the ocean (at least New Caledonia),
but most of what we see are the volcanoes that have built them-
selves up above the folded rocks to the level of the sea."
This statement gives a general outline of the geological history'
of this region, but the stress laid on the Jurassic as a period of
folding does not appear to be justified. As far as our present
knowledge goes, the periods during which folding-forces have
been effective since the Carboniferous are Permian (Permo-Car-
boniferous), late Cretaceous, and possibly late Cainozoic. The
Permian strata of New South Wales indicate folding with axes
approximately N. by W.; the Triassic, Jurassic, and Lower Cre-
taceous all show a general stratigraphical conformity, and have
all been affected to the same extent by a folding which was later
than Lower Cretaceous and earlier than Cainozoic, the direction
of the axes of folding being about N.30°W.; the Cainozoic rocks
in the Ipswich District have been subject to a minor folding-
movement with approximately meridional axes.
It is also noted that, in New Caledonia, Triassic, Jurassic, and
Cretaceous strata are apparently conformable, and have all been
subject to subsequent overthrusting from the north-east. This
strengthens the conclusion that the more important folding did.
not take place till after the deposition of the Cretaceous.
It appears, then, that there were transgressions in Triassic
time (1) from the east, extending as far west as New Caledonia,
and (2) from the south, forming the Gulf of Queensland. During
Jurassic, the eastern coast remained in much the same position,
the Gulf of Queensland disappeared or was very much reduced,
and the sea transgressed in Western Australia parallel to the
present western coast, and also in New Guinea to the Strickland
River district; during this time, also, the extent of the basins.
BY A B. WALKOM.
Ill
i
Text-fig. 5.
Map showing the distribution of land and water in the Australasian
Region in Jurassic time. (Solid black represents the present known
Regie
extent of strata).
112 GEOLOGY OF LOWER MESOZOIC HOCKS OF QUEENSLAND,
in which freshwater deposits were laid down, increased enor-
mously. The early Cretaceous period saw very great changes in
distribution of land and ^vater. The sea transgressed from the
north in the region of the Gulf of Carpentaria, at least as far as
Lake Eyre; Australia appears to have been divided into two
parts, either by the extension of this transgression to the south-
west, or by meeting another from the south-west. These suc-
cessive changes are shown on Text-figs. 3-6.
Summary.
The Lower Mesozoic rocks of Queensland comprise three di^■i-
sions, namely, the Ipswich, Bundamba, and Walloon Series, The
Ipswich and Bundamba Series are of comparatively limited dis-
tribution, and are confined to the south-eastern portion of the
State. The Walloon Series has a much greater extent; in addi-
tion to occurring in South-eastern Queensland, in association with
the Ipswich and Bundamba Series, it outcrops in a belt along
the western slope of the Main Divide from the New South Wales
border to Cape York, dipping westerly beneath the marine Cre-
taceous. It probably underlies the Cretaceous strata over the
greater part of Western Queensland. In Eastern Queensland,
there are a number of small, isolated occurrences of the Walloon
Series, as indicated on Plate i. The thicknesses of the three series
are approximately as follows: Ipswich Series, 2,000-2,500 feet;
Bundamba Series, 3,000-5,000 feet; and Walloon Series, up to
10,000 feet.
From an economic point of view, the Lower Mesozoic rocks
are of special importance in three respects, namely, (1) the greater
part of the coal produced in Queensland is from the coal-measures
of the Ipswich and Walloon Series, chiefly the former; (2\ the
large supplies of artesian water are obtained from sandstones
which are the equivalents of the Walloon Series in Western
Queensland; (3) practically all the Queensland sandstone used
for building purposes is of Lower Mesozoic age.
In South-eastern Queensland, the Lower Mesozoic rocks (and
also the Cretaceous strata) have been folded into a series of anti-
IJY A. li. WALKOM.
113
Text-fig. 6.
Map showing the distribution of land and water in the Australasian Region
in Lower Cretaceous time. (Solid black represents the present known
extent of strata).
114 GEOLOGY OF LOWER MESOZOIC ROCKS OF QUEENSLAND,
clines and synclines, whose axes are in a general N.30°W.-S.30°E.
direction. This folding is sometimes severe, the strata having
been fractured, the folding giving place to distinct faulting. The
age of this folding and faulting is Post-Lower Cretaceous and
Pre-Cainozoic ; none of the Cainozoic rocks in south-eastern
Queensland have been affected by it.
In the Ipswich district, Mr Cameron has observed a sub-
sequent folding-movement with approximately meridional axes;
this latter folding has affected the Cainozoic sedimentary and
volcanic rocks, as well as the Lower Mesozoic rocks.
In Western Queensland, the Mesozoic rocks have not been
subject to folding-movements.
The Bundamba Series is practically devoid of fossils, but both
the Ipswich and Walloon Series contain abundant plant-remains;
there are also numerous fossil insects in the Ipswich Series.
Examination of the fossil floras indicates clearly that the Ipswich
Series must be regarded as Triassic in age, and the Walloon
Series as Jurassic. The exact position of the Ipswich Series in
the Triassic cannot be definitely fixed at present, but it seems
certain that it is Upper Triassic, possibly Rhsetic, but probably
older. The flora of the Walloon Series corresponds, to almost
equal extents, with Liassic and Lower Oolitic floras. This Series
probably corresponds to a period covering these two epochs, and
when the Walloon Series is known in greater detail, it may be
possible to subdivide it accurately.
The exact position of the Bundamba Series in the Geological
Record cannot be fixed, but it is suggested that it is more closely
associated with the Ipswich Series than with the Walloon Series,
and, therefore, it is tentatively placed in the Triassic.
A comparison of the Queensland Lower Mesozoic strata with
other occurrences in Australia of similar age seems to show, (1)
that the Narrabeen and Hawkesbury Sandstone Stages in New
South Wales are older than the Ipswich Series; (2) that the
Wianamatta Stage of the Hawkesbury Series in New South
Wales, and also possibly part of the Lower Mesozoic strata of
Tasmania are of the same age as the Ipswich Series; and (3) that
BY A. B. WALKOM. 115
the following series in the other States are of the same age as
the Walloon Series: the Artesian Series, Clarence Series, and
Talbragar Beds in New South Wales; the Jurassic strata of the
South Gippsland, Cape Otway, and Wannon areas of Victoria;
the Leigh's Creek beds in South Australia; part of the Lower
Mesozoic strata of Tasmania; and the marine Jurassic series in
Western Australia.
An account has been given of the probable geological history
of Eastern Australia between the close of the Palaeozoic era and
the beginning of the Cretaceous period, and it is evident that the
Lower Mesozoic rocks are, with a single exception, of freshwater
origin. Their exact mode of origin is not definitely settled; it
has been shown that the basins in which they were deposited
must have had some outlet; deposition may tlien have been in
inland lake-basins with a river-outlet, or the deposits may repre-
sent accumulations under fluviatile conditions.
The geological history of the area is illustrated by a series of
palseogeographic maps of Australia and the South-western Pacific
region, representing the probable distribution of land and water
at different stages in Lower Mesozoic time.
EXPLANATION OF PLATES I.-IL
Plate i.
Map of Queensland showing the distribution of Lower Mesozoic rocks.
Plate ii.
Geological Sketch-Map of South-eastern Queensland showing the areas
occupied by the Lower Mesozoic rocks.
116
STUDIES IN AUSTRALIAN KEUROPTERA.
No. 5. The Structure of the Cubitus ix the Wings of the
Myrmeleonti d^e.
By R. J. TiLLYARD, M.A., D.Sc, F.L.S., F.E.S., Linnean
Macleay Fellow of the Society in Zoology.
(With three Text-figures).
In No.l of this series of Studies,* I worked out the wing-
venation of 2Iyrineleon unise7'iatus Gerst., from the pupal wing-
tracheation, and showed how, in the forewing, the apparently
simple vein M was in reality a forked vein, the vein usually con-
sidered to be Cuj being in reality the cuhito-median, Cu^ -1- M.,, and
the point of incidence of M2 on to the line of Cuj being marked
by the foot of the oblique vein O. The branch descending from
the point of bifurcation, which I called the cubital fork (cuf),
was, naturally enough, taken to be Cuo. It was on the lack of
specialisation of this branch, that I based my argument for the
recognition of the tribe Protoplectrini, of which the correct name
should be Creagrini, as previously given by Navas.
In No. 4 of this same series of Studies,! when describing the
wing- venation of the Hemerohiidce, I showed how, in the fore-
wings of Drejmnepteryx and Drepanacra, an archaic Cug was
present close to the base, and that, in addition to this, Cu^ was
strongly branched at a sharp angle, much further distad, into
two veins which I called Cuj^ and Cuu, In the allied genus
Drepanomina, the archaic Cuo is lost, and replaced by a furrow.
In the majority of the smaller Hemerobiidce, even the furrow has
disappeared. Thus, in the most highly specialised forms of the
family Hemerohiidce, all traces of the original C\x.^ are lost, and
'' These Proceedings, 1915, xl., Pt.4, pp. 734-751, PL Iviii.
\0p. cit., 1916, xli., Pt.2, pp.269-332, Pis. xii.-xix.
BY R. J. TILLYARD. 117
there is left only the distal secondary forking of Cui into Cuj^ and
Cuiij, the point of bifurcation being properly termed the i^pcondary
cubital fork {ciif).
Now the resemblance between the condition of the cubitus in
the forewings of these smaller specialised Hemprohiidce^ and in
the forewings of the Myrmelcontid(f, is too striking to be ignored.
It suggested to me at once that the veins which I had named
Cuj and Cu.2 in the Myrmflpontidcf were in reality only Cuj^ and
Cu]b respectively, and that, if the true Cu" still existed, it must
be sought for near the extreme base of the wing. 1 therefore
examined a considerable number of MyvmpUontidw^ and found
that, in many cases, there could be detected, close to the base of
the wing, a semi-opaque connecting-vein, not unlike an oblique
vein, (though not so well defined) running across from Cu^ to 1 A.
This appeared to me to be the remains of Cu.^. With a view to
proving this point, I decided to examine the pupal tiacheation
of some of the older Mynndpontidce, belonging to the suV)family
Dendroleontince, in the imagines of which this supposed remnant
of Cu.2 appeared to be much more evident than in the genus
Myrmeleon itself.
Most of the larva" of the Dcndroleontince do not make pit-falls,
but must be sought for either in loose sand, or in rubble or debris
collected at the bases of large trees. During the winter of 1916,
larvte of several genera of this type were collected around Sydney.
A number of these spun up in the following September. At the
end of a fortnight, three likely-looking cocoons were opened, but
were found still to contain larvae. They were then placed under
close observation until pupation took place. The first pupa ex-
amined proved to be very recently formed; so that, although a
good dissection of the forewing was obtained, the structure was
so exceedingly delicate that all the air passed out of the tracheae
before either a photomicrograph or a camera-lucida drawing could
be obtained. With the second dissection, I decided to take no
risks, but removed it at once to the microscope-stage, and made
a camera-lucida drawing of it. Both the pupal tracheation and
the imaginal venation showed up very clearly, and the result was
9
118
STUDIES IN AUSTRALIAN NEUROPTERA, V.,
X / //// / / / /ZZZZZ^^:^^ So
Text-fig. 1.
Traeheation of forewing in the pupa of Xanthohon hefinsi Tillj^ard; ( x 35).
T-T / I M// / /////,
Sc
R
Rs
Ml
Text-fig. 2.
Venation of Xanthohon helmsi Tillyard; a, basal half of forewing, ( x 4^);
h, basal portion of same, enlarged, ( x 10); c, base of hindwing, ( x 4iK
LETTERING OF TEXT-FIGURES.
lA, 2A, 3A, first, second, and third analis, respectively — lA-f Cus,
eubito-anal — C, costa — Cu, cubitus; Cui, Cuo, its two principal branches;
Cuu, Cuib, the two principal branches of Cui — Cuia-f-Mo, cubito-median
— cuf, primary, and cnf, secondary cubital fork— ^r, vestigial frenulum
— M, media; Mi, M„, its two principal branches — mf, median fork — Oc,
cubital oblique vein — Om, median oblique vein — R, radius — Rs. radial
sector — 7'/, radial fork — Sc, subcosta.
BY R. J. TILLYARD. 119
a complete success, which is here reproduced in Text-fig. 1. As
the venation was complete, and even the delicate pigmentation of
the imaginal wing could be noted, I was easily able to determine
this wing as belonging to the rather rare insect Xantholeon
hdmsi Tillyard, — a result as fortunate as it was unexpected, since
Xantholeon is certainly one of the most archaic of our Australian
Myrnidpontid(e.
The result of this examination can be gathered by comparing
Text-figs. 1 and 2. It will be seen that it not only bears out in
full my suspicions concerning the real position of Cuo, but shows
also a further unexpected point of interest. For the trachea
Cu2 is not a weak remnant descending to 1 A, as the examination
of the imaginal venation had led me to suspect, but is a strong
trachea which has captured the line of lA, in the same manner
that M2 has captured the line of Cui,,. In both cases, it is the
upper trachea that prevails, the lower that goes under. In both
cases, the crossing of the upper trachea on to the level of the
lower is marked, in the imaginal venation, by an oblique vei7i,
which represents, of course, the actual basal piece of the upper
vein in question. We must therefore distinguish in future
between the median oblique vein {Om) and the cubital oblique
vein (Oc). This latter is very clearly marked in Xantholeon
(Text-fig. 2), also in Glenoleon, Acanthaclisis, and other genera;
but it cannot be satisfactorily made out in our Australian species
of the genus Myrmeleo7i, which are more highly specialised.
The process by which M2 and Cug have attained their present
positions may be described as a process of "trachea-capture."
The two tracheae run at first more or less parallel. As the wing-
becomes narrower, they come to lie side by side. The oxygen-
supply conveyed by both is now no longer fully needed; so that,
in course of time, the weaker (in this case the lower) trachea
becomes aborted, being reduced to a remnant impinging upon
the strong upper trachea from below. In the resulting imaginal
wing-venation, however, where the determining factor is not
oxygen-supply, but the wing-stresses brought about during for-
ward flight, the vein formed along the coui'se of the lower trachea
120
STUDIES IN AUSTRALIAN NEUROPTERA, V.,
continues the line of the stronger upper trachea beyond the point
where the partially aborted lower trachea ceases to exist, and
must, therefore, be regarded as a composite vein, whose double
origin is now only indicated by the oblique vein placed above it.
Thus, in the case under consideration, we have two composite
veins formed, viz., the aubito-median, Cuj^ + M.,, (whose structure
I worked out in No.l of this series of Studies), and the cnbito-anal,
1 A + Cug, whose existence
has not been before sus-
pected. The oblique vein
indicating the presence of
the former is Om, that in-
dicating the presence of
the latter is Oc. The com-
plete, correct, amended
notation for the forewing
of Xantholeon helmsi Till-
yard, is given in Text-fig.
2. In Text-fig. 3, I oflPer
a phylogenetic series of
stages to show the evolu-
tion of the Myrmeleontid
type from the original archaic formation of these veins, as still
preserved for us in the archaic Hemfirohiidce^ Drepanepteryx, and
Drepanacra.
* Phylogenetic series illustrating the evolution of the cubitus in the
wings of Hemerobiidoi and Myrine.Ieontiche. — Series a, h, c, three stages
leading to the reduced Hemerobiid type, also to the type found in hmd-
n-inr/s of Xymphidce and MyrmeJeontidoe. With a, compare forewing of
Drepanepteryx', with b, compare hindwing of Drepanepteryx; with c, com-
pare wings of Drepanomina. The final stage, in which the furrow / is
completely eliminated, is not shown in this figure. — Series a, b, d, three
stages leading to the formation of the cubito-anal vein in the forewing of
the Myrmdeontidn', by fusion of the veins lA and Cuo. In c?, the basal
part of Cuo is indicated by the cubital oblique vein Oc.
Lettering as on p. 118, and, in addition, k, the point destined to become
ciif, and s, the sector of Cu^ destined to become Cuib.
Cuib
Text-fig. 3.*
BY R. J. TILLVARt). 121
In Text-fig. 2, c, I have figured the base of the hind wing of
Xantholeon helmsi Tillyard, in order to contrast the simphfied
condition of this wing with the complicated condition of the fore-
wing discussed above. Since the MyrtuphMtntidm are admittedly
derived from Nymphid-like ancestors; and since, in the Nymphidce
themselves, both the media and the cubitus of the hindwing are
already simple, unbranched veins, it must follow that this con.
dition is to be found in their derivatives, the Myrmeleontidce. It
should, however, be noted that there are signs of specialisation
even in the hindwing, in that the vein lA approaches basally
very close to Cu, very nearly fusing with it; while 8c and R are
completely fused together throughout the basal portion of the
wing.
We have now to consider whether our new discoveries on the
structure of the cubitus require us to make any revision in the
present classification of the Myrmdeontidiv. Esben-Petersen, in
Part iii. of his "Australian Neuroptera,""^ has named the area
between Cuj^ + Mo, Cujb, and the hindmargin of the wing, the
intermibital area, and has pointed out that a Banksian Line is
sometimes developed in this space; such a line, he calls the inter-
cubital line. With this I am in agreement. Now the develop-
ment of an intercubital line is evidence of high specialisation in
this area. Therefore, those genera in which it occurs, if they
have hitherto been regarded as primitive, must have their posi-
tion reconsidered. In the tribe Creag7^itii, this intercubital line
is beautifully developed, and is correlated with a condition of
parallelism between Cui^ + Ms and Cujb- It was natural, so long
as this latter vein was taken to be Cuo, that this condition of
parallelism should be considered primitive. But the archaic
condition existing for Cujb is not one of parallelism to Cuja^ + M.,,
but rather, as may be seen in Xantholeon, one of strong diver-
gence from it, Cujb running obliquely to the margin, and parallel
to the numerous branches descending from Cuia-fMa. Xantho-
leon^ Gymnocnemia^ Periclystus, and other genera show this
♦These Proceedings, 1917, xhi., Pt.l, pp.203-219, PI. x.-xv. (PL x., fig.
1, Protoplectron striatellum Esb.-Pet. ).
122 STUDIES IN AUSTRALIAN NEUROPTERA, V.
primitive condition well. From this primitive state, two lines of
development may follow: —
(1) Ciij^ may become fixed in its divergent position, and the
intercubital line may be developed in the usual manner between
the descending branches of Cui;, + M.. Various stages in this
process may be studied \vithin the genus Ghmoleon, while the com-
plete formation may be seen in Acanthaclisis and ^lyrmdeoii.
(2) Cuib may become straightened out parallel toCuj, + Mo, the
intercubital line becoming developed between them, and parallel
to both. The beginning of this line of evolution is evident in
the forewing of Frotoplectron striatellum, Esb.-Pet., which is,
therefore, fairly primitive. In P. venustum Gerst., the forma-
tion is practically completed; w-hile in Creagris ^9/?*m6ews 01.,
(Palsearctic) the development of the three parallel veins, with all
connecting structures reduced to simple series of cross-veins,
reaches the highest possible stage of evolution along this line.
It would appear, therefore, that my estimate of the tribe
Creayrini, as the most primitive of the DendroleonthLce, is not
correct. They form, rather, a separate side-line of development
away from the main stem, culminating in a very highly specialised
genus, viz., Creayins. It should be noted, in this connection,
that the larva of Creagris is the only one amongst the Dendro-
leontince that is known to make pit-falls. It will now be of
special interest to find out whether the larvte of Protoplectron do
likewise.
123
ON THE AFFINITIES OF TWO INTERESTING F088IL
INSECTS FROM THE UPPER CARBONIFEROUS
OF COMMENTRY, FRANCE.
By R J. TiLLYAKD, M.A., D.Sc, F.L.S., F.E.S., Linnkan
Macleay Fellow of the Society in Zoology.
(With three Text-figures).
In May, 1917, Mr. Herbert Bolton, M.Sc, F.R.S.E., F.G.S.,
Director of the Bristol Museum, England, published an interest-
ing paper upon the "Mark Stirrup" Collection of Fossil Insects
from the Coal-Measures of Commentry (Allier), Central France."^
As is well known, these insect-beds are of Upper Carboniferous
Age, and have yielded a very large number of fossils, most of
which have been described by Brongniart and Meunier. The
chief characteristics of the assemblage may be shortly summed
up in the statement that they appear to have been at a stage
when the separate Orders known to us to-day were only beginning
to be foreshadowed, nearly all the specimens found being of large
size, with dense wing venation, and primitive structure of head,
thorax, and abdomen. The dominant group was the Blattoidea.
No undoubted Holometabolous Insects are known to exist from
these beds, nor were any such known from any Palaeozoic rocks,
until the discovery of Permochoi'ista, a genus of undoubted
Mecoptera, in the Permian Coal-Measures of Newcastle, N.S.W.f
Eight species are represented in the "Mark Stirrup" Collec-
tion. Five of these are Blattoids, and one is a Palseodictyopteron.
The other two are of very great interest, and their affinities are
certainly problematical enough to require very careful investiga-
tion before they can be settled with anything approaching finality,
* Mem. Proc. Manchester Lit. Phil. Soc, 1916-17, Vol.61, Pt.l, No.2,
pp.1 -.32, Pis. i.-v., [May, 1917].
t These Proceedings, 1917.
124 FOSSIL INSECTS FROM COMMENTRY, FRANCE,
I I'efer to Megctgnatha odonatiformis Bolton, and Sycopteroii
symmetrica Bolton. Bolton places the former in the family
Perlidce (by which the author evidently means the Ordei- Perlaria,
and not the family Perlido' s.str.); though, in a note added to
this determination, he states that Dr. A. D. Imms suggests that
the specimen "may possibly come nearer to the Sialidce,'' by
whicli, I take it, he means the Order Megaloptera, rather than
the actual family Sialidce s.str. As for Sycopferon, this is
assigned to the "family Fanorpidce" for which, again, we are
evidently intended to read the "Order Mecoptera," since the
author compares it with Orthophlehia of the Lias. If this placing
be correct, then we have a Holometabolous insect present in the
Upper Carboniferous — an occurrence whicli is, of course, not im-
possible, but is certainly improbable, in view of our present
knowledge of the Phylogeny of the Insecta.
I have, of course, no opportunity of examining the specimens
themselves; but Mr. Bolton has given us such excellent photo-
graphic reproductions and drawings of the fossils in his paper,
that one would certainly scarcely desire anything better. As.
the determination of the affinities of these two fossils is a matter
of prime importance to students of Insect Phylogeny, and as I
have already informed Mr. Bolton by letter of the views that I
shall here express, I propose now to state my reasons why these
two insects cannot be accepted as belonging to the Orders to
which Mr. Bolton would assign them, and to indicate also to
what known Orders they may, with more probability, be con-
sidered to belong.
Megagnatha oDONATiFORMis Bolton. (Text-fig. 1).
Bolton, op. cit., pp.2-6, PL i., figs. 1-4.
From the photograph given by Bolton in PI. i , fig.l, and from
the drawing in Fig. 3, it is evident, as Bolton himself admits,
that any reconstruction of the wing-venation of this fossil must
be largely a matter of conjecture. We should, therefore, attempt
to find other characters that may lead to a solution. The figures
show that: —
(1) The wings were either absolutely, or approximately, equal
BY R. J. TILLYARD. 125
in size, and of similar shape; probably, therefore, the venations
of fore- and hind wings were the same.
(2) The insect had a small, elongated head and prothorax.
(3) The head carried a pair of straight, slender, and fairly long
antennse, and also a pair of shorter, but very prominent, some-
what curved, slender appendages, which Bolton considers, with
some doubt, to have been the mandibles.
(4) The three pairs of legs were well developed, and placed
far apart from one another.
(5) The abdomen was also moderately short, and probably
carried two very short cerci.
In the Perlaria, the head and prothorax are neither narrow
nor elongated, but are always more or less flattened down dorso-
ventrally, more or less widened; and the prothorax is never far
removed from the pterothorax. The antennae are very long, and
the cerci also, in all the most archaic forms; genero, in which the
cerci are shortened are demonstrably descended from forms which
had longer cerci. Further, fore- and hindwing are never equal
in size, or of similar shape; and forms in which there is an ap-
proximation to equality are demonstrably derived from forms in
which there has been greater inequality. Nor are the venations
of fore- and hindwing ever the same, but differ fundamentally,
as a study of the nymphal tracheation of the two wings clearly
proves. Finally, the Perlaria have mostly reduced, weak, non-
projecting mandibles; and it is demonstrable that these organs
were never, within the limits of this Order, slender and project-
ing, as seen in Megagiiatha.
Even if it were to be admitted that the restored venation in
Bolton's Fig. 4 were correct, I fail to see in it any true Perlarian
characters. It much more resembles the venation of an archaic
Termite forewing, such as Mastotermes.
I conclude, therefore, that no affinity between MeijayiuUha and
the Order Perlaria can be shown to exist.
The only possible claim to affinity with the Order Megaloptera
rests upon the projecting mandibles, since these structures also
project strongly in the a,H!:haJ\c Corydalus and allies. But the
126
FOSSIL INSECTS FRoM COMMENTRY, FRANCE,
mandibles of these latter insects are much more strongly built,
and are attached to a broad and massive head that is as unlike
that of ^fegagnatha as it well could be. We may well ask, are
these "mandibular-like structures" mandibles'? Is it not much
more likely that they are the maxillary jDalpi, whose joints, like
those of the antennae, have become obscured in the fossil? They
are too slender to have been of any use for biting; and sucking
mandibles, such as we find in the Ant-lion, do not occur, as far
as we know, in any imaginal form whatever; nor are even such
suckin^i; mandibles ever so slender as in this fossil.
Text-fig. 1.
Mt(jagnatlui odonatiformisl^ditoxi', {y-i:2). Upper Carboniferous of Coni-
mentry. From Bolton's PI. i., fig. 2. The short cercus on the left
side of the figure is not shown in Bolton's figure, but is visible in his
fig. 1 (photograph).
T am, therefore, forced again to conclude that no real aliinity
can be demonstrated between this fossil and the Megaloptera.
BY K. J. TILLYARD.
127
What, then, are the most likely affinities of this peculiar fossil?
I should answer that it shows a very remarkable resemblance to
the very ancient and mysterious Order Embioptera, about which,
unfortunately, we know very little. In Text-fig. 1, I reproduce
Bolton's figure of Megaynathci, and place opposite to it, in
Text-fig. 2, a drawing of an Embid {Oligotoma sp., undetermined)
which I happen to have in my collection. It must be at once
admitted that the resemblance is a very striking one. The form
of the head, prothorax, and legs, the great separation of the
latter, and the equality of the wings, are characters that separate
Text-fig. 2.
Oliyotonm sp., (undetermined) in Coll. Tillyard; { x 7^). Recent.
Brisbane, Queensland.
out the Embioptera very distinctly from all other groups. All
these characters are possessed by Megaynatha. Further, the
Embioptera possess elongated maxillary palpi, which, in the
position of rest, lie projecting in front of the head, much in the
128 FOSSIL INSP:CTS from COMMENTRY, FRANCE,
manner that these supposed mandibles of Mpgm/nafha do in the
actual fossil. The only two discordant characters are the much
more complex venation of the fossil, and the closeness of its wings
at their bases. The latter may be easily explained as having
been due to unequal crushing of the thorax: probably the wings
were actually well separated at their bases, as the structure of
the thorax, and the position of the legs, undoubtedly suggest.
As for the venation, it is quite evident that recent Embioptera^
like uKJst recent Termites, possess a very reduced venation, con-
taining only very little of the original elements. We see, in
JIastotermes, how complex was the venation originally possessed
by the Termites. In the Embioptera, the analogue of Masto-
termes no longer exists; or, at any rate, it has not yet been dis-
covered. But I have seen a species from Australia (of which,
unfortunately, I have no notes or figures) with a venation con-
siderably more complex than that of Oliyotoma. All students of
the Embioptera agree that they represent the last remnants of
a peculiar group of great antiquity; but, so far, their fossil history
has been almost completely missing.*
Meyax/natha is larger than Oliyotoma', but this is what we
should expect, if the two are really related. For OliyoUnna is
clearly a reduced form, as its venation proves.
T would suggest, therefore, that Meyaynatha odonatiformis is
in reality an ancient representative of the Embioptera, and
should be placed within that Order, as the sole known type of a
new family, the J/egaynafhidft', differing from all known members
of the Order by its greater size and more complex venation, as
well as, probably, by tlie shorter comparative distance between
the bases of the fore- and hind wings.
As the identification of the "mandibular-like'" structures as
true mandibles is, at the best, very doubtful, and as not a single
Odonate character is possessed by this fossil, it is much to be
regretted that its author should have chosen a generic name
* Two doubtful records from West African Copal can scarcely be
regarded as more than subfossil, and add nothing to our knowledge of the
group. A single species from Baltic Amber belongs to the genus Oliyotoma-
BY R. J. TILLYARD. 129
based upon the former character, and an entirely misleading
specific name, neither of which can be altered.
Sycopteron symmetricum Bolton. (Text-fig. 3, 6, c).
Sycopferon symmetrica Bolton, oj). cif., pp.6 8, PI. ii., figs. 1-2.
If this fossil could be proved to be Mecopterous, it would be
one of the most striking discoveries in Pal?Poentomology. For
that reason alone, we are all the more bound to examine the
evidences of its supposed Mecopterous affinities as carefully as
possible.
Text-fig.3, b, c, are portions of Bolton's PI. ii., fig. 2, which, as
far as I can see by comparison with the photograph in his fig. 1,
is accurate in all except a possible minor detail or two. The first
thing that strikes one, on examining this fossil, is the way that
its wings are folded down the back of its abdomen. If this were
the natural position of rest, it would be that of a Dipteron or a
Hymenopteron; yet both these Orders are excluded from the
question, for reasons that must be sufficiently obvious- without
stating them. We have, then, to conclude, either that SycojJteron
belonged to some extinct Order, which folded its wings in this
position (a highly improbable theory), or that it belonged to some
Order in which the wings were held roof -like over the abdomen;
in which case, the flattening down of the wings in the fossil
may well have caused some underfolding of either the costal or
the posterior border of the wings, or both.
Bolton claims for his insect a Panorpoid venation, stating that
its nearest approach is to be found in Orthophlehia of the Lias.
When, however, he goes into detail, it is quite clear that he is
unable to homologise the separate veins of Sycopteron with those
of Orthop)}dehia^ without getting into very serious difficulties. In
order to show this, I give in Text-fig.3, «, 6, the venation of
Sycojiteron, (h), as interpreted by Bolton, and the venation of
Permochorista, (a), which is the oldest fossil Mecopteron known,
and the venation of which closely resembles that of Orthophlehia.
It will be seen at once that the type of venation present in the
Mecoptera (and there is no Order in which the venation is more
130
FOSSIL INSECTS FROM COMMENTRY, FRANCE,
constant in character, differing little right through from the
Permian to recent times) is radically different from that of
Text-fig.3.
a, Forewing of Permochorista mitcheUi Tillyard, (restoration, with all
the cross- veins omitted); { x 4). Permian of Newcastle, N.S.W. h, Fore-
wing of Sycopteron symmetricnm Bolton, with his naming of the veins;
( X 7j). Upper Carboniferous of Commentry. From Bolton's PL ii., fig.2.
c. Head and thorax of same, from the author's same figure; (x7f). d,
Forewing of Amphientomum paradoxnm Br. ;(xl5). Oligocene, Baltic
Amber. From Enderlein. In b, the dotted vein x indicates the probable
position of the basal piece of Rs, not shown in Bolton's fig.2, but appa-
i-ently slightly indicated in his fig. 1 (photograph).
Sycopteron^ this latter being of a much simpler type, which does
not occur in the Order Mecoptera at all. The resemblance is
BY R. J. TILLYARD. 131
due only to the method of branching dichotomously, and the
absence of clearly-marked crossveins, — characters which are by
no means confined to the Mecoptera.
I would suggest that the true costa of this insect has become
folded under in the fossil, and is not clearly visible. Bolton
says, "The costal margin seems to have been extremely delicate,
and to have left very faint traces of its position." This might
well be true of the subcosta, which is frequently a weak vein.
In the Mecoptera, the costa is strongly formed, but there are
other insects in which it is not so. This suggestion receives
support from the fact that, if the front vein preserved in this
fossil is really the costa, then Sc, R, and M all come off from a
common stem, in a manner that cannot be paralleled except in
the Homoptera, and certainly never occurs in the Mecoptera.
Whether w^e allow that the costa was underfolded or not, the
following dilemma has to be faced: —
(1) If the fossil is Mecopterous, then the naming of the veins
by Bolton is incorrect. The five-branched vein which he has
distributed between R and Rs is certainly the media; and we
are then driven to suppose that all the rest of the venation,
costad of this, has been underfolded, except a portion of Rs, which
would be Bolton's Sc.
(2) If Bolton's naming of the veins is correct [or even if the
large extent of underfolding suggested in (1) cannot be admitted],
the fossil is neither Mecopterous, nor in any way related to the
Mecoptera, since the typical Sc, R, and Rs of that Order — the
latter alone of which is never less than four-branched* — are all
absent.
From this dilemma, there is only one escape, viz., to admit at
once that the Mecopterous atiinities, much as we must regret it,
cannot be proved, and to seek for some more likely solution of
the problem.
In Text-fig. 3, 0?, I show the venation of the forewing of the
very archaic insect Amphientomum paradoxum Br., from Baltic
* Excepting in the highly reduced, recent XannochoristidiV, where it is
three-branched.
132 FOSSIL INSECTS FROM COM M ENTRY, FRANCE,
Amber. This insect belongs to the Order Psocoptera or Copeo-
gnatha, an Order which is not Holometabolous, but which pos-
sesses a reduced venation very closely resembling that of certain
Holometabolous Orders, in particular the Hymenoptera. Nearly
all recent Psocoptera have a much more specialised venation than
Amphit^nfomum; but close relatives of the latter still exist in the
peculiar genera Echmepteryx and Ci/mafopsocus, as well as the
genus Amj)hi('7ifo7num itself, which has recently been rediscovered
in Ceylon.
It will be seen that the correspondence between the wing-veins
of Sycopteroii and Ar)ip)hiento7nuin is exceedingly close; in fact,
they can be completely homologised, provided only that Sycopf-
eron possesses the short basal piece of Rs which is absent from
Bolton's figure. A close examination of the photograph of
Sycopteron, in Bolton's PI, ii., fig. 1, suggests to me that this piece
actually does exist; at any rate, it seems to be clearly indicated
on the left wing. The fossil should certainly be further examined
to determine this point.
In Am^jhifiitoiHum, the subcosta is a very weak vein, merging
into the costa before halfway. The radius is a strong vein, forked
distally, and giving off its sector far distally, at a point only just
before its fork. Rs itself is forked, as in Sycopteron (the vein
Bolton calls R). Further, M is three-branched in Am^Dhientomum,
in the same manner as in Sycopte7-on (the vein Bolton calls Rs).
In Amphimfom^im, the cubitus is forked far distad, and is
attached to the stem of M at a point quite one-third of the wing-
length from the base; in Sycoptero7i, the attachment and forking
lie both much further basad. So also the anal veins in Amphi-
entomum are less primitive than in Sycopteron, being shorter and
less straight.
Now Amj^hientomum is evidently specialised in the following
points : —
(1) The shortening of Sc.
(2) The removal of the origin of Rs far distad, and consequent
shortening of that vein.
(3) The close union basally between M and Cu.
BY R. J. TILLYARD. 133
(4) The removal of the cubital fork far distad.
(5) The shortening and curvature of the anal veins.
If, then, ancestors of this insect existed in Palseozoic strata,
we should expect them to exhibit a generalised condition in these
five characters. This is exactly what Sycoptproit does exhibit.
Without stating positively that Sycopteron lies in the ancestral
line of Amphienfojnum, yet we can affirm that, as far as its vena-
tion is concerned, the two are certainly closely allied, and
Sycopteron is the older type.
Let us now examine the rest of the fossil for clues as to its
affinities. The first peculiarity to be noted is the huge boss-like
areas upon the back of the head (Text-fig. 3, c). If these be com-
pound eyes, then Sycopteron is certainly neither a Mecopterous
nor a Psocopterous insect; but, as Bolton mentions no facets on
these areas, we may confidently take it that they are not. They
must, therefore, be epicranial or occipital swellings. Now the
head of Sycopteron is much too prominently developed to agree
with the Mecopterous type. But almost all the Psocoptera have
a comparatively large head, with prominent development of the
posterior portion of the epicranium. If such a head were to be
crushed somewhat in becoming fossilised, its appearance would
certainly closely resemble that of Sycopteron. The small eyes
would probably be completely covered by the flattened epicranium.
The structure of the thorax in Sycop)teron is certainly primitive,
and might indicate equally well affinities with several Orders,
including both Psocoptera and Mecoptera (Text-fig. 3, c).
Again, in the Psocoptera, the hind wing is always small and
weakly chitinised. Hence, in insects of this Order which might
become fossilised with their w^ngs flattened down over the abdo-
men, no sign of the hindwing or of its venation would appear.
The Mecoptera, on the other hand, have, of all the insects, except
the Odonata and Planipennia, the most strongly chitinised hind-
wings, of a size almost equal to the forewings, and with a strongly
marked and closely similar venation. Hence, if a Mecopteron
became fossilised in the position we are supposing, it seems
inevitable that traces of the hind wings and of their main veins
10
134 FOSSIL INSECTS FROM COMMENTRY, FRANCE.
must be present. But Bolton mentions nothing of this sort, and
the photograph in his PL ii., fig.l, certainly shows nothing that
could be so interpreted. The probability, then, of this fossil
having belonged to the Mecoptera, is very much diminished upon
this count alone
Taking all the evidence into consideration, it seems legitimate
to conclude that Sycopteron is most certainly not a member of
the Order Mecoptera, but that it is very likely an archaic type
of the Order Psocoptera, related to Anijyhientomum of the Oligo-
cene, but considerably less specialised, in that it lacks all five of
the venational specialisations possessed by this genus. It should
certainly be placed in a new family Sycopteridce, having the
characters already mentioned. It should be noticed that the
size of this insect (wings 9-10 mm.) agrees very well with its
inclusion in the Order Psocoptera, all the known members of
which are small, ranging from a wing-length of only 10 mm., at
most, down to very minute and wingless recent forms.
135
ORDINARY MONTHLY MEETING.
April 24th, 1918.
Professor H. G. Chapman, M.D., B.8,, President, in the Chair.
Mr. Clyde Douglas Gillies, M.Sc, University of Queensland,
Brisbane, and Dr. Henry Priestley, B.Sc, University of Syd-
ney, were elected Ordinary Members of the Society.
The President announced that the Council had elected Messrs.
C. Hedley, F.L.S,, W. W. Froggatt, F.L.S., W. S. Dun, and
A. G. Hamilton to be Vice-Presidents; and Mr. J. H. Campbell
[Royal Mint, Macquarie Street] to be Honorary Treasurer — for
the current Session, 1918-19.
On the motion of Dr. Cleland, it was resolved : that the con-
gratulations of Members be tendered to Dr. S. J. Johnston, B.A.,
on his appointment to the Chair of Zoology in the University of
Sydney.
The President offered the congratulations of Members to Dr.
R. J. Tillyard and Dr. A. B. Walkom (in ahseiitid) on their
attaining the Doctorate of Science.
The Donations and Exchanges received since the previous
Monthly Meeting (27th March, 1918), amounting to 51 Parts or
Nos., 7 Bulletins, 2 Reports, and 5 Pamphlets, received from 30
Societies, etc., were laid upon the table.
136
THE ORIGIN OF YOLK IN THE OVA OF AN
ENDOPARASITIC COPEPOD.
By H. Leighton Kesteven, D.Sc, M.B., Ch.M.
(Plate iii.).
In January, 1905, Professor J. P. Hill presented me with
material for the study of the morphology and development of an
endoparasitic copepod infesting Ftychodera an.sfralu'usis Hill.
The parasite was found to be new, and was subsequently described
under the name of Ubius hilli Kestv., in 1913(3). The following
observations were made on the developing, ultimate oogonium,
and primary oocyte at the time U. hilli was studied (1908), and
were submitted to Prof. Hill for criticism. At his suggestion,
this paper was withheld from publication because, at that time,
the material on which the observations were made was all stained
in one way. Since then, I have obtained fresh material, which
was fixed in {a) 5% formaldehyde in seawater; and (h) Miiller's
bichromate-solution. Specimens, after cutting, were stained with
(1) Delafield's htematoxylin, (2) in Heidenhain's haematoxylin,
(3) Mayer's carm-alum, (4) Flemming's method for karyokinetic
figures.
This staining was done over three years ago, but press of
other work has, till now, prevenced me from examining the
results. Recently, I have found time to go over these sections,
and I find that I have nothing to add to the original paper. I
am satisfied, that the observations recorded truly represent pro-
cesses taking place in the developing cell, and are not artifacts, for
I find the cells to present these appearances, not only in different
specimens similarly stained, but also in the same and in different
specimens dissimilarly stained. Were I situated in a city near
a library, I doubt not I would hnd many reasons for altering
BY H. LEIGHTON KESTEVEN. 137
that portion of the following paper which deals with the litera-
ture. This paper was written in 1908, and since then I have
had no opportunity of keeping abreast of the cognate current
literature, and, at present, T am beyond the reacli of a libraiy,
I believe, however, that even though this contribution is thus
nine years old at date of printing, it is deserving of publication.
The ovarian epithelium in the young female is composed of
fairly regular, cubical oogonia.* The continued division of these
gives rise to the mass of ultimate oogonia which fills not only
the lumen of the ovary, but also the anterior portion of that of
the oviduct. It is while lying free in the lumen, in this mass,
that the ultimate oogonium advances so far towards maturity,
that it may thereafter be regarded as the primary oocyte.
AVhen first shed into the lumen of the gland, the ultimate
oogonium is a small, rounded, hyaline cell. Its comparatively
lai'ge nucleus may contain only one karyosome, or it may contain
two, three, or four of approximately equal size. The size of the
inicleus is defined by the nuclear membrane alone; that is to say,
the nucleoplasm is hyaline, and takes no stain, nor is it difl^'eren-
tiated from the cytoplasm by the presence of a discernible chro-
matin-reticulum (Fig. 1). In those cases where there is only one
karyosome, I am unable to find that this a»e differs from the three
or foul' in other cases, nor among these is there any difference
inter se.
The cell now enlarges. The nucleus, increasing in size at a
greater rate than the cell, comes ultimately to fill nearly the
whole cell.
When this process has reached the stage depicted in Fig. 2, a
chromatin-reticulum is well established, and the karyosomes have
increased in number. In short, the period is characterised by an
increase of chromatin.
As soon as this increase has reached its maximum, the reticulum
becomes broken down, till, as depicted in Fig. 3, in place of a
network, there is present a great number of fine granules of
* The description is of material stained with htematoxylin without any
counter stain.
138 ORIGIN OF YOLK IN OVA OF AN ENDOPARASlTIC COPEPOD,
chromatin, and the kaiyosomes He free in the nucleoplasm. The
period is characterised by the disintegration of the chromatin-
reticulum formed during the last period.
The fine granules of chromatin next begin to increase in size,
and lose in depth of staining, until, instead of being opaque
black points, they become semi-translucent, purple spherules. The
karyosomes meanwhile remain unchanged (Fig. 4). This pheno-
menon, I regard as the formation of the first yolk-granules.
During succeeding stages, they continue to increase in size.
The karyosomes very soon exhibit signs of activity. Each of
them, from a solid sphere of chromatin, becomes converted into
a small, spongiform mass (Fig. 5), probably due to the formation
of vacuoles within them. What this activity, which characterises
the period, means, I am quite unable to say, but I do not think
that it can affect the deductions made later.
Meanwhile, the nucleus has so increased in size, that it is now
surrounded by a mere envelope of cytoplasm, the presence of
the nuclear membrane being evidenced more by the definite out-
line of the nucleus, than by the actual visibility of the membrane.
The next period is characterised by the formation of a new
and much smaller nucleus within the old. This takes place in
three steps.
Around one, or it may be two. or three, coalescent, spongiform
karyosomes, there becomes recognisable an area of plasm devoid
of the spherules, which are scattered plentifully throughout the
rest of the nucleus, and this area of plasm takes a faint purple
stain (Fig. 6).
This purple-staining globule of plasm, with its contained karyo-
some or karyosomes, by this time quite coalesced, is next enclosed
in a distinct membrane (Fig. 7).
The karyosome once more assumes a solid spherical form, the
surrounding plasm still staining purple.
This area henceforward constitutes the nucleus of the cell; it
is shown in Fig. 8, which is a section of a mature, primary oocyte.
Meanwhile the remaining karyosomes, scattered throughout the
old nuclear area, have also shrunken to their previous size and
BY H. LEIGHTON KESTEVEN. 139
shape, and aoain become solid; during this time, none of them
were surrounded by a clear area of plasm staining purple.
The old nuclear membrane is apparently still present, in some
cases, until a much later period; but, in others, it seems to have
broken down at the time when the new nucleus first develops its
membrane.
Up to this time, the ultimate oogonium has lain free in the
lumen of the gland or oviduct; it now becomes attached to the
wall of the latter, either to one cell by a foot, as in Fig. 9, or, in
the absence of the foot, to several cells.
Growth in size appears now to take place verj^ rapidly, and
there is a great increase in the quantity of yolk-spherules. No
changes are observable in the new nucleus.
Concurrently with the increase in size of the cell and quantity
of yolk, the karyosomes left free in the old nuclear area appear
to be dissolved without showing any further signs of activity.
When the cell, now to be regarded as a primary oocyte, has
reached its full size, it is once more set free into the lumen of
the oviduct.
There is present in the mature primary oocyte only one set of
spherules, variable in size, certainly, but differing in no othei'
way from one another. All are yolk-spherules or none are yolk-
spherules. The spherules which were formed by increase in size
of the chromatin-granules are, therefore, similar to, and may be
regarded as being the same as, those formed later.
After these observations had been made, and the conclusions
given below had been deduced from them, I sought, in the pub-
lications of previous workers, for comparable observations and
deductions. I cannot better give the results of my examination
of literature than by the following quotation from a paper by
Caroline McGill(5), who, on p. 21 9 of the paper quoted, expresses
the opinion that "it seems probable that chromatin may have
something to do with yolk-formation."
Will (7) thinks that the larger nucleoli of the amphibian
germinal vesicle pass out into the cytoplasm, and there become
yolk-nuclei. MacCallum(4) concludes that, in the ova of Am-
140 ORIGIN OF YOLK IN OVA OF AX ENDOPARASITIC COPEPOD,
phibia, the peripheral nucleoli generate a substance which diffuses
first into the nucleus, and from there into the cytoplasm; finally,
it combines with the cytoplasm to form yolk. Henneguy(2) be-
lieves that the corpuscles of Balbiani in vertebrates are either
parts of the nucleolus, or the entire nucleolus, which pass through
the nuclear wall into the cytoplasm. Montgomery (6), in Pisicola,
describes the nucleus as contracting in volume, and, in so doing,
discharging all except one of its nucleoli into the cytoplasm.
Goldschmidt(l), in active gland-cells and in muscle-cells of Ascaris,
has described a cytoplasmic chi'omatin, which, instead of being-
gathered into irregular masses as in the Kissl bodies of nerve-
cells, is arranged in fibres or coarse reticula. In most instances,
however, this chromatin, which he calls 'Chromidialapparat,' is
not derived from nucleolar material, but represents nuclear chro-
matin which has made its way into the cytoplasm. In fact, in
many cases, the chromatin-fibres of the cytoplasm extend directl}'
through the nuclear membrane, and are continuous with the
chromatic reticulum of the nucleus. Since the chromatic appa-
ratus is more highly developed in active than in resting cells,
Goldschmidt concludes that it umst function in the metabolism
of the cytoplasm.
My own deductions may now be put very briefly. They are : —
1. The yolk-granules are formed by the combination of a cyto-
plasmic constituent with chromatin.
2. The first yolk-granules are formed within the nucleus.
3. The formation of the new nucleus is a pseudo-contraction of
the overladen, old nucleus.
4. This pseudo-contraction leads to the shedding of some of the
karyosomes, which are henceforward to be regarded as yolk-nuclei.
5. These yolk-nuclei are stores of chromatin, which are to con-
tinue the functional activity of the nucleus of the growing pri-
mary oocyte, that is to say, they are to supply chromatin for
that combination which is yolk-formation.
6. The ultimate oogonium is nourished by endosmosis, the
primary oocyte by the epithelium of the oviduct. In both cases,
the all-important substance recei\ ed is the cytoplasmic constituent
B¥ H. LEIOHTON KESTEVEN. 141
which enters into yolk-formation. Clii'omatin-constituents, how-
ever, must be derived from without in tlie earliest stages.
LITERATURE.
1. (liOLDSCHMiDT — ■" Dev Clironiidialapparat lehliaft funetionierender (le-
webszellen." Zool. Jahrb., xxi., 190.").
2. Henneguv — "Le corps vitellin de Balhiaiii dans Wvni des Vertebre.s."
Journ. Anat. Physiol., xxix., 1898.
3. Kesteven — A new Endopaiasitic Copepod : M(>rpholo!j;y and Develop-
ment. Proc. Linn. Soc. N. S. Wales, 1912, xxxvii.U918).
4. MacCallum — "Contribution to tlie Morphology and Physiolooy of the
Cell.'" Trans. Canadian Inst., i., 1891.
5. McUiLL — "The Behaviour of the Nucleoli durino- Oogenesis of the
Dragonti3' with special reference to Synapsis. " Zool. Jahrb., xxii.
1906.
6. Montgomery — "Comparative cj^tological Studies, witii special regard
to the Morphology of the Nucleolus." Journ. Morphoh, xv., 1899.
7. Will — "Ueber die Entstehung des Dotters und der Epithelzellen bei
den Amphibien und Insecten." Zool. Anz., vii., 1884.
EXPLANATION OF PLATE III.
Explanatory letters, — K., kar3'osomes — Nu'., the new nucleus in process
of formation — Nu"., the new nucleus nearlj'- formed — Nu. memb'., the
nuclear membrane of the old nucleus — Nu. memb"., the juielear membrane
of the new nucleus.
Fig. 1. — The ultimate oogonium when first detached from the wall of the
ovary.
Figs.2, 3, 4, 5. — Stages in the growth of the ultimate oiigonium. ,r^'-";:
Figs. 6, 7. — Stages in the formation of the new nucleus.
Figs. 8, 9. — Sections of mature primary oueytes. iO">:,^!, ''i^J /.>'. K
142
CONTRIBUTIONS TO OUR KNOWLEDGE OF SOIL-
FERTILITY.
No. xvi. The Search for Toxin-Producers.
By R. Greig-Smith, D.Sc, Macleay Bacteriologist to the
Society.
In the earlier papers of this series, it was shown that bacterio-
toxie substances were sometimes present in the clear liquid ob-
tained by shaking a soil with water and filtering it through
porcelain. The toxins were potent, for, when the same portion
of soil was extracted a second time with water, the extract was
highly nutritive. The first extract undoubtedly contained sub-
stances which functioned as toxins, as well as substances which
acted as nutrients, and, of these two, the toxins were the more
powerful.
The local soils were found to be toxic during the cold winter
months and not in the dry summer. The toxicity was variable,
and even in the favourable season, one could not be certain that
a soil, undoubtedly toxic at one time, woiild be toxic at another.
It seemed to be a matter of chance, but this was undoubtedly
due to the fact that the cause of this toxicity was unknown.
With the elucidation of the origin of the toxins, it will be better
understood when a soil is likely to be toxic or otherwise.
Soil-toxicity, as exhibited in the extracts of the soils, can be
demonstrated directly and indirectly. Directly, by adding a
certain number of a test-bacterium, and obtaining a diminution
in that nnmber after an incubation-period of, say, 20 hours at
22°. Indirectly, by obtaining an increased growth as a result of
boiling the extract and also of diluting it. A toxic soil exhibits
all three characters. There are other points connected with
toxic soils, and these will be found in my former papers.
During my earlier work, I was led to believe that the cause
pf the toxicity would be found in the products of the bacteria,
BY R. GREIG-SMITH. 143
and, in the beginning of this research, into the etiology of
toxicity, some of the more Hkely bacteria were tested with more
or less completeness. The work was tentative, the bacteria were
tried one way and another. A favourable result was occasionally
obtained, which led to repeated trials with slight variations, but
these ended in failure to obtain a truly toxic condition. Some
moulds were also tested, but the experiments with these did not
lead me to believe that the source of the toxins would be found
in the flora of the soil. The fauna remained, and when the
amcebae were tested, the first results were so satisfactory that the
source of the toxins seemed to have been traced. Unfortunately,
these results were not confirmed, and, as the work proceeded, it
became evident that the toxic effect was caused by the produc-
tion of alkali in the solutions in which the protozoa were growing.
The test-bacteria were very sensitive to changes of reaction, and,
so far as the solutions were concerned, it was made clear that a
perfectly neutral solution was exceedingly difficult to maintain,
and that any departure from a strict neutrality retarded the
growth of the bacteria and exhibited some of the effects of a
toxic solution. The main steps of the work that led to this
conclusion are recorded in the following pages. Many experi-
ments have been omitted because the results did not appear to
justify a lengthening of the paper. They were either indefinite
or confirmed a negative result previously obtained.
It is well known that some soil-bacteria are inimical to others,
and it was considered that some one group might be specially so
to bacteria generally. Instances of an inhibiting or toxic effect
exercised by some micro-organisms against others are familiar to
most workers in soil-bacteriology. It is an ordinary experience
to find certain colonies of bacteria, such as Bac. mycoides and
certain moulds, passing over or through other bacterial colonies
when spreading over the surface of an agar-plate. Occasionally,
they are seen to avoid some particular colony, and we find these
surrounded by a clear zone of agar, across which the wandering-
mould or bacterium will not pass. Apparently, the colony has
sent into the medium some toxic substance, and there does not
144 CONTRIBUTIONS TO OUR KNOWLEDGE OF SOIL-FERTILITY, xvi.,
appear to be a thinning away of the toxin, for the wandering-
colony generally becomes heaped up at the margin of the toxic
zone and then spreads round and eventually encircles the colony.
It may be that the thickening of the edge is caused by the toxin
becoming so weak as to be able to exercise a stimulating action
like other weak poisons."^ The Actinomyces-colonies are gener-
ally toxic to such wandering moulds and bacteria.
As soil contains so much sand and inert matter, it did not
appear to be a suitable medium for experiments with these pre-
sumable toxin formers, and it was considered that a fluid medium
would be better. With regard to the nutrient added to the
water, Bottomley's work with auximones had suggested the idea
that there might be a relation between them and decayed soil-
toxins. Bottomley used moist peat-moss as a culture-material,
but, as this could not be obtained, the first experiments were
made with washed sphagnum-moss suspended in water.
The moss was picked, dried, cut up, washed, and again dried.
Ten grams were put into a flask with 600 c.c. of tap-water and
sterilised.
Two bacteria, T.P.2 and T.P.4, were selected as being possible
toxin-producers. They were not identified at first, for, in ex-
periments such as these, one determines the value of the bacte-
rium first and identifies it afterwards, if its activity justifies the
identification.! The bacteria were seeded into the flasks of sus-
* Journ. Roy. Soc. N. S. Wales, 1916, p.77.
t Bac. T.P.2 is a short, motile rod measuring 0"5 : 2yut. Gelatine is
liquefied slowl3% the colonies being round, white, and slightly raised with a
ciliate edge. On agar, the growth is ivory-white, rough and cohesive.
Long, needle-shaped crystals are quickly formed in agar containing sodium
phosphate, and these are fairly characteristic of the organism. In some
media, slime is formed from dextrose, althougli none is produced on
Lipman-Brown agar. In bouillon, the groM th produces turbidity, a film
and a sediment; nitrates are not reduced. Milk is coagulated and slowlj'
peptonised. On potato, the growth is yellow-brown. The indol test is
positive, the Gram test negative, and the bacillus produces neither gas or
acid from dextrose, saccharose, mannite, or glycerine. Nitrate is not pro-
duced from ammonia salts. It appears to have Bac. imincHis as its closest
ally.
BY R. GREIG-SMITH.
145
pended moss, which were incubated at 22" for varying periods.
The extract was prepared in the usual way by filtration first
through paper, then through porcelain. One portion of the
extract was boiled for an hour under an inverted condenser. The
various portions, generally SOc.c, of the raw, boiled, and diluted
extracts, were each seeded with 1 c.c. of a suspension of Bac.
2)7-odi(jiosus, and incubated at 22° for 20 hours, when counts
were made by the plate-method. The numbers of bacteria in the
raw extracts were taken as 1 00, and those in the treated extracts
were calculated in terms of this. Fractions were omitted, and
numbers less than 1 were taken as 1. The actual number of the
water-control can be found by dividing the extract by the
extract/water ratio, for the latter was obtained by dividing the
extract-count by the water-count.
Experiment i.
Growth of Bac. prodigiosus
in extract, 20 hours at 22°.
Penicillium clado-
Micro-organism.
T.P.4
T.P.'J
sporioides.
Nature of moss.
new
new
old
new
old
Duration of test; days —
29
8
12
11
10
13
10
11
Number of test.
1
2
3
4
5
6
7
8
Extract, boiled
766
223
72,000
600
5,500
34
184
1,167
Extract, raw ...
100
100
100
100
100
100
100
100
Extract, raw, 80%
33
54
122
92
100
479
90
48
Extract, raw, 60%
15
15
133
100
100
2,240
67
40
Extract, raw, 40%
6
4
300
107
100
600
56
14
Extract, raw, 20%
2
1
167
107
135
21
36
40
Water-control ...
1
1
HI
77
88
1
25
8
Extract/water ratio ...
417
580
0-9
1-3
1-2
111
4
12
In looking over the results of Experiment i., it is seen that
moss is not a good substance for determining toxin-23roduction.
When new, it is too nutritive, and when old, that is, when it had
been used and washed once or twice, it is too poor. The dilution-
curves of tests 3, 4, and 5 are almost horizontal lines, indicating
that the extract is of a nature similar to water. The results
obtained by boiling the extract in tests 3 and 5 cannot be ex-
146 CONTRIBUTIONS TO OUR KNOWLEDGE OF SOIL-FERTILITY, xvi.,
plained; they certainly indicate a degree of toxicity which is not
confirmed by the dilution-numbers.
The numbers Avith Penicillium cladosporioides are peculiar. A
flask wdth new moss had been sterilised and allowed to stand for
several w^eeks before being infected with Bac. T.P.2. At the
end of the incubation-period, the mould was plainly seen growing
as a floating mass upon the surface; it had ousted the bacteria,
for few bacterial cells were obtained from the fluid. The rise in
the numbers upon dilution is typical of a toxin, but the reduction
upon boiling is not. Again, the dilution-effect was not obtained
in tests 7 and 8.
On the whole the experiment was unsatisfactory, and con-
sequently other media were tried.
Experiments were made with dilute solutions of nitrogenous
salts such as ammonium sulphate, ammonium phosphate, and
potassium nitrate, as well as with alkaline salts as potassium
phosphate, but there was little sign of any probable formation of
toxin with any of them.
The USE of Collodion-Capsules.
In an endeavour to improve the experimental method, use was
made of collodion-capsules, as it w^as considered that by growing
the bacteria outside and the test-organism inside the capsule, the
production of toxin might be rapidly determined. Accordingly,
capsules were prepared by coating the insides of 3/4 in. test-tubes
with 4% agar in w-ater. After drying at 37", they received one
or two coats of thick collodion, and a narrow paper- scale was
fixed inside near the middle, and a short length of thin tubing
with a thread attachment near the top. The separation of the
capsule from the tube was effected by filling the tube with water
and slowly raising it to near the boiling-point, when the collodion
separated easily from the glass. The capsules were washed in
changes of water, steamed to get rid of all traces of alcohol, and
finally inserted in wide test-tubes, which were plugged. The
medium was added, 20 c.c. in the outer tube, and 5 c.c. in the
capsule, and the whole was sterilised. The outer liquid was
BY R. GREIG-SMITH. 147
seeded with soil-organisms and incubated for some days, when a
suspension of the test-organism, Bac. prodiyiosus, was added to
the capsule. After incubation at 22° for a day or two, the cells
were distributed, either by blowing air through the culture or
by repeatedly drawing up and blowing out the liquid in a pipette.
Then 1 c.c. was abstracted, and a count of the bacteria made.
Previous to the mixing, the height of the liquid on the paper-
scale was read, and afterwards the volume was determined by
water run from a burette to the same mark.
The soil-organisms were obtained from a garden-soil that had
been kept some time in the laboratory. The numbers of bacteria,
originally high, as the soil had been heated, had fallen to that
normal for the soil, viz., to 6 to 8 millions per gram. Plates of
Lipman-Brown agar were smeared with the soil-suspension in
appropriate dilution, and, after several days, it was noted that
the flora consisted roughly of 30% of bacteria, and 70% of Actino-
myces-forms. Of the latter, 17% were Actino. chromogena^ which
darkened nutrient agar strongly; and 23% were Actino. odorifera,
which darkened the agar slightly; the remaining 30% were
indefinite. Many colonies were picked from the plates, the
micro-organisms were classified or grouped, and representatives
of the groups used for infecting the liquid outside the collodion-
capsules in the tubes.
The experimental results were noted as the progeny of one
test bacterial cell originally added to the collodion-capsule, but
as nothing will be gained by giving the exact numbers, the
general indications, as compared with control-tests obtained from
the figures in the various experiments, are here noted.
Experiment ii. Soil-extract [soil 1, water 2 parts].
Soil-bacteria, 6 days at 18°; test-bacteria, 1 and 7 days at 22".
Inhibiting strongly — Rhizobium from soil, Bac.A7.
Inhibiting slightly — Various (5) Actinomyces-forms, Bac. A5.
Accelerating slightly — Aspergillus sp., Bac. A2,
Accelerating strongly — Bac. Al.
A.S an indication of the value of the terms, it may be said
148 CONTRIBUTIONS TO OUR KNOWLEDGE OF SOIL-lJ-ERTlLITY, XVI.,
that the control-test showed that one cell had, in 7 days, in-
creased to 118,000; the Rhizobium had multiplied only to 100;
the Actino. -forms averaged 53,000, Aspergillus 240,000, and
Bac. Al had a progeny of 7 •'28 millions.
Experiment iii. Gum-acacia, 0"2%.
Soil-bacteria, 12 days at 18"; test bacteria, 5 days at 22°.
Inhibiting slightly — Aajjeryillus sp., Penicillium sp., Actino. chrom.
Tnditferent — Actino. odor.^ Bac. Al, Bac. A5.
Accelerating strongly — Bac. A17.
Experiment iv. Hay-infusion, 0*05%.
Soil-bacteria, 11 days at 16°; test-bacteria, 1 day at 16°.
Inhibiting — Bac. B4.
Inhibiting slightly — Bac. B5.
Indifferent — Bac. B2, grey and white Actinomycetes.
Accelerating — Bac. B8.
Experiment v. Hay-infusion, 0"05%.
Soil-bacteria, 11 days at 16"; test-bacteria, 2 days at 15°.
Inhibiting strongly — Bhizobium (from soil).
Inhibiting moderately — Bac. Al, Bac. A2.
Inhibiting slightly — Various Actinomycetes, Bac. A5, Bac. A7,
Bac. A9.
Experiment vi. Albumen, 0-1% in tube, water in capsule
(albumen coagulated upon steaming).
Soil-bacteria, 7 days at 15^; test-bacteria, 1 day at 22".
Inhibiting strongly — Rhizobium (from soil), Bac. A7.
Inhibiting slightly—Bac.T. P. 2, Bac.B4, Bac.C2, Bac.C3, Bac.Cl.
Indifferent — Bac. A17.
Experiment vii. Soil-extract [soil 1, water 1 part].
Soil-bacteria, 11 days at 15""; test-bacteria, 1 and 3 days at 22°.
Inhibiting strongly — Rhizobium (from the Lupin), Bac. A7.
Inhibiting moderately — Bac. C3.
Inhibiting slightly— Bac. T.B.2, Bac. B8.
Indifferent— Bac. Al, Bac.A17, Bac.C2.
BY R. GtlElG-SMITH.
149
Experiment viii. Hay-infusion, 0-1%, with dextrose 0-1%.
Soil-bacteria, 12 days at 16"; test-bacteria, 2 days at 22".
Inhibiting moderately — Penic. dadosporioides.
Inhibiting sHghtly— Actino. 11, Bac. A8, Bac.T.P..^, T.P.I, T.F.5,
T.P.6, T.P.7,S.B.2, S.B.5.
Indifferent— Actino. 10, Bac. S.B.I, S.B.I.
The results of the experiments, as a whole, showed that the
action of the soil-microbe is generally irregular. This is notably
the case with Bac. A 17, which is the same as Bac. Al. In some
cases, it increased the growth of the test-organism, while, in
others, it had an indifferent action. In the case of Rhizobium
and of Bac. AT, we have bacteria which produce a luxuriant slime
in media containing sugar, and even in those containing merely
a trace of sugar or none at all, the inhibiting property is seen to
remain. I cannot but think that they acted by extracting the
nutrient from within the capsule in order to produce their slime,
which remained in the outer tube. The inhibition is, therefore,
in all probability, due to the absence of nutrients rather than to
tlie presence of toxins in the collodion-capsule. As the experi-
ments did not promise to aid the investigation, they were dis-
continued.
The Use of vakious Media.
The action of Bhizobium led to its being tested in Hasks of
various media, and the tests are grouped together in the follow-
ing Table.
150 CONTRIBUTIONS TO OUR KNOWLEDGE OF SOIL-FERTILITY, xvi.,
Experiment ix.
Soil-orcjanism
Rhizobium.
Derivation
Soil.
Soil.
Lupin-nodule.
Lupin-nodule.
Medium
Hay-infusion.
0-1%.
Gum-acacia,
0-2%.
Dried blood,
0-4%.
Ammonium
sulphate,
0-1% with salts
Duration of test
3 days.
16 days.
15°
6 days.
11 days.
Temp, of incubation
16°
15°
16°
Number of test
1
2
3
4
Extract, boiled
Extract, raw
Extract, raw, 80% ...
Extract, raw, 60% ...
Extract, raw, 40% ...
Extract, raw, 20% ...
Water-control
Extract/water ratio...
1,684
100
170
106
80
175
58
1-7
343
100
99
69
63
44
23
4-3
325
100
106
96
67
49
15
6-5
41
100
52
15
5
18
22
4-6
There are indications of the formation of toxin in some of these
tests, although it must be said that they are not very pronounced.
Test 1 shows a narrow extract/water ratio, pointing either to the
possible absence of nutrients in the extract or to the presence of
toxins. The increases obtained in the boiled and in the diluted
extract lead one to believe that toxins were present. Test 2 had
undoubtedly been incubated for too long, but there is a suspicion
that the extract is of the same nature as test 1. Test No. 4 is
irregular, and, so far as we can judge, gives us no information.
In these tests, the growing organism had undoubtedly removed
nutrients from solution, and, by utilising them, had prevented
their appearance in the porcelain filtered extract. This was
shown by an extension of test 1 , in which the raw, uninoculated,
filtered hay-infusion gave 666,800 cells, and the same, when
boiled, gave 379,600 cells as against 100 of the unboiled bacterial
extract.
A mixed culture of bacteria, existing as an actively nitrifying
suspension of soil-bacteria, was seeded into a solution of 0*1%
dextrose in 0-1% hay-infusion. The suspension was incubated
at 15° for 7 days. On preparing and testing the extract, it was
BY R. GREIG-SMITH.
151
found to give the boiling- and dilution-reactions. The original
mixture of bacteria was plated out in levulose ammonium-sulphate
agar, and the flora was seen to consist of five organisms named
provisionally 8. B.1-5. These were tested with the same hay- and
dextrose solution, but as they showed no sign of toxin-formation,
it is unnecessary to give the details. The original experiment
was repeated, but the result was quite different from the first.
Experiment x.
Soil-organisms
Mixed bacteria.
Medium
Hay-infusion, 0'1% with dextrose 0'1%.
Duration of test
7 days.
12 days at 15°.
Extract, boiled
Extract, raw
Extract, raw, 80%
Extract, raw, 60%
p]xtract, raw, 40%
P^xtract, raw, 20%
Water-control
Extract/water ratio
917
100
176
230
615
465
28
3-5
96
100
92
59
31
7
1
472
The differences between these tests is marked, and, while there
probably was a difference in the kinds of bacteria added in the
two cases, there is the suggestion that the time of incubation
may have an influence upon the result.* In the 12-days' test, the
toxin may have been formed on or about the 7th day, and have
'■■" The experiment was repeated some months afterwards, with the fol-
lowing results : —
Soil-organisms.
Mixed bacteria.
Duration of test.
2
7 days.
Extract, boiled
Extract, raw
Extract, raw, 80% ...
Extract, raw, 60% ...
Extract, raw, 40% ...
Extract, raw, 20% ...
Water-control
Extract/ water ratio...
188
100
32
8
]
1
1
100
165
100
48
40
34
26
9
1-2
The previous experiment was not confirmed, and there is no evidence of
toxicity shown by the dilution-curves.
152 CONTRIBUTIONS TO OUR KNOWLEDGE OF SOIL-FERTILITY, XVI.,
become decayed by the l'2th It appeared advisable, therefore,
to test the solutions, from time to time, to see the influence of
the time-fa(;tor. Tliis was done in the following experiment, in
which Bac. T.P.2 was used at 16° to 1 8\
Experiment xi.
Soil-organism
Bac. T.P.2.
Medium
Hay-infusion, 0'1%.
Haj^-infusion and
dextrose, 0*1%.
Duration of test in days
3
5
r \ 11
3
5
11*
Extract, boiled
Ratio of boiled extracts
450
12-8
100
90
48
31
16
5
18
308
8-8
70 35
2 1
240
1
560
2-3
2,092
8-7
Extract, raw
Extract, raw, 80%
Extract, raw, 60%
Extract, raw, 40%
Extract, raw, 20%
Water-control
Extract/water ratio . . .
100
80
107
111
82
56
1-8
100 100
122 99
118 76
.159 63
111 58
122 32
0-8 3-1
100
55
32
17
8
1
113
100
36
24
19
7
9
11-5
100
84
107
123
73
192
0-5
" The numbers in these are irregular, largely owing to the counts being
low.
During the filtration of the infusions through the porcelain, it
was noted that, in the dextrose-media, a considerable amount of
slime had been formed. Attention was not called to this in the
three days' culture, but, in the five and eleven days' culture, the
slime was pronounced. It is clear from the results that the
organism had produced a certain amount of toxin about the
sixth day in the simple hay-infusion, and that it had largely
disappeared by the eleventh day. The ratio of growth, which
is a rough index of the approximate amounts of nutrients in the
extract, declined as time went on, and so did the effect of boiling
the solution. In the hay- and dextrose-solutions, boiling showed
a steady increase of nutrient produced as the age of the culture
increased, while, without dextrose, the reverse was the case.
The organism T.P.2 was originally obtained upon an agar-plate
which had been seeded with a suspension of soil-bacteria. It
was conspicuous in producing a zone across which neither moulds
nor spreading bacteria would go. The single colony upon puri-
fication was found to consist of two closely allied forms pro-
BY R. GREIG-SMITH.
153
visionally named T.P. 1 and T.P.2, the former producing a white
colony, the latter an ivory-white on nutrient agar. T.P.I lique-
fies gelatine quickly and foims no slime on dextrose- media, while
Bac. T.P, 2 liquefies gelatine slowly, and forms a slime from
dextrose. The belief was raised by other experiments that the
formation of slime withdraws nutrients from the medium, and
yields a poor extract; but whether the toxin-formation has any-
thing to do with the production of slime, has yet to be deter-
mined. Bac. 'J\P.l, the slime-free ally of Bac. T.P.2, was tested
under the conditions of the last experiment, viz., in hay-infusion,
with and without dextrose, and at periods of 3, 6, and 10 days,
but no evidence of toxin-formation was obtained.
An attempt was made to determine the effect of slime-forma-
tion upon the production of toxin by using a race of Rhizobium
from the soil, which formed a luxuriant slime on solid dextrose-
media.
Experiment xii.
Medium.
Hay-
infusion.
0-1%.
Hay-infusion with
dextrose, 0-1%.
Soil-organism.
Rhizobium .
Rhizobium .
Duration of test in days.
3
6
13
3
6
Extract, boiled ...
Extract, raw
Extract, raw, 80%
Extract, raw, 60%
Extract, raw, 40%
Extract, raw, 20%
Water-control ...
Extract/water ratio
82
100
13
1
1
1
1
1,900
5
100
10
5
4
3
9
49
205
100
84
77
58
50
35
2-8
20,840
100
3
2
2
2
1
72
45,030
100
40
36
49
47
37
3
A 13-days' test with hay-infusion and dextrose was found to
be sterile, and has been omitted. Although these tests were
intended to see the effect of slime-production upon toxin- forma-
tion, it happened that no slime was obtained on the porcelain
filter from the dextrose solution, and, but for the growths on
agar-slopes, one would have thought that the cultures were dead.
However, the slow reduction of the extract/ water ratio as time
went on, and the great increase obtained on boiling the dextrose-
solutions were again noted.
154 CONTRIBUTIONS TO OUR KNOWLEDGE OF SOIL-FERTILITY, xvi.
1 have found, in the past, that soils are more toxic in the
winter than in the summer-months, and it is to be expected that
temperature will have some influence in either the production or
the decay of the toxin. The influence of temperature was tested
in the following.
Experiment xiii.
Medium
Hay-infusion, 0*1%.
Soil-organism
Bac. T.P.2.
Temperature
15-5°.
22°.
Duration of test in days . . .
2
4
8
2
7
Extract, boiled ...
20
74 1 123
30
16
Extract, raw
100
100 100
100
100
Extract, raw, 80%
33
77 t 48
46
29
Extract, raw, 60%
12
64 1 19
26
12
Extract, raw, 40%
3
45 '' 12
1 /
7
Extract, raw, 20%
1
45 , 10
13
5
Water-control ...
1
53 11
6
5
Extract/water ratio
248
2 9
15
18
The experiment was rather disappointing, as there was little
evidence of toxin-formation at the lower or the higher tempera-
ture. In four days at 15'5° it is shown by the low extract/water
ratio that much of the nutriment has been locked up in the bodies
of the bacteria. As the ratio is higher on the eighth day, one
would imagine that there had been some dissolution of the cells.
In an earlier experiment, there had been signs that some
formation of toxin had occurred in a solution of gum-arabic
infected with Bac. '1\P.2. This led to the employment of an old
culture of Rhizobium which had been grown in hay-infusion and
dextrose, and which contained a quantity of slime. It was
sterilised and infected with Bac. T.P.2. It showed no signs of
toxicity on the third day.
An old culture of Bac. T.P.2 in hay-infusion was sterilised and
infected with Bac. T.P.2, but there were no signs of toxicity on
the second or fourth day.
A solution of Gum-Acacia, 0-2%, when infected with Bac.
T.B.2, showed no toxicity on the second or fourth day.
BY R. GREIG-SMITH. 155
A rod-sliaped bacillus, Bac. A17, which had appeared to
stimulate the growth of Bac. prodigiosus in collodion-capsules,
was tested in hay-infusion, but there was no evidence of toxin
formation in the boiled or diluted extracts.
In view of the indeterminate nature of the results hitherto
obtained, it was deemed advisable to augment the intensity of
the growth of the bacteria by increasing the amounts of the
nutrients and by aerating the media during cultivation. To gain
this end, beakers containing cotton-wool were sterilised, and
media, such as 1% hay-infusion with and without 1% dextrose,
which had been seeded with various organisms, were added in
sufficient quantity to moisten the wool. After incubation at
22°, the cotton-wool was squeezed and washed, and the liquids
made up to a definite volume before being filtered through
porcelain.
A soil-Rhizobium, Bac. T.P.2, Penicillium cladosporioides, and
an Actinomyces, A 10, were tested at difierent times, such as 3,
6, and 11 days; but, in none of the tests, was any evidence of
toxin-formation demonstrated. This also applied to old cultures
of bacteria reinfected with moulds. The method was useless,
therefore, for the object in view.
The experiments, however, raised the idea that small changes
in the reaction might be responsible for much of the irregularity
in the results. Bacterial cultures, in the absence of a ferment-
able sugar, are generally more or less alkaline, while soil-extracts
are supposed to be more or less acid. The soils used in this set
of researches were acid, that is to say, when a piece of litmus-
paper was inserted or pressed into a paste or porridge made by
mixing the soil with water, it became reddened in the course of
a few minutes. The aqueous extract of such soils should, there-
fore, be acid, and it appeared that, if the extract could be pre-
vented from becoming alkaline through the action of bacteria,
there would be a better chance of demonstrating any development
of toxicity. At any rate, the cultural fluids would be more in
harmony with the conditions that pertain in the soil. To efiect
this faint, permanent acidity, it would be necessary to add some
substance which would absorb any alkalinity produced, in the
156 CONTRIBUTIONS TO OUR KNOWLEDGE OF SOIL FERTILITY, xvi.,
same manner that chalk maintains a faint aciditv in feimenting
solutions of sugar. The only substance that promised to achieve
this purpose was humic acid, and its use appeared advisable, as
it is ever present in soils.
A quantity of humic acid was accoi-dingly prepared from
rotted bamboo-mould by treatment with sodium h\'drate, followed
by acidification, washing with water by decantation, filtration,
and finally by dialysis. The acids were dissolved in normal soda
and sterilised.
Tests were made with strung hay-infusion with and without
dextrose by the cotton-wool method, but, although several
organisms were used, no results of any importance were obtained:
the solutions were far too nutritive. A converse test with tap-
water and no nutrient showed that the humic acid acted as a
poison. Although an opalescent suspension of bacteria had been
added, no living bacteria were to be found by the third day. In
one case, a pink yeast had obtained access to the fiask, and, on
the thirteenth day, the extract showed a rise upon boiling; and,
upon dilution with dilute hay-infusion, the same sterility,
towards the introduced bacteria, was noted after a few days.
The humic acid was undoul)tedly the cause of the plienomenon.
It had been dissolved in normal soda, and the solution was
added to the flask of medium, and was followed by the quantity
of normal hydrochloric acid necessary to neutralise the soda.
The humic acid remained largely dissolved, and acted as a soluble
acid. In one test, that of the pink yeast on the thirteentii day,
the acidity of the extract to phenolphthalein was = -t-0"5' by
Fuller's scale, that is the extract contained Go c.c. of normal
acid per litre. The experiments suggest that, to beefiective, the
humic acid should be insoluble in water.
Experiments with Humus.
As the toxin-formation in soil must be related to the changes
in the organic matter, an attempt was made to see in how far
humus would be useful in elucidating the problem. Rotted
bamboo-grass was sifted, partly dried, mixed and sifted again to
obtain a uniform mould. Portions weighing 10 grams were put
into deep Petri-dishes, and sterilised by heating for two hours at
BY R. GREIG-SMITH.
157
130". When cold, lOc.c of a suspension of Bac. T.P.2 were
added to each portion. After various periods of incubation at
18**, a portion was transferred to a shaking bottle, treated with
500 c.c. of distilled water, and shaken 50 times at 10-minute
intervals for an hour. The suspension was then filtered through
paper and porcelain, and the usual procedure followed.
Experiment xiv.a.
Soil-organism
Bac. T.P.2.
Duration of test
3 days.
3,870
100
50
13
9
5
1
150
6 days.
Extract, boiled
Extract, raw
Extract, raw, 80% ...
Extract, raw, 60% ...
Extract, raw, 40% ...
Extract, raw, 20% ...
Water control
Extract/water ratio...
210
100
79
50
21
4
2
52
As the experiment did not seem to be going to give any useful
information, the remaining portions were extracted with different
quantities of water to test the influence of various strengths of
extract.
Experiment xiv.b.
Growth of Bac. prodigiosus in extracts of leaf-
mould infected with Bac. T.P.2, and incubated
for 11 days at 18°.
10
gram
portion
to water.
500 CO.
300 c.c.
100 c.c.
100
100
100
35
240
487
5
276
1,563
23
3
46
4
36
2
Extract, raw
Extract, raw, 80%
Extract, raw, 60%
Water-control
Extract/water ratio
The results of Experiment xiv.a, are much the same as have
been obtained in solutions, viz., a lessening of the nutritive and
of the boiling effects, as the period of incubation proceeded. In
Experiment xiv.b, we have the dilution-effect becoming more pro-
nounced as the water used in making the extract became less,
12
158 CONTItlBUTIONS TO OUR KNOWLEDGE OF SOIL-FERTILJTY, xvi.,
Portions of the air-dried mould containing 14 grams of dry
matter were put into Petri-dishes and sterilised. They were
infected with cultures of Bac. T.P.2, and of Actinomyces 10, and
the moisture raised to 40%. The extracts were prepared by
shaking each portion with 500 c.c of distilled water.
Experiment xv
Soil-organism
Bac. T.P.2.
Actino. 10.
Control.
Duration of test in days
4
6
18
24t
10*
18t
24t
Extract, boiled
44
164
119
47
71
90
138
Extract, raw
100
100
109
100
100
100
100
Extract, raw, 80% ...
849
532
270
66
292
171
119
Extract, raw, 60% ...
3,710
1,880
1,845
54
574
268 1 68
Extract, raw, 40% . . .
9,500
3,090
6,637
37
1,038
160 j 57
Extract, raw, 20% ...
8,070
2,500
16,090
17
562
o
25
Water-control
47
20
153
1
2
1
1
Extract/water ratio . . .
2
5
0-6
1,130
44
275 460
Reaction
+ 0-6
-I-0-6
+ 0-55
+ 0-3
—
-i-0-4 +0-2
* Growth of a green
Penicillium at edge of dish.
tPeri
neated
with P(
snicillii
im.
The growth of the accidentally introduced Penicillium resulted
in a flattening of the dilution-curves, and in a diminution of the
acidity. From this, it must be inferred that the acidity was
chiefly responsible for the toxicity of the extracts. To test the
matter, a further experiment was made with partially neutralised
mould, and with neutralised extracts.
Four-punce glass bottles were used instead of Petri-dishes, and,
into each, 18-2 grams of mould, containing 14 grams of dry
matter, were introduced. They were sterilised at 130° for two
hours, moistened with 3 c.c. of sterile water, and steamed for an
hour. The steaming proved to be unnecessary, as portions of
the heated soil proved to be sterile. One of the portions was
infected with a Fusarium, another with a Rhizobium, a third
served as a control, a fourth was treated with enough lime-water
to neutralise the apparent acidity, and the last was subjected to
the vapour of ammonia for a couple of hours. The moisture in
them all was brought to 40%. The extract of the raw mould
was found to be very acid, 1000 c.c. containing the equivalent of
BY R. GRKIG SMITH.
59
one c.c. of normal acid, using phenolpbthalein as an indicator.
The extracts were neutralised with sodium bicarbonate. The
Fusarium, the Rhizobium, and the control-tests were incubated
for five davs at 22°.
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The point to be noted from this set of tests is, that the neu-
tralisation of the acidity of the extract has converted the toxic
160 CONTRIBUTIONS TO OU K KNOWLEDGE OF SOIL-FERTILITY, xvi.,
into a nutritive condition, not only in the raw extract, but also
in the 80% dilution. The numbers obtained in the extracts of
the control and of the infected leaf-moulds are of the same order,
and indicate that the toxic property is inherent in the sterilised
medium. The suspicion is strengthened, that the toxins are
really acids developed during the sterilisation of the organic
matter. It is evident that the treatment of the leaf-mould with
lime-water or with ammonia was not suflacient to neutralise the
excessive acidity of the humus, for the numbers show a position
intermediate between the acid and the fully neutralised extracts.
In a new set of tests, the sifted mould was treated with water
to eliminate the sand with which it was mixed. It was then
dried, and again sifted through a finer sieve to remove the bulk
of the fine, light fibres. It contained moisture 10-2%, and ash
43-8%, leaving 46% for the organic matter. Portions containing
10 grams of organic matter were put into 4-ounce bottles, and
sterilised at 130° for two hours. During the sterilisation, an
odour of burnt sugar was noticeable.
A portion of the sterilised leaf-mould was tested against a
portion of the unsterilised for acidity. Each was shaken 300
times with 500 c.c of water, and filtered. This was repeated a
third time. The acidity of the filtrates was tested with N/lOO
soda, using phenolphthalein as an indicator. The figures repre-
sent the number of c.c. required to neutralise the 500 c.c. of the
extract.
mould, heated.
mould,
not heated.
difierence.
1st 500 c.c.
2nd 500 c.c.
3rd 500 c.c.
47-5
16-25
10-0
7-5
5-0
375
40-0
11-25
6-25
The curves of these numbers were plotted, and it was seen that
the curve of the heated mould would meet that of the unheated
mould at the fifth 500 c.c, and that, at the fourth, there would
BY R. GREiG-SMITtt.
16;
be a difference of 3 c.c. The total differences would, therefore,
amount to 60 5 c.c. of N/100 acid for the total acidity developed
during sterilisation and removable by washittg ivitk icater. In
terms of the organic matter of the mould, this means that 100
grams during sterilisation developed an acidity equal to 605 c c.
of normal acid, equivalent to 0-36% grams of acetic acid.
A portion of the sterilised leaf-mould, containing 10 grams of
organic matter, upon being gradually moistened with water, was
found to form a fairly coherent ball when pressed in the hand
after the addition of 1 5 c.c. of water to each portion. Accord-
ingly, 15 c.c. of water containing 6 c.c. of N/10 sodium bicar-
bonate were added to each portion, which was thoroughly mixed
and pressed down. Suspensions of the bacteria in 1 c.c. of water
were subsequently added, mixed, and pressed.
After an incubation of five days, extracts were made, and
their effect upon the growth of Bac. j^vodigiosus noted.
Experiment xvii.
Soil-organism...
Bac. T.P.2.
Rhizobium. None.
i
Reaction of extract
acid, neutral.
acid.
neutral, acid.
neutral.
Extract, boiled
Extract, raw ...
Extract, raw, 80%
Extract, raw, 20%
Water-control
Extract/water ratio
106
100
132
1,677
1,471
0-07
77,930
70,580
4,390
60
100
281
5,342
200
0-5
41,960
35,420
12,740
152
100
108
1,000
1,316
0 07
85,800
89,310
9,950
I
The numbers in the control and in the T.P.2 tests are virtu-
ally the same, and evidently no growth of the bacillus had
occurred. The numbers with Rhizobium are different, not only
in the acid extract, but also after it had been neutralised with
soda. The fact that Rhizobium can grow in the partially acid
leaf-mould is a point worth noting. It is needless to discuss the
possible formation of toxins with these leaf-moulds, and the re-
162 CONTRIBUTIONS TO OUR KNOWLEDGE OF SOIL-PERTILITY, xvi.,
maining portions of the tests were discarded until further work
had been done.
The addition of the bicarbonate of soda to the portions of leaf-
mould did not neutralise the soluble acidity, for when the raw
extracts were tested, it was found that the T.P.2 test had an
acidit}^ equal to + 0 62° in the extract. The control had +0-63'',
and lihizobium +0-35". It is, perhaps, to be expected that, in
a substance like rotted leaf-mould, there will be a mixture of
humic acids, some soluble in water, some partially soluble, and
some insoluble in water. The humates are so complex that it
was a mistake to consider that even all the water-soluble acid
would have been obtained by water-extraction, unless the time-
factor had been taken into account, and days instead of hours
been occupied in the extraction.
The necessity for having a neutral vegetable-mould was
emphasised, especially in the last two tests, and some experiments
were made with the object of getting a better idea of the true
acidity.
The Acidity of heated Leaf-mould.
A number of portions of the leaf-mould, each weighing four
grams, were bottled, and some were heated for two hours at 130°.
A sterile and an untreated portion were repeatedly shaken up
with a standard alkaline solution at intervals during the time
of contact, and filtered. The filtrates were tested for residual
alkalinity or, when neutral liquids were used, for acidity. 'J'he
numbers represent cubic centimetres of normal acid in, or derived
from, 100 grams of the dry, organic matter of leaf-mould.
Calcium bicarbonate. — The acidity was first tested by the
method of Hutchinson and MacLennan,* which consists in
having the portions in contact with a solution of bicarbonate of
calcium for a certain time, and subsequently determining the
amount of lime that had been removed from solution
Heated 93-3
Control 91-7
* Journ. Agric. Sci., vii., 75.
BY R. GREIG-SMltH.
163
Sodium bicarbonate. — Portions were moistened with 10 c.c. of
alcohol, and treated with 100 c.c. of water containing 1 gram of
bicarbonate of soda, for five days.
Heated 165
Control 121
Other portions were moistened with 5 c.c. of alcohol, and
treated with 200 c.c. of water containing 0 8 grams of bicarbonate.
Heated 82-4
Control ... 75-5
Alcohol. — Portions were shaken up with 200 c.c. of neutral
spirit and allowed to stand overnight.
Heated 9-4
Control 4-0
Water, — Portions were shaken up with 500 c.c. of water and
allowed to stand overnight.
Heated 241
Control 9-2
Liine-24jater.—V ovtiow^ were treated with 200 c.c. of N/21
lime-water and allowed to stand overnight.
Heated 383
Control 362
The numbers obtained with lime-water were so high, that the
experiment was repeated. The four grams of leaf-mould were
shaken with 300 c.c. of approximately N/24 lime-water, and due
allowance was made for the amounts removed in the portions of
fluid abstracted.
Normal alkali, in c.c, absorbed by 100 grams of dry organic
matter.
Days.
1/8
1
2
3
4
5
7
9
Heated
Control
340
35o
403
395
423
410
432
418
442
426
448
432
4o«
440
4(32
444
This confirms the previous test in showing that a compara-
tively large amount of lime is removed from solution, and that
164 CONTRIBUTIONS TO OUR KNOWLKDGE OP SOIL-FERTILITY, xvi.,
more is absorbed by the sterilised than by the unsterilised mould.
The organic matter has absorbed from 12 to 13% lime [CaO], and
about 90% of the total was absorbed within two days.
The difference in the amount of base absorbed from the
hydrate, as against the bicarbonate, led to a test being made
with bicarl)onate of magnesia. A quantity of freshly precipitated
and waslied carbonate was suspended in water, and a current of
carbon dioxide was passed through for several hours. The solu-
tion was filtered, and 200 c.c were added to 4 grams of soil in
stoppered bottles. The solution of bicarbonate of magnesia was
approximately twelfth normal The bottles were shaken fre-
quently, and the portions abstracted from day to day and boiled
with an excess of N/20 sulphuric acid for ten minutes, and
titrated with ]S'/20 soda in presence of phenolphthalein.
Magnesium bicarbonate'— l^ovmal alkali, in c.c, absorbed by
100 grams of dry organic matter.
Days. 3
5
7
10
Heated...
Control...
99
89
99 105
89 99
104
99
The numbers are closely akin to those obtained with calcium
bicarbonate, and much under the tests with lime-water.
Baryta-water. — The action of lime-water was controlled by a
test made with 'baryta- water, in which 300 c.c. of approximately
N/15 alkali were added to each 4-gram-portion of mould.
Normal alkali, in c.c , absorbed by 100 grams of dry organic
matter.
Days.
1 2
3
5
6
7
8
9
12
Heated...
Control ...
389 408
365 , 382
419
391
434
400
442
405
445
417
448
418
414
425
464
433
These numbers run closely with those of the lime-water test
BY R. GREIG-SMITH. 165
The set of experiments show that vegetable-mould, oiiginally
of ail acid reaction, is made more acid by the action of beat,
such as by sterilisation for two hours at 130°. The acid sub-
stances are partly soluble in alcohol and in water. They consist
of two kinds, one of which is capable of being neutralised by the
bicarbonates of the alkaline earths. The other makes itself
evident when in contact with the hydrates of the earths. The
mould which was examined removed from four to five times more
alkali from the hydrate than from the bicarbonate.
At the close of the baryta- water experiment, the control-soil
was rapidly filtered on the pump, washed with a small quantity
of water, transferred back to the bottle and shaken with 300 c.c.
of water, and tested from time to time.
3 hours ... ... . . 26*5
1 day ... ... ... ... 36
2 days 36
Of the 433 c.c. removed from 100 grams of dry, organic matter^
36 were given up to water, and we may, therefore, consider the
difference of roughly 400 c.c. as being in combination.
The sterilised leaf-mould was also filtered and shaken with 200
c.c. of twentieth-normal hydrochloric acid. This removed the
following amounts of normal lime.
3 hours ... ... ... 328
1 day ... ... ... ... 343
2 days 343
Deducting this from the total baryta absorbed, viz., 464 c.c,
we have 120 c.c, which is close to the amount absorbed from the
bicarbonates. It is evident that the kind of humic acid, which
fixes the bulk of the lime from a solution of the hydrate, forms
with it a feeble combination. This is destroyed by mineral acids
but is not attected to any extent by water. The combination
seems to be too detinite in its quantitative relations for a simple
case of adsorption.
From the experiments, it was concluded that the organic
matter of the leaf- mould contained humic acids, and, of these,
about one-quarter were capable of decomposing the bicarbonates
of the earths, while three-fourths were too weak to do this but
166 CONTRIUUTIONS TO OUR KNOWLEDGE OF SOIL-FERTILITY, xvi.,
could combine with the earthy hydrates. Heating the mould
increased the amount of acid, and the increase was largely soluble
in water.
Humus with Calcium Carbonate.
According to the earlier lime-water test, in which 100 grams
of dry organic matter absorbed 383 c.c. of N/1 alkali from lime-
water, it appeared that a neutralisation of the acids of the leaf-
mould would be obtained by adding 2 grams of calcium carbonate
to each portion containing 10 grams of dry organic matter, and
that neutralisation would be certain if 2*5 grams were added.
A set of bottles were prepared, each receiving the equivalent of
10 grams of dry organic matter, and 2*5 grams of calcium car-
bonate as precipitated chalk. The bottles were sterilised for
two hours at 130", then moistened and stirred with 15 c.c. of
water, and steamed for an hour. Subsequent tests showed that
sterilisation had been complete, and that the extracts furnished
by the tests were neutral to phenolphthalein. Sets were infected
with certain micro-organisms and incubated at laboratory tem-
peratures for varying times. It will be remembered that the
organisms T.P.2 and Actino. 14 were chosen because they in-
hibited the growth of spreading bacteria and moulds on agar-
plates. A test made at the beginning of this experiment showed
that T.P.2 had lost its toxic power, while Actino. 14 had not.
In view of this, the bottles of T.P.2, which had been infected,
were incubated in an atmosphere of carbon dioxide, to see if
this would have any influence in restoring the toxicity. Several
of the portions, which were tested on the eighteenth day, were
unfortunately destroyed.
BY I{. GREIG-SMlTH.
167
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168 CONTRIBUTIONS TO OUR KNOWLEDGE OF SOIL-FERTILITY, xvi,,
A general glance over the results leads to the belief that no
toxins are formed by the organisms. It may be that the excess
of carbonate of lime has brought about a condition in which the
preservation of the toxins is not possible. The outstanding
feature of the experiment is the increased nutritive effect ob-
tained with Fenicilliu'tit dados par ioides. The extract/water
ratio increased very rapidly, indicating that the mould had been
actively attacking the organic matter and producing substances
which stimulated the test-organism, Bac prodigiosvs. to an in-
creased production.
The amount of unacted-upon carbonate in the treated leaf-
mould led to the belief that too much had been added, and that
an excess of carbonate was not desirable.
Humus neutralised with Limk-water.
A quantity of sifted leaf-mould was soaked in lime-water for
an hour, 5 litres of lime-water being used for every 500 grams of
leaf-mould. A trial test showed that this proportion furnished
a neutral extract, although, when the bulk was prepared, the
extract of the mould was faintly ncid. This was possibly caused
bv the subsequent sterilisation. iVt the end of an hour, the sus-
pension was stirred and decanted on to a filter, and the organic
matter washed, and at the same time separated from the heavier
sand b}' repeating the procedure with water. The residual
organic matter was dried, sifted, and bottled.
At this stage, the mould contained moisture 40" 42%, asli
18-80%, and organic matter 40'48%, so that 24-5 grams contained
10 grams of organic matter. This quantity was put up into a
number of four-ounce bottles, which were heated for two hours
at 130°. Most of the moisture was driven off by the heating. A
test showed that the heated mould formed a coherent mass, upon
being pressed in the hand, when a weighed portion contained 20
grams of water, and this was accepted as being the quantity
requisite for a full bacterial growth. After the heating, the
water was raised to 19c.c., the remaining 1 c.c. being left for
the infecting suspension. The water was thoroughly incorpo-
rated, and the bottles were steamed for an hour. Sets were
infected with several micro-organisms and kept at room-temper-
ature, 15°.
BY R. GREIG-SMITH.
169
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1 70 CONTRIBUTIONS TO OUR KNOWLEDGE OF SOIL-FERTILITY, xvi.,
The behaviour of Pen. cladosporioiaes was different in tViis ex-
periment from that in the presence of an excess of chalk. It did
not decompose the organic matter so quick!}', but the rise in the
dilution-curve was more pronounced. This suggested the idea
that, if the nutritive effect could be kept down, the curve might
be more accentuated Use was made of Rhizobium for growing
with the Penicillium because, when the former was tested at the
same time, viz., on the tenth day, the nutritive ratio was lower
than either the latter or the control-test. The double growth
kept down the ratio, but there was no elevation of the dilution-
curve. The growing Penicillium brought about an alteration in
the reaction of the extract. The reaction of the control-test
seems to indicate, that the bacteria and moulds cause the medium
to furnish acid extracts which, in time, may become alkaline.
The effect of using different amounts of water in preparing the
extract was shown in the case of the mould infected with Fusa-
rium species. Two tests were extracted at the same time, one
with 500 c.c, the other with 250 c.c. of water. The stronger
extract gave the higher nutritive ratio, as 204 : 42, while the
weaker solution showed a tendency to produce a rise in the
dilution-curve.
On the whole, the neutralised leaf-mould did not come up to
expectations in serving as a suitable medium for the demonstra-
tion of toxin-production by the selected micro-organisms.
When an extract is diluted and subsequently sown with
bacteria, it will give a count in proportion to the amount of
dilution. But if the extract is acid, the dilution will be less
acid, and the count will not be in proportion, but will be more
or less according to the effect of the change in the acidity upon
the growing bacterium. It is difficult to say just what this
difference from the normal would be, especially with such weakly
acid extracts as in Expt. xix. The neutralisation of an extract
so strongly acid as -f TO is well seen in Expt. xvi., in which the
dilution to one-fifth raised the count from 150 to 300-fold. A
set of experiments were made to obtain some definite informa-
tion regarding the influence of slight variations in the reaction.
hy k. gre1c4-smith. 171
The Influence of Pveaction upon Bacterial Growth.
An extract of the same vegetable-mould as in Expt. xix. was
prepared, and the acidity was determined in the following
manner. Twenty-five c.c were pipetted into a beaker, an equal
volume of distilled water was added, and the beaker was covered.
It was boiled for ten minutes to expel the carbon dioxide, and
rapidly cooled. Three drops of phenolphthalein were added, and
N/100 soda was run in until a difference in the tint was manifest.
An end-point was not aimed at; just the difference in the tint
from a yellowish to a faint brown. From the reading, 0*05 c.c.
was deducted as being necessary to produce the change of tint
over the neutral point. For example, 25 c.c. of an extract gave
a tint with 015 c.c. of N/TOO soda. Deducting 0*05 c.c, the
reading becomes O'l c.c. for the neutral point. With 0-4 c.c,
the tint was a pronounced red, and the final deep purple was
obtained with 0-9 c.c. The neutral quantity 0*1 c.c, when cal-
culated to normal acid per litre, gives 0-04 c.c, which, by Fuller's
scale, is represented by -f 0'04. A good, white light is required
during the titration to observe the change of tint. The method
appears to be correct, for when the extracts were neutralised by
the findings of the method, they always gave the highest counts
after incubation with the test-bacterium.
After determining the reaction of the extract, quantities of
100th normal lactic acid or bicarbonate of soda were added to
10 c.c portions of the extract, and sterile water to bring the
volume up to 1 1 cc. One c.c. of a suspension of Bac. jyrodigiosus
was added, and this brought the volume up to 12 c.c, upon which
the quantities of acid and alkali, which were added, had been
based. The bottles were incubated at 22" for 20 hours, when
counts were made, and these were calculated in terms of the
neutral extract.
172
CONTRIBUTIONS 10 OUR KNOWLKDGE OF SOIL-FERTILITY, xvi.,
EXPEBIMENT XX.
Reaction of extract
Grow
'th of Bac. pi'odigiosus, 20 hours at 22^
(Fuller's scale).
a
b
e
average.
-0-3
56
43
53
50
-0-2
50
67
60
60
-0-1
93
71
70
78
0 0
100
100
100
100
+ 0-1
86
88
77
83
+ 0-2
33
44
21
33
+ 0-3
3
10
3
5
The numbers in the three tests are not uniform, but they serve
to show the probable variation that the effect of dilution has
upon an acid or alkaline extract. For example, in the ten days'
test with Penic. cladosj). (xix.), the numbers with 80% w^ere 187,
and with '20%, 38. The reaction-curve passes through 86 for
+ 0-1,91 for +0-08, and 99 for +0-02. A slight calculation
shows that the numbers at the 80% dilution are higher by 6%,
and at the 20% dilution higher by 15% than they should be on
account of the reduction of the acidity by the mere dilution. In
these weakly acid extracts, however, small differences such as
these are negligeable, as the counts themselves are liable to
greater fluctuations.
A stage has been reached in the investigation at which it is
made clear that bacteria and moulds do not produce toxins, or,
if they do, the toxin is not capable of being demonstrated either
in nutrient solutions or in vegetable-humus. Any resemblance
to toxicity is probably caused by an alteration in the reaction of
the medium, and, to such alterations of reaction, the test-organ-
ism is very sensitive.
The Growth of Amceb^e.
The attempt to obtain toxic substances among the byproducts
of certain bacteria and moulds had not been so successful as had
been wished ; in fact, it had been decidedly unsatisfactory.
Either the methods of producing the desired bodies were at fault,
or the micro-organisms were not such as would give the desired
results, although they had been selected as being the most likely
to do so. It is known that bacteria can give out bacteriolytic
BY R. GREIG-SMITH. 173
substances, as for example, Bac. pyocyaneus, and moulds may be
capable of doing the same. But these are not the only inhabit-
ants of the soil. There are certain protozoa, and with the failure
to obtain a decided and undoubted production from the bacteria
and moulds, the attention was turned to the soil-fauna.
With regard to the possible activity of the members of the
soil-fauna in this direction, we are faced with the fact, that the
introduction of certain species of protozoa, e.g., the Amoebse and
Colpodse into sterile soils, does not bring about the condition that
holds before sterilisation. The bacterial numbers, which become
greatly increased as a result of the sterilisation, do not become
reduced to the previous level when the protozoa are present.
It is true that Goodey* has lately shown that the inhibiting
factor, which determines the reduction of the bacterial numbers,
becomes evident when the numbers of amoebae approach and
exceed 50,000 per gram of dry soil, but, at the same time, he
says that the sterilised or disinfected soils are not suitable media
for the production of the factor. The treatment of the soil with
heat or with the volatile disinfectants so alters the soil, that it is
not immediately suitable for developing the inhibiting factor,
whatever it may be.
I had considered the possibility of soil-amceba? being able to
produce substances akin to the immune bodies of the animal
pathologist, but, as experimental work failed to reveal any sign
of toxic substances, and such immune bodies must be included
in this category, I simply record the fact that such a possibility
had been entertained, and that experiments with, and without,
"vaccines" had been negative.
Some difficulty was met with in obtaining a "pure" culture of
soil-amoebse. Upon a former occasion, they had been cultivated
in quantity in 1% liay-infusion, without any trouble, but, upon
repeating the procedure, it was found that the medium was not
well adapted for the purpose. The single cells generally disap-
peared overnight. The trouble was traced to the medium being
too nutritive, thus permitting the accompanying bacteria to
become too numerous and render the medium too alkaline. A
* Proc. Roy. Soc, 89 (B.616), p.297.
174 CONTRIBUTIONS '10 OUR KNOWLEDGE OF SOIL-FERTILITY, xvi.,
hay-infusion, in which the bacteria had grown for some time, had
changed from having a faint original acidity of -I-0-25 to an
alkalinity of —2*4. It was naturally assumed that the alka-
linity developed overnight had been the cause of the disappear-
ance of the ama3b8e, but, when either saccharose, lactose, lactic
acid, humic acid, or sodium phosphate was added, the trouble
still remained, although it was not so pronounced in the presence
of humic acid. The addition of potassium nitrate, mono- or
di-hydrogen phosphate, or ammonium phosphate to make a 005%
solution with the hay-infusion, also resulted in the disappear-
ance of the amcebse. It seemed at one time as if the smallest
droplets gave the most satisfactory growths, that one amoeba in
a small droplet had a better chance of growing than one cell in
a large droplet, and it was concluded that the hay-infusion was
too strong. It is a recognised rule, in growing amoebae, that a
poor medium should be used in order to prevent the protozoon
being overwhelmed with bacteria, but 1% hay-infusion is by no
means considered to be a rich medium, especially when made
from a rather poor sample of couch-grass. However, experiments
j showed that 0-2% hay-infusion was well adapted for growing
amoebae in mass-culture, although it produced rather delicate
forms in single-cell work. One of my most successful starter-
cultures was obtained by gradually adding 1% hay-infusion so
that the bacteria were kept under as much as possible. An
equally good starter was obtained by growing the cells in a 2%
infusion of exhausted leaf-mould, the same as was used in some
experiments about to be recorded.
During this part of the investigation, the effect of adding
sodium chloride to the hay-infusion was tested. In one instance,
the addition of 0-2% was beneficial, while a larger quantity de-
stroyed the amoebae. In another instance, the addition was
injurious. A good culture-fluid was found in tap-water contain-
ing 0-05% asparagin with 0-11% K^HPO^. Although two of
the original five cells died overnight, the remaining three cells
increased to 39 in another day. In view of this, an experiment
was made in which the asparagin was replaced by chloride,
sulphate or nitrate of ammonia, nitrate of potash and urea. A
BY R. GREIG-SMITH. 175
number of amoebse were added to each droplet, and a trace of
chalk. All the cultures did well, and, in course of time, the
mobile forms encysted. The cells of the urea-test were vegetat-
ing long after the others, and the amoebse in the ammonium
chloride test also persisted longer, and finally disappeared with-
out forming cysts.
A preliminary experiment was made with an extract of a
partially exhausted leaf-mould, using a growth of amoebae derived
from a single cell. Although the solutions were tested from
time to time in the customary manner for the influence of boiling
and dilution, no definite information was obtained, and it was
concluded that the culture-solution was too poor in nutrients to
show or develop any signs of toxicity.
Vegetable-mould neutralised with lime (p. 168) was then used.
Each test contained the equivalent of ten grams of dry organic
matter with an amount of water sufficient to cause the soil to
adhere loosely to the sides of the containing bottle. This meant
73% of moisture. It was apparent from the results that the
mould became too acid for the continued growth of the amoebae.
Forty thousand were added to each bottle, and, in five days, they
had increased to 1*6 millions, in twelve days they had decreased
to 136,000 mobile forms, and, by the twentieth day, they all had
encysted. While this was going on, the extract, originally
neutral, became more and more acid, doubtless due to the carbon
dioxide, produced by the activity of the bacteria introduced with
the amoebae, reacting with the calcium humate, and liberating
free humic acid.
The test was repeated with similar results. On the sixth day,
the amoebae had increased forty-four fold, and the reaction of the
extract was - 0-04 ; on the thirteenth day, the amoebae had
encysted, and the reaction was -i-O-l. The bacterial numbers
gave no information.
Better results were obtained with an infusion of hay made b}^
diluting a 1% infusion with tap-water to make a 0-2% solution.
Tests were made on the sixth, twentieth, and forty-second days,
but, as the two latter gave somewhat normal dilution-curves,
they are not recorded.
176 CONTRIBUTIONS TO OUR KNOWLEDGE OF SOIL-FERTILITY, xvi.
EXPERIMEJ^T xxi.
Soil-protozoon,
Amoeha Umax.
Numbers corrected
for alkalinity.
Extract, boiled
575
575
Extract, raw...
100
100
Extract, raw, 80%
387
277
Extract, raw, 50%
962
489
Extract, raw, 20%
3,252
1,205
Water-con tz^ol
5,522
1,821
Bacteria added at start
712
204
Extract/water ratio
0-018
—
Reaction of extract
-0-38
-00
The figures are instructive, inasmuch as they show a consider-
able reduction in the number of bacteria originally added. There
was something in the solution which was strongly toxic towards
the test-organism added to the extract. It is unfortunate that
the extract was so alkaline, for this undoubtedly clouds the issue,
but, even when an allowance is made for it, according to the
information previously obtained with extract of vegetable mould
(p. J 72), a strongly rising dilution-curve is still apparent.
The alkalinity was determined by boiling 25 c.c. of the extract
with 35 c.c. of distilled water and 1 c.c. of centinormal sulphuric
acid for ten minutes, rapidly cooling the solution, adding 1 c.c.
of centinormal soda, and titrating back until the tint became
that of the control. Phenolphthalein was used as the indicator.
It gave a true indication of the reaction in extracts of leaf-mould,
but, as will be shown subsequently, it is not so good for solutions
of hay-infusion.
The experiments so far showed that some means must be
adopted to eliminate the excessive alkalinity of the fermented
cultures. On a previous occasion, humic acid had been used,
but, as it had been soluble, it had not served the desired purpose.
On standing, however, a solution of the acid had precipitated,
and this precipitate of insoluble humic acid was washed and
used. The employment of soil for maintaining a neutral reaction
was suggested by the fact that, when it is put into hay-infusion
and incubated, there is developed a mixed flora and fauna, the
appearance of which is so healthy, that a pronounced acidity or
alkalinity of the infusion is unlikely.
BY R. GREIG-SMITH.
177
In one test, a small amount of sodium phosphate was added to
the hay-infusion, but, from the appearance of the amoeba?, the
salt seemed to have enhanced the alkaline effect, at any rate it
favoured bacterial growth and rapidly destroyed the amoebae.
The neutralisation of the alkalinity with lactic acid proved to
be useless, for, in three days, the culture was as alkaline as
before the addition.
The advantage to be gained by using insoluble humic acid or
soil was tested with solutions obtained by diluting a 1% infusion
of hay with nine volumes of water, that is they contained one
part of hay per thousand.
Experiment xxii.
Protozoon
Amreha Umax.
Medium
Hay-infusion with
humic acid.
Hay-infusion with soil.
Test
1
2 ( 3
4
5
6
7
8
Duration of test in days
10
14 1 24
8
12
15
22
36
Extract, boiled
63o
503
869
955
1,749
2,160
637
3,477
Extract, raw
100
100
100 i
100
100
100
100
, 100
Extract, raw, 80%
107
115
108
117
151
106
87
122
Extract, raw, 50%
104
182
117 !
160
169
313
109
190
Extract, raw, 20%
89
203
129 1
264
451
665
156
494
Water- control
55
86
137 !
302
398
1,036
288
332
Bacteria added at start
9-6
40 20
52
188
99
42
35
Extract/ water ratio . . .
1-8
ri
0 09
0-33
0-25
0-1
0 34
0-3
Reaction of extract . . .
+ 0-0
+ 0-02
-0 04
+0-02
-0-02
-0 02
-0 05
+0-03
The tests were started with 500 c.c. of 0T% hay-infusion, either
a small quantity of washed humic acid (about 0-03 gram) or 25
grams of sterile soil and 10 c.c. of an amoeba-culture. The latter
represented from 30,000 to 50,000 mobile forms as estimated by
the counts of later starter-cultures. The amoebae in the earlier
periods were not counted, but they grew well, and a count made
on the sixteenth day showed 10,000 per c.c. in the humic acid,
and 5,600 in the soil test. On the twenty-fourth day, the flasks
contained 6,600 and 3,600 per c.c. respectively.
The results show a low reaction, and it was assumed that the
fluids were approximately neutral. The humic acid tests do not
178 CONTRIBUTIONS TO OUR KNOWLEDGE Of SOIL-FERTILITY, xvi.,
exhibit any pronounced degree of toxicity as judged by the effect
of boiling, or dilution, or by the relation of the raw extract to
the water-control. They are of the indefinite type which leads
one to further experimenting in the hope of obtaining something
more definite.
In the soil-tests, we have a direct evidence of toxicity in the
twelve days' culture. The filtered extract was directly toxic,
reducing the number of bacteria added at the start from 188 to
100. The effect of boiling the extract for an hour under a con-
denser, and of diluting the raw extract is also well shown in the
increased nutritive effect. The toxic nature of the twelve days'
culture is confirmed by that of the fifteen days', for, although the
added bacteria are not decreased, they did not increase.
A trial was made to see if humus, such as vegetable-mould,
would be useful for maintaining an approximate neutrality.
Four grams of lime-treated mould were tried against 50 grams
of sterile soil. In fourteen days, the humus test showed a reac-
tion of - 0-4, and the soil - 003. Humus was therefore useless
for the purpose. There was no evidence of toxicity in the
extracts.
While dilute hay-infusion has been found to be very good for
growing the amoebae, and for obtaining evidence of toxin-forma-
tion, it seemed advisable to see if a simple nutrient would be as
good. The preliminary tests in the cultivation of the amoeba-
had shown that they grew well in urea and ammonium chloride
and, with these, calcium nitrate was included as a nutrient,
which would probably not produce an alteration in the reaction
of the culture-medium. Solutions of urea, 0*02%, ammonium
chloride, 0036%, and calcium nitrate, 0-056% in tap-water, were
prepared. These contained equivalent quantities of nitrogen.
To 500 c.c. of these solutions, 50 grams of soil and 25 c.c. of an
amoeba-culture containing 75,000 mobile forms were added.
Unfortunately the amoebae did not increase in numbers and were
rarely more than 330 per c.c. Extracts were prepared on the
seventeenth da}^ when all hope of their increase had been given
up. The extracts had the following reactions, urea, - 1-36,
ammonium chloride, +0*24, and calcium nitrate, +004. There
6Y R. GREIG-SMITtl.
179
was no evidence of toxicity other than could be accounted for
bv the reaction. The experiment did not indicate that any of
these chemicals would be of any value in the research.
The Influence of Aeration.
With the idea of determining the influence of aeration upon
the production of toxin, a quantity of dilute (0-2%) hay-infusion
was infected with a culture of ameebse, 50 grams of soil were
added, and the bottle containing the test was attached to an
aspirator, which caused a few bubbles of air to pass through the
liquid every few minutes. A control-bottle was allowed to stand
in the laboratory. They were ordinary litre- bottles, and the 500
CO. of infusion was 2| inches deep. Portions of the fermented
liquids were filtered through porcelain on the fifth day, and
again on the eighth day, with the following results.
Experiment xxiii.
Treatment
Aerated.
Not aerated.
Duration of test in days . . .
5
8
5
8
Extract, boiled ...
33
122
1,690
9,550
Extract, raw
100
100
100
100
Extract, raw, 80%
113
126
102
137
Extract, raw, 50%
117
144
160
429
Extract, raw, 20%
108
356
229
600
Water-control ...
74
242
152
574
Added at start ...
17
21
38
49
Extract/water ratio
1-3
0-4
0-6
0.2
Reaction
+0-2
+0-06
-0-06
- 0-02
The solutions were twice the strength of those of Experiment
xxii., but whether this accounted for a slow growth of amoebse in
the non-aerated test or not, cannot be said. None were seen on
the fifth day, 330 on the eighth, and 2,300 per c.c. on the
twelfth. In the aerated test, the amcebge grew well: 3,000 were
noted on the fifth, and 9,300 per c.c. on the seventh day. Aera-
tion induced an acid, and its absence an alkaline reaction. The
acidity was so high on the fifth day that, on the sixth, the aera-
tion was stopped. This appeared to have had an effect upon the
dilution-curve.
180 CONTRIBUTIONS TO OUtt KNOWLEDGE OF SOIL-FERTIMTY, XVI.
The greater growth of amoebie and the smaller probability of
the production of a toxic effect in aerated cultures raised the
idea that, like yeast-cells, the reproductive may be inversely
proportional to the physiological activity. So another experi-
ment was started, and, as a variation, a flask was included
which was infected with a culture of bacteria only, derived from
a protozoon-free droplet of a starter. The extracts were tested
on the sixth day.
Experiment xxiv.
Micro-organism
Amrebo
. liviax.
Bacteria only.
Method
Aerated.
Not aerated.
Not aerated.
Extract, boiled
1,168
2,039
3,453
Extract, raw ...
100
100
100
Extract, raw, 80%
7o
109
106
Extract, raw, oO%
99
123
87
Extract, raw, 20%
103
199
260
Water-control ...
87
100
78
Added at start...
15
17-5
13-7
Extract/water ratio
115
ro
1-27
Reaction
-fO-04
-0 05
+0
The numbers bear out the contention that, if toxicity is to be
obtained, it will not be as the result of aeration. The reaction
went on as in the previous case, aeration producing acidity, and
no aeration, alkalinity in hay-infusion. The examination of the
culture-fiuids showed that the am eel Ke had begun to encyst in the
aerated, and that 1,000 mobile forms per c.c. were in the other.
A further test was made upon the same lines; as a variation,
a deep layer of fluid was used without aeration in order to accent-
uate the conditions. The method at this time had been to use
700 c.c. of fluid contained in a bottle of about 1,200 c.c. capacit}',
and, in this, the fluid had a depth of 9 cm. In the deep test,
1,700 c.c. were used, and in an ordinary Winchester this had a
depth of 16 cm. In the latter, the amoebae grew slowly, the first
indication being obtained on the thirteenth day, when 330 per
c.c. were noted. On the seventeenth, they had risen to 1,000.
The aerated test showed 5,300 on the fourth, and, without
aeration, the first evidence, 330 per c.c, was obtained on the
eleventh day.
BY R. GRKIG-SMirH.
181
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182 CONTRIBUTIONS TO OUR KNOWLEDGE OF SOIL-FERTILITY, Xvi.,
Taking the figures as a whole, there is not sufficient difference
between them to justify the consideration that any toxin had
been produced by the amcebse, for rises in the dilution-curve are
obtained in their absence. In view of later experience, the rises
might well have been caused by the reaction of the culture-fluid.
About this time, it became evident that the method of de-
termining the reaction by the use of phenolphthalein, as the
indicator, might be faulty when hay-infusion was used, and that
results based upon the reaction of a medium such as extract of
vegetable-mould did not hold for another medium, such as hay-
infusion. The method had been to add a few drops of phenol-
phthalein to a portion of the extract, and boil for ten minutes.
If the solution became purple, another portion was boiled with
acid, and titrated back. If it only became slightly reddened or
tinted, it was titrated without boiling with acid. When methyl-
orange was employed as the indicator, the reaction-numbers were
much higl^er.
It became necessary to examine the influence of various
strengths of hay-infusion, as a direct toxic action had been found
in a 01% solution, and had not again been obtained in 0-2% solu-
tions. The bottles received 700 c.c. of infusion, 60 grams of
sterile soil, and 20 cc. of a starter containing 112,000 mobile
amoebae.
Experiment xxvi.
Strength of infusion . . .
0-05%.
0-1%.
0-2%.
Duration of test
5
8
7
10
7
10
17
Extract, boiled
610
135
729
353
1,016
2,038
1,028
,, raw...
100
100
100
100
100
100
100
„ 80%
—
111
94
219
98
425
111
,, ,, 50%
377
118
174
216
143
586
167
,, ,, 20%
150
69
298
28
343
136
250
Water-control
122
61
219
86
400
154
181
Added at start
16
8
28
11
51
20
28
React'n, phenolphthalein
-0-02
-0-03
-0 02
-0-02
-0 02
-005
-0 06
, , methyl -orange . .
- 0-52
-0-23
- 0-32
-0-30
-0-52
-0-39
-0-52
Extract/water ratio
0-8
1-6
0-5
11
0-2
0-6
0-5
Amtebsi per c.c.
500
1,000
1,830
2,000
660
5,600
2,000
6Y R. GREIG-SMITtt. 183
There was no direct toxic eflfect in any of the tests, and, so
far as the possibility of obtaining such by using different
strengths of infusion are concerned, the strongest appears the
most likely. The ten-days' extract of the 0-2% solution gave more
favourable numbers than any of the others. In this, the amoebae
were the most numerous.
A further test with 0-1% hay infusion was made, but in it the
amcebae did not grow quite so well as on the former occasion
when the direct toxic action was obtained. The counts showed
830 per c.c. on the seventh, and 1,000 on the eighth day. No
evidences of toxicity were obtained on the eighth or twenty-first
days, and it must be concluded, that a definite toxicity cannot
be demonstrated by growing bacteria or amcebse in the usual
culture-duids.
The Influence of Reaction.
Much has yet to be found out regarding the influence of re-
action upon bacterial growth, and the reason for the rise in the
dilution-curves of th« extracts. As the infusions and culture-
solutions are generally made with tap-water to supply a small
quantity of saline matter accepted as being necessary for the
growth of micro-organisms, a beginning was made with it. Tap-
water is known to be alkaline, but the extent of the influence of
its faint alkalinity is not know^n. The nutrients were increased
by preparing infusions of hay, and these were made sujQEiciently
weak to enable useful counts to be obtained. A 1% infusion of
hay was diluted with 99 parts of porcelain-filtered, distilled and
tap-water, thus obviating heat-effects. Portions were treated
with lactic acid and with ammonia so as to bring up the final
volume to a definite reaction, and sown with a definite number
of cells of Bac. produjiosus. They were incubated at 22° for 20
hours, and counted. In tabulating the results, the highest
counts were taken as 100.
184 CONTRIBUTIONS TO UUR KNOWLEDGE OF SOIL-FERTILITY, xvi.,
Experiment xxvii.
Reaction.
Distilled water.
Tap-water.
+ 0-3
11
16
+ 0-2
18
29
+01
44
100
+0-05
59
—
00
100
5
-0 05
44
—
-01
28
01
-0-2
7
0-05
-0-3
0-6
002
Reaction of the untreated diluted infusion
to methyl-orange ...
±0-0
-0-20
To phenolphthalein, direct
—
-0-03
To phenolphthalein, indirect ...
—
-014
The distilled water test shows that the bacteria grow best in
a neutral solution, and the tap- water test that neutrality is ob-
tained when approximately +0-1'' of acid has been added. The
curves of these numbers are steep on both sides of the approxi-
mately neutral line, indicating that a slight difference in the
reaction of an extract will make a great difference in the growth-
numbers. As it is impossible to obtain a neutral reaction in the
cultivated extracts, it would be necessary to neutralise them
before dilution in such tests as have been made. It is not clear
how this could be done, for, in the solution under examination,
the bacteria showed 0'1° of alkalinity, methyl-orange showed 0-'2°,
phenolphthalein by direct titration after boiling, 003°, and by
indirect or back titration, 0*14°.
The experiment would be incomplete without the inclusion of
others showing the influence of diluting distilled and tap-water
infusions with distilled water, as is customary.
BY R. GREIG-SMITH.
185
Experiment xxviii
.
Distilled water.
Tap-water.
Raw
Boiled.
a
b
a
b
Solution, boiled ...
,, unboiled
„ 80%
50%
„ 20%
Distilled water-control
Added at start ...
Solution/water-ratio
Reaction to method-orange
Reaction to phenolphthalein, direct
Reaction to phenolphthalein, indirec
26 —
100 100
75 86
— 42
27 18
2 3-5
0-4 2
47 28
-0 -0
- 0 - 0
-0 -0
1,020
100
140
1,160
2,880
263
50
0-4
-0-16
-0 05
-0-16
131
100
234
207
182
32
20
2.5
-0-24
-0-11
-0-24
The boiled tap-water infusion, "b," was boiled three times
upon successive days, as is usual in preparing such culture-
media. Tests "a" were made eleven days before tests " b.'' Bv
an accident, the "b"' tests were incubated at 26° instead of 22"
as in " a," and this should be borne in mind when considering
the increase of the unboiled numbers over the start.
The distilled-water numbers are considered to be normal, for
they show a gradual fall as the nutrients are weakened by dilu-
tion with water. The tap-water curves rise as the alkalinity is
weakened. The numbers of the raw tap-water test indicate
that, after all due allowances are made, it is of a toxic nature,
which is probably not entirely traceable to the alkalinity. The
numbers of the boiled- water test are not so pronounced, and are
much the same as liave been obtained in previous experiments
with bacterial and protozoon cultures. Tt should, however, not
be forgotten that many of these gave normal curves.
A set of experiments were started when it had become evident
that, in all probability, the reaction had more to do with the
symptoms of toxic effect than anything else. In the endeavour
to get round any individual action of the bacteria, flasks of
dilute (0-1%) hay-infusion, made with distilled water to avoid
the action of tap- water, were seeded with an amcebse-culture, and
with a bacterial culture derived originally from a protozoa-free
186 CONTRIBUTIONS TO OUR KNOWLEDGE OF SOIL-FERTILITY, xvi,,
droplet of soil-suspension. The latter were thus controls. The
amoebEe-cultures were twice seeded with amcebse, once, at the
start, and again on the second day, as the first seeding did not
seem to have been successful. They grew slowly at 18"^ and
exhibited a twenty-fold increase on the tenth day, and a fifteen
and ten-fold increase on the thirteenth day with the humic acid
and humic acid -|- soil respectively. They were tested on the
fourteenth day.
Experiment xxix.
Humic acid.
Humic acid +soil.
Amcebffi.
No
amoebse.
Amoebfe.
No
amcebse.
Extract, raw
„ 50%
Water-control ...
Added at start ...
Extract neutralised with lactic acid
Extract/water-ratio
Reaction to methyl-orange
100
192
133
16
104
0-75
-0-15
100
125
82
10
127
1 2
-0-28
100
184
98
11
198
10
-0-21
100
183
88
10
243
ri
-0-28
The addition of soil to the culture-fluids had no influence, one
way or the other, in increasing or decreasing the numbers, and
its use appears to be of no value. The experiment shows pretty
clearly that any toxic effect is not caused by the amcebse, but
rather by the bacteria which always accompany the protozoa.
It is doubtful if the alkalinity, as indicated by methyl-orange, is
a true index, but, if accepted as true, the neutralised solutions
were certainly more nutritive than the unneutralised, for the
nnmbers were higher. With a better indicator, a lower reaction
might have been obtained and higher numbers furnished in the
neutralised tests. At any rate, it is pretty safe to conclude, that
the rise in numbers upon dilution is largely, if not entirely, due
to the lesseninof of the alkalinity of the filtered extracts.
Reaction-Experiments.
The reaction of the soil-extract is never constant, but varies
from day to day. probably within certain limits. For example,
an extract was made on November 30th, 1917, by taking 300
BY R. GREIG-SMITH.
187
grams of garden-soil and 300 c.c. of distilled water. The two
were shaken 300 times during an hour, and filtered. The re-
action to methyl-orange was -0-14, although a paste of the soil
was acid to litmus.
A similar extract, made on December 6th, had a reaction to
methyl-orange of - 0-24. This extract was examined, with the
following results.
Experiment xxx.
Soil-extract, boiled
21
,, raw
100
„ 80%
100
. 50%
64
„ 20%
12
Water-control ...
0-5
Added at start ...
0-7
p]xtract/water ratio
206
Quantities of lactic acid and of carbonate of soda were added
to vary the reaction, and the treated extracts were seeded with
the test-organism in the usual manner. The following numbers
were obtained after the usual 20 hours' incubation at 22".
Experiment xxxi.
Acid or alkali added.
Net reaction.
Bacterial grow^th.
+0-4
+016
1-5
+0-3
+006
6-3
+0-2
-0-04
17
+0-1
-0'14
49
+0-05
-019
100
0-0
-0-24
81
-0-05
-0-29
31
-0-1
-0-34
15
-0-2
-0'44
1-3
The true neutral point was reached by adding 006° of acid,
that is to say, the methyl-orange indication was - 0-19 in excess.
The effect of shaking up various quantities of soil and water
was tested, to see the differences in the reaction of the extracts.
Round numbers were taken, but, as the solid contained 8% of
moisture, a correction was made.
188 CONTRIBUTIONS TO OUR KNOWLEDGE OF SOIL-FERTILITY, XV:
Proportion of soil to water, round numbers ... 2:1 1:1 1:2
Proportion of soil to water, calculated for dry
soil... ... .. ... ... 1 :0-63 1 :ri7 1 : 2-26
Reaction to methyl-orange ... ... —0-24 -0-18 i —0-12
Reaction to phenolphthalein, direct ... -0*044 I -0*016 | -0*008
2:1
1 :0-63
1 :1
1 :ri7
-0-24
-0 044
-0-18
-0-016
Curves of these numbers show that the reaction of the soil-
water, equivalent to a ratio of 1 : 0*087 for the dry soil to 8% of
moisture, would be between -0*4 and -0-5 for methyl-orange,
and about - 0*09 for phenolphthalein; yet the soil was acid to
litmus.
The reaction to phenolphthalein was examined somewhat
closely, after the suspicion was raised that the reaction of the
solution had more to do with the phenomenon of toxic action
than had been supposed to exist. It was found to be untrust-
worthy, as much depended upon the rate at which the solutions
were boiled, previously to cooling and titration. For example,
covered beakers containing 50 c.c. of tap-water were boiled
slowly and rapidly, cooled and titrated.
Duration in minutes
10
20
30
Slow boiling
Rapid boiling ...
-0 022
-0-076
-0-048
-0-105
-0-071
-0-118
Much, therefore, depends upon the method and time of boiling
in expelling the carbon dioxide. In the tests previously recorded,
the extracts were boiled in an open beaker for ten minutes at a
speed intermediate between slow and rapid boiling. They pro-
bably do not indicate the true alkalinity, but, for that part, it
has been shown that the same would have occurred by using
methyl-orange.
A twenty-one-days' culture of bacteria and amcebse was filtered,
and the extract treated with lactic acid in decreasing quantities,
seeded with tlie test-organism, incubated and counted. It had
a reaction to methyl-orange of -01 6°, and to phenolphthalein,
direct, of - 004".
BY R. GREIG-SMITH.
EXPKRIMKNT XXxii.
189
Lactic acid
added.
Neb reaction.
Bacterial counts.
0-31
+ 015
s
0-2«
+ 0-10
19
0-21
+ 0 05
41
016
00
46
0 11
-0 05
40
0 06
-0 10
100
0 0
-016
53
The experiment shows that, in the case of a fermented culture-
fluid, the lessening of the alkalinity by 0*06° produced neutralitj'.
The methyl-orange indication was -0'10°too high. A high
indication was also obtained with soil-extiact in Experiment
xxxi., and with tap- water in Experiment xxvii.
The acidity of the extracts of sterilised vegetable-moulds (Ex-
periments XV., xvi.) led to an examination being made of the
extracts of soil which had been sterilised in the same manner,
viz., for two hours at 1 30".
Two hundred gram-portions were put into sterile bottles with
10 c.c. of a suspension of Amoeba Umax, containing 10,660 mobile
forms per c c, and 40 c.c. of water. The soil thus had an ex-
cessive amount of moisture, 20%; it usually contains from 8 to
10%. The bottles were kept at laboratory-temperature, which
varied from 18-23°.
Extracts were made in the usual manner by adding 200 c.c. of
distilled water, shaking frequently during an hour, and filtering
through porcelain. Uhe extracts were seeded with Bac. pro-
digiosus, incubated and counted.
Experiment xxxiii.
Duration of test in days ...
3
63
Extract, raw
Extract, raw, 50%
Waler-coiitiol ...
Added at start ...
Extract/water ratio
Reaction to methyl-orange
100
240
1
0-16
110
+ 03
100
48
1-2
014
153
-0-8
14
190 CONTRIBUTIONS TO OUR KNOWLEDGE OF SOIL-FERTILITY, xvi.
The point to be noted is the change in the reaction of the ex-
tracts. It changed from being comparatively strongly acid on
the third day, to strongly alkaline on the fifty-third. When the
extract of the latter was treated with lactic acid to neutralise
the apparent alkalinity, the number obtained was 007 as against
100 for the non-neutralised extract; that is, for every seven
bacteria in the neutralised, there were 14,000 in the alkaline
extract. The acid had clearly been added in excess, tlie —OS'
being far from a true indication.
This confirms the result obtained in another place, that the
reaction of a soil-extract, as judged bj^ methyl-orange, is not the
true reaction. Mobile amoebae were seen in the soil on the fifty-
third day.
Conclusion. — It has been shown that certain soil-bacteria,
moulds, and araoebse, all reasonably supposed to be capable of
furnishing substances of a toxic nature, were grown in various
media and under varying conditions; and, in all cases, the signs
of toxicity which became manifest could be attributed to an
alteration in the reaction of the media.
The test-organism, Bac. prodigiosus, grows best in a neutral
medium, and an indicator is required which will indicate strict
neutrality. The methyl-orange numbers are too high, and the
phenolphthalein too low. Small divergences from the neutral
point strongly affect the growth.
The humus of leaf-mould contains two types of humic acid;
one absorbs alkali from alkaline carbonates, and the other from
alkaline carbonates and hydrates. These were present to the
extent of one part of the former to three of the latter. Heating
the humus increases the amount of acid, and the increase is
largely soluble in water.
The effect of reaction is quite of a different order from the
evidence of toxic action obtained in former researches.
I am indebted to Mr. W. W. L'Estrange for kind assistance
during the later stages of the work.
191
ON CERTAIN SHOOT-BEARING TUMOURS OF EUCA-
LYPTS AND ANGOPHORAS, AND THEIR MODI-
FYING INFLUENCE ON THE GROWTH-HABIT OF
THE PLANTS.
By J. J. Fletcher and C. T. Musson.
(Plates iv.-xxvi.)
The Eucalypts, representing about 230 recognised species,
contribute one of the dominant, phanerogamic elements to the
Australian flora. They are an assemblage of plants remarkable
in many ways, widely distributed over an entire continent, ex-
tending also to the circumjacent islands; and now acclimatised
to some extent in other countries. One of the astonishing
things about them is the liability of the seedlings of so many
species to shoot-bearing galls or tumours of an uncommon type.
Their specially distinctive characters result from a fortuitous
combination of some simple, natural, and favouring conditions
present in quite young seedlings. Firstly, they originate in the
axils of the cotyledons only, or, in addition, in a few pairs of leaf-
axils successively above these, where the buds are, as paired but
at first independent, proliferating outgrowths of cambium-tissue;
and, as a rule, the outgrowths, or the axillary stem-nodules, as
we may call them at this stage, succeed in taking possession of
the dormant buds, and incorporating them in the stem nodules.
This is how the latter, as well as the composite tumours to which
they may give rise, come to have buds or shoots.
(Secondly, the young seedlings usually have opposite and
distichous leaves; and, correspondingly, the stem-nodules are
also opposite and distichous; but as, under favourable conditions,
the latter grow faster than the stem thickens, the paired nodules
meet and fuse, and the fusions then encircle the stem.
Thirdly, as a rule, the first and second internodes do not length
too much or too soon to permit of the concrescence of the fused
192 TUMOURS OF EUCALYPTS AND ANGOPHORAS,
pairs, giving rise to composite, shoot-bearing, stem-encircling
tumours. The third and succeeding internodes are metre vari-
able, but often permit of the addition of a fourth and a fifth
pair of fused nodules to the concrescence of the first three pairs.
More than five pairs, if developed, are apt to be left isolated, and
are only of minor or of no importance. Three to five pairs are
the really effective and important ones.
Fourthly, the hypocotyl is not too long, so that, as the com-
posite, stem-encircling tumour increases in size, and grows down-
wards, it is able to tamper with as much of the root-system as it
encounters, encircling and fusing with the upper portion of the
tap-root, and the proximal portions of any lateral roots that come
in its way, intercepting more or less of the water they may
contain, at the expense of the seedling-stem, and to the ad-
vantage of the tumour and its inhabitants. In this stage, they
are composite, stem-encircling, shoot-bearing, root-incorporating
[but not root emitting] tumours.
The stem-nodules of seedlings of refractory species, or of re-
fractory individuals of any species, may not get beyond the first
stage. In such cases, the proliferation slows down, and the
axillary stem-nodules come to a standstill. A mild attack has
run its course without accomplishing anything of importance, or
interfering with the growth-habit of the seedlings.
Seedlings of susceptible species, from accidental or other
causes, may not get beyond the second stage, through the com-
posite, encircling tumours failing to incorporate roots. In such
cases, the composite tumours may last for some time, but not as
long as those which can complete the third stage.
In the Non Mallee Kucalypts, the last stage may last for some
years, until the plants are getting ready to enter on the young
sapling stage, but not indefinitely. In the Mallees, however, it
may persist throughout life, and the tumours may attain an
enormous size. But the distinction between the two groups of
Eucalypts does not correspond to inherent differences in their
tumours, as such; but to differences in the kind of roots incor-
porated in the two cases. In other words, the Non-Mallees have
ordinary roots; the Mallees have water-storing roots.
BY J. J. FLETCHER AND C. T. MUSSON. 193
In all three stages, the buds may remain dormant, or they
may develop shoots, according to circumstances. In the fifst
case, an appropriate stimulus will readily cause shoots to develop.
Anomalies, eccentricities, and examples of erratic behaviour are
common, and very instructive.
In addition to the axillary stpm-nodules, unpaired, not de-
finitely localised nodules sometimes develop at a lower level than
the cotyledons, either on the hypocotyl or on the taproot, of some
seedlings. These are very suggestive of wound-infection, and
are sometimes obviously pathological. This is particularly the
case in most seedlings of two of the Bloodwoods, in which very
extensive root-invasion may take place.
In all these cases, we can find no escape from the conclusion,
that the exciting cause of the nodules is attributable to parasitic
soil-organisms, probably of the same species, finding vulnerable
points for attack under slightly different conditions; and causing
the cambium-tissue to proliferate and grow outwards. The
nodules are not caused by insects, or other visible parasites. They
are outgrowths of proliferating tissue, and not natural growths.
They may be a nuisance to the plants, and may temporarily or
permanently interfere with the natural growth-habit. More-
over, if possible, when opportunity offers, they are brought to a
standstill.
The root-nodules of four young seedlings, of which three are
figured, in every case have incorporated the basal portion of a
young root. They suggest analogy to what Beyerinck has ob-
served in studying the root-nodules of Leguminosse — "'J he
splitting of the primary bark for the emission of the side-roots is
the special means of entrance of JJ. radicicola^' [Erwin Smith,
" Bacterial Diseases," ii., p.l03J.
The vulnerability of the axils may be due to a weak spot
which arises in connection with the differentiation of the buds,
or which is exposed about the base of the petioles when the
young leaves expand. Until serial, transverse, and superficial,
longitudinal sections through the axillary region of series of
seedlings have been studied, it is useless to speculate further.
In the belief that soil-organisms are responsible for the problem
194 TUMOURS OP EUCALYPTS AND ANGOPHORAS,
we are interested in, we began a search for records of similar or
analogous cases.
Dr. F. Erwin Smith, in the second volume of his important
treatise on " Bacteria in relation to Plant- Diseases " (1911), dis-
cusses, in detail, wound-infections (p. 51 ), and also infections
through natural openings, grouped as nectarial, waterpore-.
stomatal, and lenticellate infections But we failed to find any
reference to axillary infections. As we came to know later,
axillary infections had not then come under the author's notice.
It was with great interest that we unexpectedly met with a
paper by Mr Clayton O. Smith, of California, in which he not
only mentions the possibility of axillary infection in the axils of
the cotyledons, in stone-fruits, but he gives particulars about the
axillary nodules of some Eucalypt seedlings which came under
his notice. We quote all that the author has to say about these,
and about cognate matters, for three reasons. The paper is not
contained in the Society's librar}', and we know of only one copy
of it in .Sydney. It contains the first satisfactory record known
to us of the realisation of the fact, that the stem-nodules are
axillary; as well as the only records, that we know of, of the
successful inoculation of Austrahan plants, including Eucalypts,
from cultures of soil-organisms. It is also of interest to note
what an experienced plant-pathologist thought of the stem-
nodules. Mr. Clayton Smith does not mention the fusion of tlie
axillary knots, or that they had shoots, or that they incorporated
roots. E. tereticornis is the only species mentioned, and seed-
lings of this species are sometimes more or less refractory. It
may be, therefore, that with only limited macerial available, and
without Australian experience of Eucalypts to guide him, he
may have unknowingly experimented with seedlings that were
not as satisfactory for the purpose as others might have been.
"Further Proof of the Cause and Infectiousness of Crown
Gall.' By Clayton O. Smith. Univ. Cal. Publications, College
of Agric, Agric. Experim. Station Bulletin No.235 (Dec. 191*2).
"Bacterial Kature of [the] Disease" (p. 534).— There is now
abundant proof, that these knots [Crown-Gall on the 30 Hosts
enumerated] are caused by a bacterial organism that enters the
BY J. J. FLETCHER AND C. T. MUSSON. 195
tissue through some injury, or possibly at the point where the
cotyledons of sucli seeds as those of the stone-fruits are attached
to the young plant. In Eucalyptus seedlings, the natural knots
often appear opposite each other where the cotyledons have pre-
viously been attached, also the quince knots appear first at the
node about the old leaf-scar. All the evidence we have, goes to
show that some injury or weakness is necessary for infection to
take place.''
[Legend of Text-fig.21, p.549].— "Artificially caused galls on
forest red gum \^Eucalyptus tereticornis]. Crown Gall has not
been known to attack the various species of Eucalyptus in nature.
The significance of swellings found frequently at the crown of
young gum trees is not yet understood. They do not appear to
be detrimental to the tree. "
Victoria Bottle Tree {Stercnlia diversifolia) [tig.20], and Flame
Tree {S. acerifolia) were successfully inoculated (p. 552).
"Forest Red Gum {E. tereticornis) [fig.21]. — Seedlings of from
four to six feet were inoculated. The first successful inocula-
tions were made May 16, 1910. On March 25, 1912, there was
one large knot and one very small one at points of inoculation.
September 2, 1911, inoculated a seedling about one half inch in
diameter. February 20, 1912, there were two small knots. On
March 26. 1912. one of these knots had grown rapidly in size,
the other had not changed."
" Inoculations were made on small seedlings, July 20, 1910, on
the branches. Typical roundish knots or galls had developed
on September 5, 1910."
[Silky] Oak {Grevillea robusta) (p.552), gave negative results,
" This is probably due to the slow growth of the tree."
Under the heading of Other Galls that are at 'present little
under stood {p. bb'2}. — ^'Eucalyptus A''no^(pp. 55.3-554)— Galls occur
frequently on the small seedlings. These are hard knots that
occur always at the nodes. There will often be a gall on each
side of the plant where cot3ledons or the opposite leaves have
previously been. We have had small seedlings with these natural
galls under observation for about three years. In some cases
there is an increase in size in the knots, which still continue to
196 TUMOURS OF EUCALYPTS AND ANGOPHORAS,
be hard, and do not in any way appear to weaken the trees. In
many cases, the diseased trees have completely outgrown the
disease, and in these instances there has been but little increase
in size of [the] original, knots. From our observations, the dis-
ease does not seem to seriously injure the growth of the tree.
Numerous attempts to isolate a pathogenic organism have re-
sulted in failure, although we still believe the galls may be
caused by such an organism."
Quite recently three papers by Dr. Erwin F. Smith, of the
U.S. Department of Agriculture, have come to hand, which have
a very important bearing on the problem of the shoot-bearing
tumours of Eucalypts.* 'J'hese also are to be found only in
a few libraries. '1 hey are important because they relate to the
first records of natural, axillary, shoot-bearing tumours caused by
soil-organisms; and axillary and other special tumours produced
by artificial inoculations. We give the following extracts from
these : —
Dr. Erwin Smith says in his paper published in August, 1916:
" Last winter, I discovered that when growing plants are inocu-
lated in the vicinity of dormant buds, a new type of tumor is
produced. This tumor bears, on its surface, diminutive abortive
shoots (vegetative or floral) and in its interior, along with the
cancer-cells, numerous fragments of embiyonic tissues, variously
fused and oriented, often upside down and curiously jumbled
These tumors have never been seen by the writer in nature, or
at least if seen, not recognised as crown-gall tumors, but un-
doubtedly we shall now find them." [Then follows reference to
a footnote "As this goes to press, I have had confirmation of
this belief, having received from a florist in Massachusetts, a
crown-gall of the rose showing abortive shoots growing out of
stem-tumor."] Dr. Smith continues— " These tumors have all
* ''Further Evidence as to the Relation between Crown-Gall and
Cancer,'* Proc, Nat. Acad. Sci. U. S. A., ii., p. 444 (August, 1916).
"Mecanism of Overgrowth in Plants," Proc. Amer. Phil. Soc. Philadelphia,
Ivi., No. 6, p. 439 (August, 1917). "Embryomas in Plants (produced by
Bacterial Inoculations)," Bulletin Johns Hopkins Hospital, xxviii., No.
319, p.279 (September, 1917).
BY J .f. FLETCHER AND C. T. MUSSON. 197
been produced with Barter ium tumefadens The plants
chiefly experimented on have been Pelargonium, Nicotiana, T.y-
copersicum, Citrus, and Ricinus. All of these and some others
(Mangifera, Allamanda, etc.) have yielded teratoid tumors
from inoculations in leaf-axils/'
In a later paper, Dr. Erwin Smith refers again to the acci-
dental circumstance which suggested to him the desirability of
inoculating plants in leaf-axils and growing-points, in addition
to internodes, as he had been doing for years. " We had found
indeed, as early as 1908-9, and had produced by bacterial inocu-
lation, plant-tumors bearing roots, but the full meaning of this
discovery, as related to cancer, did not occur to me until early
in 1916. when I found crown-gall tumors bearing leafy shoots on
some of our inoculated hothouse geraniums. Beginning with
this discovery, I made numerous inoculations in the leaf-axils of
various plants, which resulted in the production of leafy tumors,
and subsequently I produced them on leaves and on cut inter-
Tiodes where no buds occur normally. Tumors bearing roots have
also been produced by us on the top of plants, and in one cut
internode of Tobacco I succeeded in producing a tumor which
bore flower-buds. These perishable root-bearing and shoot-bear-
ing tumors I regard as plant-embryomas, and have so described
them [Journ. Cancer Research, April, 1916, p. 241]."
We have not had access to the paper last mentioned. Rut in
the Johns Hopkins Bulletin for September, 1917, Dr. Erwin
Smith has given further particulars, and numerous illustrations
of the embryomas resulting from his inoculations, together with
two (fig.63) of "Pelargonium teratoid tumors originating natu-
rally ... on gardener's cuttings bedded in earth for propagation.
The specimens came from a gardener's house near Jjaltimore."
We give the following brief extract from this paper — "In
April 1916, I announced the discovery of a new type of
crown-gall, i.e., one containing numerous leafy shoots, and showed
that I could produce it at will by making my bacterial inocula-
tions in leaf-axils where there is a dormant bud I have
since discovered that leafy crown-galls occur in nature on various
plants, e.g , on the rose, and on the carnation."
198 TUMOURS OF EUCALYPTS AND ANGOPHORAS,
If the shoot-bearingj axillaiy nodules of Austialian Eucalypti
are caused by parasitic soil -organisms, under natural conditions,
then they seem to be comparable with the five cases of " leafy
tumours" [on Geranium, Pelargonium (two), Rose, and Carnation]
due to axillary infection by the organism causing Crown-
Gall under natural conditions cited, by Dr. Erwin Smith.
Were it not that, by a fortuitous combination of circumstances,
the axillary stem-nodules are able to fuse in pairs, the fused pairs
to concresce, and the re-inforced, composite, stem-encircling
tumours thus enabled to incorporate roots and so last for some
considerable time or even permanently, both the nodules, and
any shoots they might develop, would be short-lived and abortive:
as they actually are in refractory seedlings; and as the shoots on
the lower pairs of concrescences also are.
They are not exactly comparable with the embryomas pro-
duced by Dr. Erwin Smith's inoculations in leaf-axils and grow-
ing-points. But the circumstances and conditions in the two
cases are not parallel. Erwin Smith's experiments were mostly,
but not entirely, carried out with soft-tissued plants, which
responded promptly; the organisms were introduced by needle
inoculations right into the tissues of the plants, causing profound
disturbances; and the inoculations were made in upper axils.
But in the natural inoculations in the lower axils of the young
seedlings of Kucalypts, which furnish some of the most \alued
hardwood timbers, we are inclined to think that the organisms are
confined to the outgrowths, and the encircling tumours to which
they give rise, and probably do not invade the tissues of the seed-
lings. The tumours do not kill the seedlings, or even seriously
damage their tissues. 1 hey are a drag on the normal develop-
ment of the plants, especially so when shoots do not develop, and
bv interfering with the water-supply, and also by iheir shoots
preventing the development of the normal branching. In the
Mallees, so much water is intercepted by the tumours, that the
seedling-stem is dwarfed; and, by the persistence of the shoots,
the growth-habit is permanently distorted, so that the plants are
prevented from realising their potentialities as trees. The seed-
ling-stem may possibly be sometimes crowded out and got rid of.
BY J. J. FLETCHER AND C. T. MUSSON. 199
But the stem-nodules, as well as the composite tumours to
which they give rise, are complex tumours, composed of both
somatic cells and germ-cells ; and the latter are totipotent,
because in the persistent composite tumours of the Mallees, the
tumour-shoots complete their growth, flower and fruit, and pro-
duce seed. Even in the Non-Mallees, if the seedling-stem is lost,
two tumour-shoots may take its place, attain to tree-size, and
flower and fruit. But they do not prematurely disclose their
embryonic possibilities, in the way that some of Erwin Smith's
artificially produced monstrosities did. Also the production of
these tumours in Eucalypts under natural conditions is a matter
of long standing. The Mallee Scrubs, which must have been
the developments of centuries, were in their prime, when civilised
man first saw them, nearly 101 years ago.
We may next briefly review the Australian contributions to a
solution of the problem we are interested in. 'Jhese relate
chiefly to the Mallees. The earliest descriptions of the make-up
of these curious plants, by Allan Cunningham,* Tenison-W oods,t
and Baron von Mueller,! are subject to the drawback that, when
these botanists visited the untouched Mallee-Scrubs, there was no
o])portunity of seeing uprooted plants. Two of the Baron's de-
tinitions will suflfice. Under E. oleosa (Dec. vii.) : " Shrubby or
somewhat arborescent It is this species which forms a
large ingredient of the ' Mallee-Scrub,' constituting tall bushes
branched from the root." Under E. striata (Dec. x.): ".Shrubby
or somewhat arborescent ; habit that of the Mallee-Euca-
lypts, with many stems from one root."
Mr. J. Ednie Brown, § in 1882, gave his interpretation of the
" root " of the Mallees, from whicii the stems are said to rise, as
"being apparently a kind of dwarf trunk, the stems of our de-
scription [under E. gracilis. "A Mallee tree with several stems"]
representing its branches."
* Oxley's Journal (in the entry for June 10, 1817), p.63 (1820).
t ''Geological Observations in South Australia,"' p. 83 (1862). Also,
Proc. Linn. Soc. N. S. Wales, vii., p. 566 (1883).
X "Eucalyptographia," under the various Mallees (1879-84).
§ Forest Flora of South Australia, Part v. (1882).
200 TUMOURS OF EUCALYPTS AND ANGOPHORAS,
A little later, Mr. Tepper, of Adelaide, offered another ex-
plaiiation of if*^ — ''Eucalyptus oleosa is a species with an under-
ground rootstoek, from which numerous small stems, generally
crooked and semi-sarmentose, spring. When these are destroyed
by fire, &c , a host of fresh ones spring up from the caudex."
But no details are given of the characters of the supposed root-
stock.
We owe to Mr. N. B. McKayt an interesting description
of tiie underground, composite, stem-encircling tumoui- of a
Mallee, even though it is incomplete, inasmucli as no particulars
about the shoots, or the incorporated roots are given. We have
looked in vain, however, for a more satisfactory one. He says
— '"Tlie indigenous timber-growths on the Mildura horticultural
areas consist, for the most part, of blue bush, 'bull* mallee,
balar, pine, and needle-bush A well established bull mallee
is a problem to the ordinary grubbing contractor. The butt is
a great flattened bulb of curly timber, sometimes 8 or 1 0 feet
through. It is set firmly into the soil, and even if all the roots
were cut off, the tree would stand in its place just the same, as
the upper growth is very insignificant compared with the base.
Chopping mallee out is an obvious impossibility, and, as the
wood is full of moisture, it would be an endless task to attempt
burning it out. Dynamite and rack-a-rock have proved equally
useless. Before, or rather behind, the traction-engine, the diffi-
culty disappears When all the stumps are uprooted, the
adhering earth is knocked off, and they are readily burned."
Three items in this are worth notice. Firstly, that the butt
is a " flattened '' bulb : that it is to say, it increases in size hori-
zontally rather than vertically, Secondly, it is a bulb of "curly
timber": it is not a case of a simple, localised thickenii g of the
stem, but results from the proliferation of cambium-tissue.
Thirdly, '• the wood is full of moisture" : it is a water-charged
tumour, because of the incorporation of water-storing roots.
" "Remarks on the 'Manna' or Lerp Insect of South Australia." Journ.
Linn. Soc. Lond., Zoology, xvii., p. 109 (1883).
t Eighth Progress Report of the Royal Commission (of Victoria) on
Vegetable Products, 1 890.
BY J. J. FLETCHER AND C. T. MUSSON. 201
The late Professor R. Tate, in a paper entitled "A Review of
the Characters available for the Classification of the Eucalypts,' *
etc., under the section " Habit," expressed some vie"\vs. which
should have provided a new starting-point for considering the
make up of the Mallees. Unfortunately, they were so severely
condensed as to be cryptic, and quite failed to influence his
successors. Tate says —
" The Eucalypti comprise two habits of growth, viz., trees and
shrubby trees, to which I «pply the vernacular names of Gums
and Mallees. I do not know if I am correct in so doing, as I
have failed to find any definitions of these well-l<nown terms."
" I have constantly observed in seedlings and growths of one or
two years of such gums, as E. vostrata, leucoxyloti^ virwinalis, a
large inflation of the base of the stem, either at the surface or
just below the surface of the soil. In the species named, this is
eventually outgrown; but. in the mallees, it f ersists and increases
in size proportionately with the development of the branches
which are emitted from it — in the mallee, this rudely globose
bole is partially subterranean."
It is the third and fourth sentences that are important. These
embody Tate's version of the problem we are interested in, com-
pressed to an irreducible minimum. The chief diflficulty arises
from the fact that no attempt is made to explain the nature or
the significance of the basal inflations. A non-committal name
is given to them, but they remain of problematical import. Nor
is any reason given for their being transient in the seedlings of
Non-Mallees, and persistent in the Mallees. Nevertheless, he
recognised the dift'erence. Apparently his seedlings were not
young enough to show the axillary stem-nodules before they had
fused in pairs, and the fused pairs had Goncresced; and they
were not old enough to show that basal inflations were able to
incorporate roots. Jt was. presumably, from necessity, and not
from choice, that he confined his observations to seedlings of
"one or two years." If he had mentioned this, and supple-
mented it by pointing out the importance of studying seedlings
in trying to understand such complicated structures as the adult
* Report Aust. Assoc. Adv. Sci., vii., p.544 (1898).
202 TUMOURS OF EUCALYPTS AND ANGOPHORAS,
Mallees present, those who came after him might have been
induced to follow his lead, and to extend his incomplete observa-
tions. Even what he actually says ought to be sufficient to make
anyone cautious about identifying the persistent basal inflation
of the Mallees as a " rootstock," without first investigating the
transient basal inflations of such Non-Mallees as have them.
Unfortunately his excessive reticence obscured what merit
his observations may have; and his successors, in attempting to
explain the make-up or constitution of the Mallees, either
adopted Mr. Tepper's idea that the Mallees have " rootstocks,"
without offering any explanation of their peculiarities; or else
they make use of one or other of the older definitions, which,
though excusably deficient when they were first offered, are now
out of date.
As far as we can ascertain, Mr. Maiden is the only writer who
has taken any notice of Tate's statement quoted above. On the
first page of his "Critical Revision of the Genus Eucalyptus "
(1903), the author gives substantially Tate's views, almost in the
original words, but without any comment other than " This
classification is chiefly of practical use in Professor Tate's own
State (South Australia) and in Western Australia."
In Plate 57, fig. 12 (of Vol. ii.) of the same work (1911), an
illustration of a young seedling of E. paniculata, with a pair of
axillary stem-nodules still unfused, will be found. 'J his, we
belie^e, is the only illustration of a Eucalypt seedling with stem-
nodules, or any stage of them, which has hitherto been published.
In the explanation of the Plate (p. 131), fig. 12 is referred to as —
" Bulbous swelling in seedling This swelling is very com-
mon in seedlings belonging to this genus, and the cause has not
been investigated so far as I am aware. It is presumably to be
attributed to the action of bacteria." Allowing for the fact that
there is a pair of independent, bulbous swellings, and not merely
one, and that they are the first stage in the formation of Tate's
transient basal inflation of a Non-Mallee, we agree with Mr.
Maiden that they are presumably attributable to the action of
bacteria. But we should say exactly the same about a similar
seedling of a Mallee, with the first pair of axillary nodules or
bulbous swellings present.
BY J. J. FLETCHER AND C. T. MUSSON. 203
The difficulty which confronts anyone who tries to interpret
the axillary, bulbous swellings of seedlings of Eucalypts, whether
Mallees or not, and Angophoras, from the standpoint that the
Mallees have rootstocks, is exemplified by the following quota-
tion from a recent paper by Dr. Hall* — "The origin of the
peculiar rootstock of the Mallee can be well seen by observing
the seedling. In nearly all the Eucalyptus seedlings, and also
in the Angophoras, there is developed, especially if growth is
checked, a small woody swelling in the stem at the point of
attachment of the cotyledons. A number of buds will develop
on this, and shoots start from them. Jf the growth of the seed-
ling proceeds in the form of one main stem, this swelling is soon
obliterated, but, in the Mallee, these secondary shoots grow
almost as quickly as the main stem, and so, instead of a tree in
the ordinary sense, we have an enlarged rootstock, from which
spring numerous stems, all more or less of the same size." The
author is here trying to explain Tate's problem of the transient
and persistent basal inflations of the Non-Mallees and the
Mallees. If the small woody swelling at the attachment of the
cotyledons of the seedlings of Mallees is the initial stage in the
formation of a persistent rootstock, is not the woody swelling of
the seedling of Non-Mallee Eucalypts and Angophoras also the
initial stage in the formation of a transient rootstock ?
The following statement from Tubeuf and Smith's "Diseases
of Plants" (p. 299) may be mentioned. " In the Botanic Garden
at Amsterdam, the roots of several species of Eucalyptus ex-
hibited woody tumours from which proceeded outgrowths re-
sembling 'witches' brooms.' 'J hese contained the mycelium of
an Ustilago which produced spores in the cortical tissues." If
the tumours here referred to are of the same kind as those we are
interested in, we consider that the presence of Ustilago under
the circumstances mentioned, is to be regarded as merely in-
dicative of a saprophytic intruder. As pointed out by Erwin
Smith and his colleagues, a varied assortment of lodgers com-
monly infest crown-galls.
* "The Evolution of the Eucalj^pts in relation to the Cotyledons and
Seedlings." These Proceedings, 1914, p,ol7.
204 tumours of eucalypts and angophoras,
Non-Mallee Eucalypts.
Seedlings of some species of Eucalypts are exempt from
tumours. We have met with six of these, referred to later on.
As mentioned in our introductory remarks, tumours, which
accomplish all they can do, pass through the three stages of —
(1) axillary, shoot bearing stem-nodules; (2) composite, shoot-
bearing, stem-encircling tumours; and (3) composite, stem-encir-
cling, shoot-bearing, root-incorporating tumours.
Axillary Stem-nodules. — Five young seedlings, with one or
more pairs of stem-nodules, with the cotyledons, or some of the
leaves in situ, are shown in Plate iv. The cotyledons and lowest
leaves have usually disappeared by the time the nodules become
recognisable, as in the two series of seedlings shown in Plate v.
When they are absent, therefore, it is to be understood that the
opposite and distichous arrangement of the nodul^-s corresponds
to a similar arrangement of the cotelydons and leaves, in whose
axils the nodules develof ed. Figs. Al and A2 are seedlings of
E. corymbosa, each of which has only one pair of nodules. Both
nodules of the older one have a shoot with several pairs of leaves.
Older seedlings of this species are shown in Plate xi. Figs. Bl,
B2, and B3 are seedlings of E. hemiphloia, one with two, and
two with four pairs of stem-nodules. These supplement the
series of the upper row of Plate v.; they are shown here simplj^
because they happen to have one or several leaves in situ.
The upper row of Plate v. shows a gradational series of five
voung seedlings of F. hemi])hloia not old enough to show more
than three pairs of nodules. The three youngest (from right to
left) show the successive development of one, two, and three
pairs of axillary nodules. The first pair of the second seedling-
have made good progress, and are obviously more advanced, that
is older, than the second pair. This is not so evident in the
third seedling, which has a bend in the axis. The fourth ex-
ample shows a further advance. The nodules of the first pair
have grown unequally, so that one is bigger than the other,
though of the same age; while fusion between them has made
some progress. The second and third pairs have made fair
progress. The fifth example shows still further advance.
BY J. J. FLETCHER AND C. T. MUSSON. 205
The first pair have nearly completed their fusion, except
superficially; the second pair are still small; while the presence
of the third pair, in the axils of the second pair of leaves,
can be made out. One shoot has developed on each nodule
of the tirst and second pairs. Another seedling with two
pairs (Bl), and two older ones with four pairs (B2 and B3j are
shown in Plate iv. The first pair of B3 have made good pro-
gress, and have nearly completed their fusion. Another, but a
rather poor specimen which could not be photographed while it
was fresh, with five pairs of stem-nodules, and three root nodules,
is shown on the left of Plate xii. All the specimens of this
series were nursery-seedlings; and we have been able to ascertain,
that seedlings 3-4 months old, with from 4-7 pairs of foliage-
leaves, may or may not have one pair of recognisable nodules.
Bush-seedlings of E. corymbosa, about 3-4 months old, with four
pairs of foliage-leaves, will usually show one pair of recognisable
nodules, as in fig. Al.
The lower row of Plate v. shows a similar series of bush-
seedlings of E. sideroxylon. The younger ones show, in order,
one, two, and three pairs of stem-nodules. The fourth shows
a more advanced stage of three pairs, the nodules of the
second pair being about as large as those of the first. The fifth
seedling has four pairs. One of the first pair has grown more
than its fellow. One of the second pair seems to have come to
a standstill, while the other has grown downwards into the
interval between the nodules of the first pair, and shows signs
of commencing to fuse with one of them. The third and fourth
pairs are still small.
An interesting growing seedling of E. tereticornis, with three
pairs of nodules, and no shoots, is shown in Plate vi., fig 2. The
opposite and decussate arrangement of the pairs of nodules is
well seen. The first pair have developed well, and fairly equally;
and have fused basally, leaving only a superficial interval still to
be filled. The visible member of the second pair- is small, and
seems likely to fuse with the first pair before fusing with its
fellow. A notched, white card has been placed in position to
give a white background.
The total number of pairs of stem-nodules present in seedlings
depends on the number of axils aff'ected, and on the age of the
seedling. Seedlings of some species, as a rule, may have more
15
206 TUMOURS OF EUCALYPTS AND ANGOPHORAS,
than those of other species at about the same stage; but seedlings
of the same species may similarly differ. Occasionally one maj'
find a seedling of a species lia})le to have them, and old enough
to show them, without any at all. One nodule of a pair is often
missing. Apparently, therefore, sometimes the conditions which
lead up to the production of nodules fail, or inoculation was pre-
vented, or did not take place. Sometimes, after the first pair,
the change from a pair of opposite, to two alternate leaves may
result in the appearance of two incomplete pairs.
Plate vii. shows a set of miscellaneous, hardy bush-seedlings
from poor virgin soil, which are remarkable for the numbers of
pairs of axillary stem-nodules present: for the slow progress of
the lower pairs in most of them, considering how many pairs are
present, and consequently for delayed fusions and concrescences;
and for variation in the lengthening of the internodes. The first
five (left-right) are somewhat older than might be expected from
their small size. The first three are E. eugetiwides, the third of
which shows fusions and concrescence of nodules, while the other
two have done very little even in the fusion of nodules of the
same pair. Some of them have shoots. One nodule on the lower
side is missing from the third pair of the first seedling. The
fourth and sixth are JE. j'iperita, and show well, what is very
characteristic of seedlings of this species, numerous pairs, most
of which are crowded up through the non-lengthening of
other internodes besides the first and second. A later stage is
shown in Plate xii., tig.2; but, in this case, matters were com-
plicated by the death of the seedling-stem at an early stage, and
its replacement by two tumour-shoots, as is usual; these aie
remarkable for having pairs of axillary nodules (some stage of
three pairs in each case) the lower ones fairly close together.
Seedlings of this species sometimes have very shapely, large
tumours.
The fifth and seventh are seedlings of E. hcemastoma. One
nodule is missing from each of the fifth and sixth pairs of the
smaller specimen. The larger one shows considerable lengthening
of the third and fourth internodes; also a root-nodule (r.n.) from
which on one side, near the base, a root (r.) emerges, which does
BY J. J. FLETCHER AND C. T. MUSSON. 207
not come out very well in the photograph. The nodules of the
lowest pair furnish a good example of unequal progress.
The remarkable seedling in Plate x., fig.«, has eight pairs; the
seventh and eighth are indicated, but had not completed their
growth when the specimen was secured. This was a pot-seed-
ling, about 2 feet high, and somewhat older than the others
shown in Plates ix. and x. It is the most remarkable, refractory
seedling we have seen.
The remarkable seedling of E. euyenioides, about 4 feet high,
shown in Plate xix., has twenty nodules (possibly twenty-two),
fourteen stem-nodules, in addition to the concrescence, compris-
ing at least three pairs (possibly even four): four pairs are in-
complete, one nodule of each having failed to develop — potenti-
ally about twelve pairs. The nodule marked 1 is solitary, and
was about to fuse with the concrescence. The rest were not
likely to have accomplished anything whatever, even in the way
of fused pairs. Another, fine, but much younger specimen of
the same species, about 2 feet high, has twelve pairs, but the
uppermost are not very far advanced, and there are indications
of some others to follow; the first three pairs have nearly com-
pleted their concrescence. This specimen was obtained too late
to be included in the series shown in Plate vii. For its total
number of complete pairs, this is the most remarkable seedling
we have seen.
Sometimes, but rarely, an extra nodule makes its appearance,
so that instead of the first pair, there is a whorl of three. When
this happens, the second node may, or may not, also show a whorl
of three. We have four seedlings, two of which show one whorl
of three; and two have two whorls of three. 'J he explanation
of this condition is that the cotyledons of both Eucalypts and
Angophoras occasionally show anomalies, such as three cotyle-
dons, fused cotyledons, or with one cotyledon partially or com-
pletely "split." When this happens, the first pair of leaves may
be normal, or dislocated, or replaced by a whorl of three leaves.
The significance of these anomalies, for our purpose, is the pro-
vision of an additional one or two axils for the collection of soil
or dust; or the loss of an axil.
208 TUMOURS OF EUCALYPTS AND ANGOPHORAS,
Plate xi. shows a series of seedlings of two of the Bloodwoods
{E. corymhosa and E. eximia) which differ from any other seed-
lings of Eucalypts that we have seen, in that they have, usually,
only one pair of effective, axillary nodules, though there are
sometimes two, and occasionally more in late stages; but they
almost always have an unpaired, not definitely located nodule
below the level of the cotyledons. Figs. 1 and 2 represent two
young seedlings, somewhat older than the two shown in Plate iv.
One of them has only one axillar}^ nodule; we have other seed-
lings like it. Both are too young to show the unpaired nodule.
When it does appear, it is sometimes much closer to the axillary
nodules than at others. Occasionally, it is far enough away to
indicate that it possibly developed on the taproot, rather than
on the hypocotyl. It seems to be probably a case of wound-
infection due to a possible tendency of the bark of these young
seedlings to crack in growing. The nodule extends upwards,
fusing harmoniously with the fused, axillary stem nodules; and
grows downwards also, partially, or by degrees, entirely encircling
the axis. The fusion usually ends abruptly. It may be an inch
long, or four inches, and in one case about eight inches. In late
stages, it increases in diameter, but continues to be cylindrical
in shape. One specimen has two unpaired nodules, one below
the other. The lower one is evidently a root-development.
The presence of more than one pair of axillary nodules is
unusual in E. corymhosa. But we have one advanced seedling
which has five pairs [first pair (one with a shoot) fused, and the
fused pair fused with the unpaired nodule, as usual; one nodule
of the second pair with a shoot, and fused with the concrescence,
its fellow very small, and solitary; the nodules of the third,
fourth, and fifth pairs small, unequally developed, and not fused].
One seedling of E. eximia (Plate xi.. No. 6) has two pairs of
axillary nodules, which is unusual, (though we have another like
it) and a pair of root-nodules; but, apparentl}', the unpaired
nodule is absent in this case, unless it is represented by one of
the root-nodules. Both root-nodules have incorporated a root,
but that of the smaller one was accidentally broken ofi' short,
and its stump is hidden from view.
BY J. J. FLETCHER AXD C. T. MUSSON. 209
In seedlings of some species and in some individuals of the
same species, the axillary stem-nodules appear sooner, and pro-
gress faster, than is the case in others, or the period of incuba-
tion is shorter Sometimes this may be due, in some degree, to
unfavourable conditions; but in other cases it seems to be a con-
stitutional matter. Hence, it is possible to distinguish between
susceptible, and resistant or refractory species, or individuals.
The majority of the species whose seedlings are known to us, are
susceptible, some more so than others The best examples of
refractory species that we have met with are E. robusta and E.
lo7i(/i/olia, of both of which we have seen nursery-seedlings in
quantity; but we have been unable to get bush-seedlings. Ad-
vanced seedlings of these two species are shown in Plates ix and
X. The seedling of E. robusta, with eight pairs, (Plate x., fig. a)
is the most remarkable refractory seedling we have seen.
Axillary nodules develop not only on the stems of seedlings,
but also on tumour-shoots, showing that the first few leaf-axils
of tumour shoots repeat the conditions («ffered by the early axils
of seedling-stems. Fig.3 (not numbered) of PI. vi., shows the
tumour of a seedling of E. resini/era, whose stem, with one pair
of unincorporated nodules, and any shoots that may have been
present, was scorched and killed b}- a bush-fire. Two tumour-
shoots, as is usual in such a case if the seedling stem is centrally
situated, replaced the stem; these were about 7 and 10 inches
high, and were doing well when the specimen was taken. Both
have the first pair fused with the tumour. The larger has a
second pair, unfused, and unequally developed. The smaller has
a second pair, one only of a third pair, a fourth pair, and also a
fifth pair. [The numbers on the Plate indicate only the con-
spicuous ones; the fifth pair of the smaller one, not being very
well shown in the photo, was not numbered]. The tumour itself
may represent probably three fused and concresced pairs.
Another ca.se is shown in fig. 2 of PI xii. This also is a case
of two tumour-shoots taking the place of a seedling stem after
injury, not by fire in this case. Each of them has three pairs of
axillary nodules.
Still another case is illustrated in PI. xviii. The large tumour-
2l0 TUMOURS OF EUCALYPTS AND ANGOPHORAS,
shoot at the back, a little to the left of the stem, has six nodules,
of which four are visible — the first pair not fused; the second
pair dislocated, probably due to a pair of leaves becoming alter-
nate; one only of a third pair, and one of a fourth pair, with a
shoot. Another tumour-shoot, hidden in the photo, also has a
good pair.
These three seedlings show, then, that axillary nodules may
develop on tumour-shoots which develop in the ordinary way, or
on second-growth shoots, whose development has been accident-
ally stimulated.
The Nodule- Shoots. — Until the morphology of the seedlings,
and the histology of the nodules and composite tumours have
been investigated, one can attempt to interpret macroscopic
characters only with reserve. The shoot-bearing character of
the nodules is due to the fact that the axillary outgrowths take
over and incorporate thedormant buds (or the bud-forming tissue),
which, otherwise, would be smothered. But so many shoots
sometimes develop on a single nodule, or on a fused pair, that we
are inclined to think that the nodules sometimes carry away
bud-forming tissue, rather than simply differentiated buds; and
that the stimulus which is responsible for the proliferation of the
cambium to form the nodules, may also cause the bud-forming
tissue to proliferate, and spread.
On the other hand, sometimes the growing nodules fail to take
over the buds. Three good examples are shown in the two
Stringybark seedlings in PI. xii. Another is shown on the stem
of the seedling of fig.l of PI. xiv. Also two others in the re-
fractory seedling, c2 of PI. x.; for the two futile nodules of the
third pair, and the two branches just above them, belong to the
same pair of axils. There are some other examples on the re-
markable seedling in PI. xviii., particularly the pair of which
one is numbered 4. In such cases, the bud and the nodule
develop separately; if the bud perishes, the nodule is left stranded
on the stem. If both develop successfully, it is noticeable that
the nodule is usually on the outer side of the branch in the ex-
ternal angle between the branch and the stem, fused to both.
This suggests that, as the nodule develops on the outside of the
BY J. .1. FLETCHER AKD C. T. MUSSOX. 211
bud, and between the latter and the attachment of the peticle
of the leaf, the supposed weak spot in the axils, which makes
inoculation possible, may arise in connection with the dijBferentia-
tion of the bud; or that there may be a weak spot about the
attachment of the petiole, which is exposed when the leaf ex-
pands; or, perhaps, both are concerned, ►^erial, transverse, and
longitudinal sections are required to settle the matter.
In general, the conditions which favour or retard the develop-
ment of ordinary branches in leaf-axils, favour or retard the
development of nodule shoots; and, particularly, sufficient room
to allow of free exposure of the plants to light: and damage to
the growing-point, or removal of the greater part of the stem.
Defoliation, under some conditions, may also stimulate the pro-
duction of nodule-shoots.
Two very young seedlings of E. hemiphloia, with only the first
pair of stem-nodules present, are shown in figs. CI and C.2 of
Plate iv. One shoot promptly developed on each nodule after
the growing-points suffered injury, in each case. Sometimes two,
or even three, shoots will develop on at least one nodule of seed-
lings like these.
PI. xxv., fig.l, shows a flourishing pot-plant of E. tereticornis,
growing under very favourable conditions, which exhibits pro-
fuse branching, as well as nodule-shoots.
An advanced pot-seedling of E. hemiphloia^ with a well-devel-
oped, complete encircling tumour, without any shoots, was unin-
tentionally neglected, and the pot allowed to become so dry, that
the leaves wilted, and were cast off. On receiving attention, the
plant revived; but, before the new leaves appeared, five young-
shoots promptly developed on the tumour.
Though the buds usually remain dormant on the nodules of
refractory seedlings like those shown in Plates ix. and x., injury
to the growing-point, or removal of the greater port of the stem
will cause shoots to develop.
The presence of shoots is of great importance both to the
stem-nodules, and to the resulting, composite tumours, for
they are not then entirely dependent on the seedling-stem for
nutriment. A large, revived, belated nodule (A2), whose fellow
212 TUMOURS OF EUCALYPTS AND ANGOPHORAS,
is missing, is shown on the advanced seedling in PI. xix. The
seedling-stem (s.st.) perished, whereupon the main nodule-shoot
took up the running, and this gave the nodule a chance of
making a fresh start. This is the largest, individual nodule we
have yet seen. The photo is reduced by somewhat more than
one-third. Composite tumours at first are usually more or less
pyriform in shape, especially if they have no shoots, or only
insignificant ones (PI. xii., fig. on the right). Some of the later
stages of the B. sideroxy Ion-series show how the upper portion
of the tumour fills out when there is a good series of shoots
round the summit. As long as the shoots last, and continue to
grow, the composite tumours may be expected to progress pro-
portionately.
The number of nodule-bearing shoots on a seedling, like the
number of shoots on individual nodules, if the conditions are
favourable, is sometimes surprising. The oldest nodules are
likely to show ihem first, and to have most shoots. The basal
pair, or only one of them, will very often show one or more,
when the others are without them. But the nodules of any
pair, or one of them, may have shoots, when the others have
none. Both, or one, of several pairs may have at least one shoot.
Or the buds may remain dormant, and no shoots at all may
develop, as in most refractory seedlings.
It would be unusual to find more than one branch in a leaf-
axil, though, of course, there may be reserve-buds. Three shoots
are often present on a single nodule, but there may be as many
as seven, or even more, in difi'erent stages. Fig. 1 of PI. vi., shows
a very attractive bush-seedling of Anyophora lanceolata, pro-
bably not under two years old, photographed while fresh. As
shown, it is less than half the natural size, the stem-height
above the encircling tumour being about 10^ inches. The
tumour is the result of the complete fusion of the first pair only.
The nodules of the second pair, still ver}^ small, unfused, and
without shoots, are to be seen, rather indistinctly, a little above,
but there is no third pair. Eight shoots are present, four on
each side, but two of one group are dead.
An inspection of the figures of the E. sideroxylon-sevies will
fiY J. .1. FLETrHEK AND C. T. MUSSOX. 213
give an idea of the way in which shoots develop about the
summit of the tumours, when they are complete in late stages
When specimens like these are scorched by a bush-fire, and the
stem, and any shoots that may be present, are killed, provided,
of course, that the underground portions, including the tumours,
are not hopelessly injured, fresh second growth shoots may come
up freely, when the conditions become favourable. Kig.3 of PI.
vi.. is an example of a scorched seedling. Two second-growth
shoots only are present here. The growth-habit of such plants,
especially if they are scorched a second time, or oftener, and
recover, if they succeed in attaining any size, necessarily is much
modified. In the quotation from Mr. 'J epper's paper, the second-
growth shoots of the Mallees, under similar circumstances, are
mentioned.
The shoots of all the pairs except those which supply them at
the summit of late stages of the encircling tumours, are doomed
to perish at an early stage, except under exceptional circum-
stances, because the tumours are gradually pulled underground.
Even those on the summit cannot last indefinitely, as a rule,
unless water-storage roots are incorporated.
The nodules, when quite small, are smooth. As they increase
in size, the surface becomes warty, due to local proliferation. As
the warty protuberances increase in size, they meet and fuse, and
fill up the vacant spaces. In this manner, by constant local
proliferation at the periphery, the nodules, or the composite
tumours to which they give rise, increase in size. The latter,
when doing well, are excessively warty. This is indicated in
some of our illustrations, but the warts do not always show up
as conspicuously as they do in the specimens.
The nodules and the surface of tumours, when fresh, are readily
sliced with a knife; but, when dry, they are hard and woody.
The tumours when fresh, and doing well, contain a good deal of
of moisture; but, when drying, gaping, longitudinal cracks, often
from top to bottom, appear.
Later Stages. — Growing nodules very readily fuse with any
other suitable living tissue with which they make contact; so
214 TUMOURS OF EUCALYPTS AND ANGOPHORAS,
that finally, when the fusion is complete, there is no visible line
of demarcation. As the nodules of a flourishing pair progress,
they increase steadily in size, growing upwards slightly, but
more evidently outwards, backwards, and downwards, until they
meet and fuse, thus encircling the stem. Similarly the fused
pairs grow downwards, covering up and fusing with the portion
of the stem involved, until the several fused pairs have con-
cresced. As the hasal portion of the concrescence grows down-
wards, it finally encircles and fuses with the upper portion of
the taproot, and the proximal portions of any lateral roots that
it may encounter. We have an example of two seedlings grow-
ing so close together that the encircling tumours came into con-
tact and fused. If two such seedlings survived and attained
tree-size, they might furnish an example of apparently one tree
with two stems.
The opposite and decussate arrangement of the stem-nodules,
corresponding to the disposition of the cotyledons and leaves, is
ati ideal arrangement for the production of well-balanced, sym-
metrical, composite, encircling tumours, provided— (1) that all
the pairs of stem-nodules are complete; (2) that the nodules
develop promptly; (3) that they grow comparatively equally and
uniformly, and make the necessary fusions, and the fused pairs
the necessary concrescences, at the right time, and in the right
way; and (4) that the internodes, especially the lower ones, do
not lengthen too soon or too much. But if one or several of
these provisoes fail, the final result will be correspondingly modi-
fied. If plenty of material is available, very suggestive and
instructive anomalies, of almost every conceivable kind, may be
obtained.
Incomplete pairs of nodules are common. If several nodules
or pairs are missing, any resulting composite tumour will be
correspondingly smaller. A good example is shown in PI. xiii.,
fig.2, of the E. sideroxyloii-sevias (about half nat. size). This is
a concrescence of the fused first pair and of one nodule only of
the second pair, and this, though it is included, did not make
much progress and contributed very little, and is still recognis-
able (in the specimen though not in the photograph).
BY .7. .t. FLETCHER AND C. T. MUSSON. 215
One nodule missing from the first pair, which is chiefly re-
sponsible for the basal portion of the encircling tumour, is a
fruitful cause of the production of lopsided tumours. An ex-
ample is shown in PI. xiv., fig. 3, in which the left nodule of the
first pair is missing. Unfortunately the photograph is slightly
blurred. A missing nodule from the uppermost pair, may inter-
fere with the symmetry of the summit of the tumour. We have
a seedling with three incomplete pairs only, one nodule from the
top and the bottom pairs on the same side being missing. At
the best, all that these could give rise to, is a half-encircling
tumour.
Fig. c of PL viii., appears to be a case in which one nodule of
the first pair (on the left) did not develop. The seedling-stem
having been broken off, a normal branch in the axil without a
nodule then assumed the erect position, and took the place of
the seedling stem. The single nodule present, made fair pro-
gress. It shows indications of about a dozen shoots, some of
them dead. The large shoot, just to the left of the stump of
the stem, belongs to the latter.
Sometimes a nodule, instead of growing downwards and fusing
with the stem, will grow downwards and outwards, away from
the stem, even in erect seedlings. Three bent seedlings are shown
in PL viii. They have stem-encircling tumours, resulting from
the fusion of the first pair, which has grown outwards and
downwards away from the stem. There is only one pair in fig.a.
Figs. 61, 62, show some indication of others poorly developed.
There are no incorporated roots in these specimens, nor, we
think, was there any likelihood of its happening. In arranging
the specimens for being photographed, some of the lateral roots
were unintentionally left in unnatural positions.
Inequality in the comparative rate of growth of the nodules
of the same pair, as well as in the comparative growth of suc-
cessive pairs, is also a fruitful source of anomalous developments.
One nodule of a pair may come to a standstill, while the other
goes on growing; or both may progress, but one of them faster
than its fellow (as in No.7 of PL vii.). Nodules of the same pair
may fuse sooner on one side than on the other; or they may fail
216 TUMOURS OP EUCALYPTS AND ANGOPHORAS,
to fuse at all on one side. Encircling tumours, as they grow
downwards, may progress faster on one side than on the other.
If the nodules do not develop promptl}-, while the tissues are
quite young, that is, if the period of incubation is longer than
usual, the nodules seem to make slower progress, when they do
appear. Meanwhile, the lower internodes may have lengthened
to such an extent that the concrescence of pairs may be delayed
or even prevented. Angophora seedlings frequently oflfer very
characteristic examples of this kind of thing.
The E. sideroxi/lon-series (PI. v., lower row, and Pis. xiii.-xvii.),
of which the earliest, the last, and as many intermediate stages,
as space will allow, are shown, are intended to indicate the course
of events in a susceptible species, W ith the exception of the
tumour in PI. xvii., all the photographs were taken when the
specimens were fresh. They are variously reduced, from about
one-third to one-half, or even more in the larger ones. They are
intended to show the composite tumours, with recognisable por-
tions of the seedling-stems and tumour-shoots, and, especially,
various stages in the capture of lateral roots.
The smaller of the two crossed seedlings (PI. xiv., fig.2) shows
the first and second fused pairs of nodules concresced; the third
pair have not yet completely fused; one nodule from the fourth
pair is missing. 'Ihe growing-point was injured. The taproot
is caught between a lateral root and the tumour, so that it can-
not be freed without trimming the latter. There was enough
soil between the two tumours to prevent fusion. The larger
specimen has lost its seedling-stem, and two of the tumour-shoots,
one on each side, are taking the lead. No roots had been
incorporated in either specimen.
The seedling in PI. xiii., fig. 1, with a stem-height of 20 inches,
has three pairs almost completely concresced, and two pairs not
yet fused. Two roots are just incorporated, and another was
ready to be.
The two seedlings on the right (PI. xiii., tig. 3) show incom-
plete concrescences at the top; and an early stage of the incor-
poration of a root.
Fig.2 of the same Plate (reduced by nearly one-half; is a fine
BY J. J. FLETCHER AND C. T. MUSSON. 217
seedling, with a stem-height of 37 inches, the two longest tumour-
shoots about 13 inches. It is a good example of a small tumour
resulting from a shortage of axillary nodules. OnU' three de-
veloped, of which the only one of the second pair (hidden at the
back of the photo) made little progress, and contributed practi-
cally nothing to the tumour. Small though it is, the tumour has
captured three good, lateral roots Two others, which were
accidentally broken off, and whose bases are hidden by the
lateral root on the right, were about to be captured. The seed-
ling-stem had a good canopy of foliage, and had got well ahead
of the tumour-shoots, so that it would probab y have got rid of
the latter sooner than seedlings with larger tumours.
The seedling in fig. 1, PI. xiv., (reduced by about one-half) has
a concrescence of probably two pairs, with three incomplete pairs
above (one of which is bidden). The downward growth of the
concrescence on one side^ is much in advance of that of the other,
as shown on the left. The uppermost is a good example of a
belated nodule which failed to get possession of the bud; both
it, and the branch above it, belong to the same axil. Several of
the shoots have done well. Two lateral roots have been incor-
porated; and several others would ha\e been later on.
Fig.3 of PI. xiv., is an example of a lopsided tumour due to the
absence of one nodule of the tirst pair. Unfortunately the photo
is slightly blurred. It has not incorporated an}^ roots. Shoots
are numerous. The seedling-stem may possibly be missing.
Fig. 1. of PI. XV., is a very complete concrescence (reduced by
somewhat over one-third). Two lateral roots are just incorpo-
rated.
Fig. 2 of the same Plate (only slightly reduced) has the con-
crescence incomplete behind except at the top. One good root
has been incorporated on the right. The root on the left with
a bifurcation, and the one above it, would have been incorporated
if the concrescence had been complete behind.
Fig.3 of PI. XV., is a good example of a late stage.
Fig.l of PI. xvi , shows a fine seedling, with a stem height of
5 feet above the tumour. The latter is shown (nat size) in PI.
xvii, The strongly developed taproot is now well established,
218 TUMOURS OF EUCALYPT8 AND ANGOPHORAS.
the lateral roots being insignificant by comparison. The seed-
ling-stem has been able to increase its initial lead, and to develop
its upper branches, so that their crown of foliage fairly completely
overshadows the tumour-shoots, preparatory- to getting rid of
them by the process of natural pruning. The lateral roots are
so insignificant that the tumour must be largely dependent on
the seedling-stem for most of its water. When the leaves of the
crown of foliage belonging to the seedling-stem and its branches
have overshadowed the tumour-shoots, and are transpiring freely,
the latter seem to suff'er in consequence. When the shoots have
been got rid of, the tumour, as such, gradually comes to a stand-
still. The tumour of this example is shown in PI. xvii. (nat. size).
and is a fine specimen.
Fig. 2 of PL xvi., shows the size of two, more advanced seed-
lings, 6-7 feet high, side by side. The one on the left, which had
got rid of its tumour-shoots, was dug up (inadvertently the tap-
root was cut off" rather too short), and held beside a slightly
taller, undisturbed specimen, which had one shoot left. This
was afterwards uprooted and examined. It will be noticed that,
at this stage, the tumour is completely out of sight underground.
These photos were taken early in October, 1914, at which time
the plants were about six years old. Three and a half years
later, other examples of the same batch as those in fig. 2, were
promising saplings 9-12 feet high.
The last stage of an advanced seedling of E. tereticorids is
shown in fig. 2 of PI. xxi.; this has one small shoot left. The plant
was uprooted carefully, placed on a stump, and photographed
immediately. It was growing near the specimens shown in PI
xvi. In this case, the seedling-stem appears to be excentric,
consequent upon lopsided growth of the tumour: and the shoots
were all on one side. One nodule was not incorporated in the
tumour. The difi'erence in size between the taproot and the
incorporated or other lateral roots is well shown.
PI. XX., shows another but older stage of the same species; but,
in this case, matters are complicated by the death of the seedling-
stem. A few inches of the latter were in situ when the plant
was found, but were accidentally broken ofi" in packing-up a
BY J. J. FLETCHER AND C. T. MUSSON. 219
number of specimens. The stump of it is just discernible in the
photo. There were a number of shoots, but, in this case, one of
them took the lead, and was 6 feet high. At the base of it is a
remarkable, encircling insect-gall. At first, we took this to be
a case of the fusion of pairs of axillary nodules on a tumour-
shoot, followed by concrescence; and that it might bean example
of what Tate meant, when he said seedlings and ''growths" have
basal inflations. On cutting it tiansversely, four radial cavities
were exposed, two of which contained living coleopterous pupae,
one in each cavity; the other two contained only exuviae or
excrement. Apparently, the mother-insect oviposited at intervals
all round the base of the shoot.
Of the two advanced seedlings of E. eugenioides in Plates xviii.
and xix., the first has not yet got rid of the tumour shoots, which
are situated at the back, as the specimen is shown. The other
one had got rid of them; in this case, the death of the seedling-
stem gave rise to complications.
Tate, in his brief way, said that Non-Mallees, which have
basal inflations, grow out of them. Apparently, he was relying
on experience, and not speaking from observation. The only
author who mentions what takes place, as far as we can find, is
Ednie Brown, who in his "Forest Flora of S.A.", under E.
leucoxylon (Part ii.), says — "During the first two years of its
growth, the plant has a low-lying or spreading habit, not at all
prepossessing in its favour, when looked upon in the light of a
future timber-tree. About the third year, however, a straight
and upright 'leader' comes away from the centre of the apparent
bush, which, after this, soon assumes the form of a promising
young tree.''
What is here described as a straight and upright leader, is
simply the seedling-stem coming into its own, and getting rid of
the tumour-shoots. Otherwise, it the seedling-sttm is destroyed,
two, strong leaders usually come away; that is two, tumour-
shoots take the lead, as we have already pointed out, and as is
shown in several of our Plates. 'J he author was apparently not
aware of the presence of what Tate, some years afterwards, called
the basal inflation of seedlings of this species. We should think,
also, that he probably underestimated the age of his plants.
220 tumours of eucalypts and axgophoras, "
Mallees.
The Mallee-Scrubs of the interior, where alone fruitful in-
vestigation can be carried out, are remote from, and inaccessible
to us; and, consequent!}^, our material for the study of the
tumours of this group is very incomplete. Nevertheless, there
is something to be learnt from it We regret that we have been
unable to procure seedlings of the big Mallees, such as E. oleosa^
E. dumosa^ and several others
Plate XX., shows four advanced seedlings of three species.
Figs. \a and 16 are seedlings of E. f^tricta from the higher part
of the Blue Mountains (about 3,000 feet), 2^-3 feet high, from a
batch of seedlings, that we had had under observation for three
years, from the time when they were too young to show nodules.
The younger one (la) has five pairs of axillar}- stem-nodules, the
concrescence of the three oldest pairs not yet completed. Ihe
lengthening of the third and fourth internodes has kept the two
upper pairs apart. Another example, about the same size, has
a more complete concrescence of three or four pairs, then two
incomplete pairs separated by the lengthening of the internodes;
and above, two pairs, close together, followed by an incomplete
pair; but all these are small, and there are no fusions among
them. The older one (16) has a more complete concrescence in
the aspect shown; but one nodule of the first pair is missing (at
the back), and the encircling tumour is consequently lopsided,
and smaller than it otherwise would have been. Three and a
half pairs seem to be present, and a fifth, small pair, doing little,
are indicated at a higher level. One shoot is present, and one
root was in process of incorporation.
Fig. 2 is a seedling of E. Moorei, about 2 feet high, also from
the Blue Mountains, The concresced portion, not quite complete
at the summit, represents three or possibly four pairs. The
uppermost pair is incomplete, the nodule at the back missing.
We have younger seedlings of both species in various stages with
up to five and six unfused pairs. We are inclined to think that
the slovv progress of t}ie nodules of the seedlings of these two
species is, in some degree, due to the low ground-temperatures
and frequent frosts on the higher part of the Blue Mountains
during the months of April-October.
BY J. J. FLETCHER AND C. T. MUSSON. 221
Fig. 3 is a seedling of E. fruticetonmi from Wyalong, for
which we are indebted to Mr. Cambage — the unh' seedling he
could find. We cannot make out the presence of more than one
pair of stem-nodules, which fused on one side (in front, in the
photo), but not on the other. The interval separating them at
the back is about \ inch wide; and there is a depression at the
top which is suggestive of the loss of a branch. The tumoui'
had about eight shoots, and there are three branches low down
on the stem. No roots had been incorporated. One specimen
is not enough to enable one to Judge whether this is merely an
exceptional case, or whether one pair is the usual number for
this species. The " Mallee-root " shown in PI. xxiv., also from
Wyalong, likewise has a tumour of one pair not fused on one
side. We do not know the species, but it may perhaps be E.
fruticetoriim: and represent the latest stage of a seedling like
Fig.S.
Plate xxi , fig.3, shows the most satisfactory example we have
had from Wyalong (much reduced). This is a specimen of one
of the small Whipstick Mallees {E. viridis) locally called Blueleaf-
Mallee. For the stage at which it has arrived, it may probably
be regarded as a fair example of a susceptible Mallee It is not
as plump as it might be, but it was collected in a drought. It
was trimmed for convenience in transmission to Sydney, but the
following dimensions may help to understand its importance :
stem-height above tumour, about 6^ inches; diameter of stem
above tumour, ^; length of tumour, about 3|; diameter, 1|;
diameter of taproot below tumour, y^^. The best aspect of it is
shown; at the back, the downward growth is not so satisfactory
as in front. The seedling-stem appears to be excentric because
of the smaller development on the right side. It has two
branches low down; and there are three tumour-shoots. Four
lateral roots have been incorporated; but they are insignificant
in comparison w^th the well-developed taproot. All the stem-
nodules have been included; the concrescence may well represent
five pairs; there is a cavity near the summit, which appears to
have been caused by borers.
Fig.l of PI. xxi., shows the youngest of four examples of E.
Behriana, received from Wyalong. Unfortunately, the stem of
16
222 TUMOURS OF EUCALYPTS AND ANGOPHORAS,
every one of these was hopelessly damaged; and consequently
the specimens are not typical examples, but they are all we can
get. The youngest ( x ^ about) has the seedling-stem broken off
short, just above the level of the second pair of stem-nodules, one
of which is now missing. The nodules of the first pair are back
and front, and unfused. The posterior one has one shoot. The
front one also had one, but this was broken off short. The
surviving nodule of the second pair developed five shoots, the
largest of which has two branches, and made good progress. It
has fused with the nodules of the first pair, and has grown down-
wards to much below the level of the first pair. This nodule and
its shoots w^ere keeping the plant going.
Two older examples differ in age. The smaller has half an
inch of the stump of the stem left. There appear to be two
pairs of nodules. The first pair inade some progress, but did
not fuse. One of them has three shoots, two of which are 5- (5
inches long. This nodule was keeping the plant going. One of
the second pair fused with the nodule with shoots; the other
made little progress. The bark on the nodules is very thick, and
adherent.
The larger one is more difficult to understand, as the stem is
almost completely missing. There are shoots up to 10 inches, in
two places, on the margin of what is left. There is one pair of
futile nodules, one much better developed than the other, on
opposite sides of what is left of the stem, which is about 1 inch
thick. Both this specimen and the preceding one have a long
piece of the root intact. The smaller one has two, nearly oppo-
site, lateral roots, and one above and one below these, not far
away. But the larger one has three promising lateral roots at
different levels, two on opposite sides of the taproot, and a middle
one in a direction at right angles. An encircling tumour would
have to make considerable growth before incorporating all of
them.
For the large specimen shown in PI. xxiii., in the natural
position, as we think (rather than view^ed with the taproot
vertical), we are again indebted to Mr. Cambage. The best
aspect of it is shown. A farmer, in grubbing up the plant, with
one blow from his axe, delivered just behind the base of the big
BY J. J. FLETCHER AND C. T. MUSSON. 223
shoot shown in the figure, split off a large piece at the back, on
which there was probably another shoot, situated on the right
of the base of the dead seedling-stem. At the back, just to the
left of the mark X on the photo, there is a longitudinal concavity,
showing the site of the missing shoot. The remains of the
original stem, now exhibiting signs of decay, indicate that it must
have been about 3 inches in diameter when it perished. The
taproot is well developed. One lateral root, with a branch, is
seen to the right. This can hardly be said to be incorporated.
At the back, there are four roots or indications of them, one of
which is incorporated, and one partially, while two others are
too low down. The development of the shoot present evidently
stimulated the growth of the tumour all round its base. The
rest seems to have come to a standstill after the death of the
seedling-stem. What strikes us about this specimen, is the
paucity of shoots: and that there is not such a satisfactory
arrangement of lateral roots, at about the same level, as is shown
in the specimen in PI. xxiv. The loss of the seedling-stem doubt-
less complicated matters, and makes the interpretation of this
incomplete specimen difficult.
E. Bekriana is described as being sometimes a Mallee, 8-15
feet high, and sometimes a tree up to about 35 feet high. Our
four specimens seem to us to be more or less refractory cases,
whose root-system is not quite like that of the typical Mallees,
in respect of the main lateral roots advantageously situated for
incorporation in the encircling tumours. If either or both these
conditions occur often, such may be responsible for the fact that
E. Behriana is sometimes a tree. This species is well worth in-
vestigation with adequate material.
Magarey,* in describing the method of the Blacks in obtaining
water from the roots of the Mallees, says — " The roots of these
water-trees run out from the stem for 40 feet to 80 feet, and lie
at a depth only of from 2 inches to 9 inches below the surface.
The position of these lateral roots is frequently marked by a
'rise' or 'bulge' of the soil right over the root. The roots are
* "Australian Aborigines' Water-Quest." Proc. R. Geog. of Aust.,
South Australian Branch, iii., p. 67 (1899). Also Rept. Aust. Assoc. Arlv.
Sci., vi., 1895, p. 647.
224 TUMOURS OF EUCALYPTS AND ANGOPHORAS,
easily raised. A native goes to a water-tree, and tries the ground
at from 4 ft. to 5 ft. from the stem; or if guided by a 'bulge' or
a 'crack,' finds the root at once" (p. 69) "Each such
mallee tree has usually from four or five to seven or eight side-
roots running out from the stem at a few inches' depth from the
surface" (p.70).
Plate xxii. shows a " Mallee-root." We do not know the
species, but it was one of the smaller ones, and may, perhaps, be
E, fruticetortim. It is not a particular!}- fine specimen, but it is
the only one we can get. In the absence of any published illus-
tration of the kind, as far as we know, we make no apology for
showing this one; as there is something to be learnt from it, as
there is from any undamaged "Mallee-root." The specimen was
placed on its side, and photographed from in front. The scale
is shown by the rule at the bottom. We interpret this to be a
case of a not quite completely encircling tumour, resulting from
one pair of stem-nodules which fused on one side (at the top of
the photo), but did not fuse on the other side; hence the breach
in the continuity of the tumour (at the bottom). There are four,
important, more or less horizontally running, lateral, water-
storing roots, two (at the top) coming off very close together;
and two others (below) at a fairly wide angle. They are almost
on the same level round the taptoot. The tumour, composed of
only one pair of nodules, was unable to encircle any of them
completely; but it fused with the upper part of all of them,
sufficiently, probably, to tap the water. The site of the seedling-
stem is indicated by the light spot, which represents the posterior
aperture of the hollow, flattened taproot, of which about six
inches are left. The site of one shoot is well shown, over the
lower lateral root on the right. The rest of the upper part of
the tumour was so badly smashed, that it is difficult to locate
exactly the situation of any other shoots that may have been
present; but, notwitlistanding the fact that the two upper roots
are very close together at their junctions with the taproot, we
believe that there was one over each of the other lateral roots.
It seems, to us^ a reasonable conclusion, that an uninjured, flour-
ishing Mallee will, as a rule, have as many shoots as there are
lateral, water-storing roots wholly or partially incorporated in
BY J. J. FLETCHER AXD C. T. MUSSON. 226
the encircling tumour, and that the former are situated over the
latter.
In an adult Mallee, the shoots and the stem, if present, are
all approximately the same height, and diameter. But in the
seedling shown in PI. xxi., fig.3, the seedling-stem has so far
maintained its initial lead. There must be some intermediate
stage, therefore, when the shoots are able to overhaul the stem.
Possibly this may come about when the lateral, water-storing
roots are sufficietjtly developed, to supply more water in the
aggregate, than the taproot alone is able to do. Such questions
as these cannot be profitably discussed without satisfactory
material, and observation on Scrub plants.
F. Mueller, Maiden, and Cambage, and others, have recorded
instances of Eucalypts of species which are ordinarily Mallees,
sometimes being trees; but we have failed to find any explana-
tion of such cases. From our point of view, it is not difficult to
understand that infection may fail to take place occasionally, or
that, from accidental causes, or because the individuals were re-
fractory, it may have been followed by little in the way of results.
The root-waterstoring arrangements of Eucalypts, whether
Mallees or not, as well as of other Australian plants, in arid
regions, are in need of scientific investigation. Little is known
of this important subject, beyond what explorers, travellers, and
early colonists gleaned from the Blacks, and have put on record.
Naturally the Blacks chose the roots of the big Mallees.
But it is a reasonable supposition, in the absence of scientific
evidence, that the smaller VVhipstick Mallees also stored water,
though not on a scale sufficient for the -Blacks to attempt to
exploit it. Inferentially, all the Mallees must have water-storing
roots, otherwise it is inexplicable how the tumours of the Mallees
can keep their shoots, and persist.
Eucalypts may have water-storing roots, however, without
being Mallees. The only author who mentions this fact, that
we are aware of, is K. H. Bennett. In his description of the
method of obtaining water from Eucalypt roots, as practised by
the Aborigines of the arid country between the Lachlan and
Darling Rivers,* he says "The Eucalypti consist of a gum (the
* These Proceedings, 1883, viii., p. 214.
226 TUMOURS OF EUCALYPTS AND ANGOPHORAS,
largest of the back country trees), a box, and a mallee. The
first-named was the most preferred, as yielding the greatest
quantity." The name of the species is not given, but it is said
somewhat to resemble "the red gum in appearance — the leaves
being narrower and of a silvery colour," and to grow "chiefly on
sandy or light loamy soil, and throws out numerous lateral roots
at a depth of from six to twelve inches from the surface of the
ground." As the Blacks could not get at the v/ater in the water-
charged tumours of the Mallees, it is intelligible enough that
they preferred to operate on the roots of a water-storing Gum.
It is desirable that seedlings of this species should be examined,
so that it may be determined whether they are exempt from
axillary stem-nodules, or refractory; or, if susceptible, how they
are able to get rid of the tumour-shoots. Seedlings of some of
the Eucalypts of Central Australia mentioned in iSpencer and
Gillen's "Across Australia," also are well worth attention, par-
ticularly those of ^. terniinalis and E. eudesmoides, because these,
perhaps, may be examples of water-storing Gums, like Bennett's.
We are not aware that analyses of the root-water of the
Mallees have been published. Magarey mentions that, when
freshly obtained, the root- water is clear, but that, after standing
for some time, it becomes discoloured, and turbid (^.c , p.70). Is
the turbidity merely due to a chemical precipitate on exposure
to the air, or is it a biological phenomenon ?
Another matter that is deserving of consideration is, the pos
sible significance of the abundance of Lerp-manna on the foliage
of some of the Mallees,-at certain. seasons and under certain con-
drtions, as. indicating the presence o£ -some form of sugar in the
sap, likely to be a source of nutriment to parasitic micrabes.
Particulars about its occurrence are given, by Tepper, in the paper
already mentioned; and in the Catalogue of tlte Victorian Ex-
hibition, 1861, Report on Class iii.,-p.25,. under the head of
" Manna."
PI. xxv., fig. 2, and PI. xxvi., show views of two growing
Mallees, with as much soil, as was possible with pocket-knives
and hands, scraped away from the base. The first and second
of the three probabh' represent E. oleosa. They were taken in
the Scrub, 50 miles north of Adelaide, by Mr. A. G. Edquist, to
BY J. J. FLETCHER AND C. T. MUSSON. 227
whom we are indebted for them. The third, in which the hand-
kerchiefs of the party formed the background, is of a Mallee
{E. sp.) at Wyalong, for which we are Indebted to Mr. W. J.
Moffat, who also, most kindly, did his best to obtain seedlings
at Wyalong for us, and who sent us the Mallee-root figured.
The published illustrations of Mallees, that we have seen, fail to
give any idea of what is out of sight underground.
Angophoras.
We have obtained good series of seedlings of four species, A.
cordi/'olia, A. r/ifermerlia, A. laiiceolata, and A. suhvelutina.
They are alike, in that they are somewhat refractory. The stem-
nodules are slow in appearing, that is the incubation-period is
longer than in the Eucalypts we know best. Some bush-seedlings
of a batch, that were over a year old, showed just recognisable
steni-nodules; but others showed nothing at all. Nevertheless,
when they are old enough, it would be surprising to find them
without some. The nodules are not only slow in appearing, but
they grow rather slowly after they do appear. Hence it is
common to find examples on which only the first pair have fused
(PL vi., fig.l); and one of these often grows more than the other,
so that the fusions are lopsided. Meantime, the internodes have
had time to lengthen,- so that, even if there were fusions of the
pairs above the first, there is little chance of concrescences. We
have some advanced seedlings, however, which show large and
complete tumours. Anomalies are common. We have one
seedling with six incomplete pairs, and nothing else. We have
not seen young root-nodules, like those figured in three Eucalypt
seedlings; and we have not seen later stages of them in Euca-
lypts. But we have examples of Angophora seedlings with what
appear to be late stages, unmistakably on the root, and incor-
porating roots.
Though, in this paper, we are confining our attention to Euca-
lypts and Angophoras, we may just mention that we have speci-
mens of about ten species of other genera which appear to be
similarly afflicted. We have been able to get young stages, with
paired, a^^illary nodules, of some of them; but it is difiicult to
get young enough seedlings of the others. We hope to offer
some observations on these on another occasion.
228 TUMOURS OF EUCALYPTS AND ANGOPHORAS,
List of Species, exempt or liable, as far as known.
We give a list of the species of which we have seen seedlings,
or in two cases, of seedlings which have come under the notice
of Mr. Carabage. We offer this list in the hope that biologists
in the other States will investigate the condition of the seedlings
of Eucalypts accessible to them, which we cannot get, and,
especially seedlings of the Mallees; and record their observations.
Group i., Exempt *S'pecies. — Six species are known to us at
present— jE'. oreades from the Blue Mountains, E. pilularis, E.
punctata, E. sp., from the foot of the Blue Mountains on the
western side; together with E. gigantea Hooker {E. Delegaiensis
R. 'V. Baker), and E. fastigata, both from the Federal Capital
Territory [collected by Mr. Cambage]. We have obtained seed-
lings of the first two in abundance; of the third, a fair number;
of the fourth, only one, but it is a fine example. Mr. Cambage
has kindly given us four examples of E. gigantea, and twelve of
E.fastigata, most of them quite old enough to show that they
are free from stem-nodules.
The seedlings of these six species, as far as we have seen, are
exempt from stem-nodules. We prefer to speak of them as
exempt, rather than as immune, until their axillary conditions
have been investigated, and inoculations carried out. The seed-
lings of E. oreades, as well as the two lots of Mr. Cambage's
seedlings, up to the stage presented, are just ordinary seedlings,
inviting no comment. But seedlings of E. pilularis, E. punctata,
and E. sp., as they increase in size, invariably as far as we have
seen, gradually come to show a pyriform thickening of the base
of the stem (of the wood to some extent, but especially of the
bark, as may be seen in transverse sections) over a distance of
several inches, according to age. This is not pathological, and
seems capable of a simple explanation. The capacity of the
root-system of these seedlings for receiving the elaborated sap
seems to be unequal to the capacity of the stem for delivering it;
so that there appears to be a stagnation of the sap about the
base of the stem, such as might be caused by a constriction or a
cincture. The enlargement of the stem ends at its junction with
the taproot, and does not involve the lateral roots. Transverse
cracks appear in the bark, and flakes often come away, when
BY J. J. FLETCHER AND C. T. MUSSON. 229
large specimens are dried, though tlie wood does not crack. But
the large, composite tumours of seedlings liable to them, crack
longitudinally, sometimes almost to the centre, when drying, as
already mentioned. Mr. Cambage's specimens are not old
enough to show this development, if they have it in later stages.
Group ii. — Two l^loodwoods, E. corymbosa and E. exiniia,
which have but one efl'ective pair of axillary stem-nodules, as a
rule; though other unimportant ones may appear in late stages:
but the fusion of this pair, almost invariably fuses with an
indefinitely located, unpaired nodule at a lower level; and the
resulting, composite tumour extends downwards, partially or
entirely surrounding the axis, and incorporating lateral roots.
Group iii. — Species liable to attack, but susceptible or refrac-
tory in varying degrees. This group will include all the Mallees,
and the three species mentioned by Tate (E. viminalis, E. ros-
trata, and E. leucoxylon), of which we have not seen specimens.
' County of Cumberland and the Blue Mountains. — E. hcema-
■Homa, E. tereticornis, E. Deanei, E. saligna^ E. maculosa^ E.
rubida, E. Luehmanniaria, E. squamosa, E. crebra, E. ^^aniculata,
E. siderophloia, E. sideroxylon^ E. amygdalina, E. piperita, E.
hetniphloia, E. longifolia (nursery-seedlings), E. i^esinifera, E.
robusta (nursery-seedlings), E. cajntellata, E. eugenioides, E.
Sieberiana, E. Moorei, E. stricta: and seedlings of sevei-al species
not identified. We have not been able to get seedlings of E.
botryoides and E. maculata.
Berrima and Moss Vale. — ^. amygdalina, E. coriacea, E. dives,
and E. Macarthuri [all collected by Mr. Cambage; who has also
given us two specimens of E. coriacea from Jindabyne, K.S.W.I.
We are indebted to Mr. E. Cheel for a fine late stage of E.
6ieberiana from Hill Top.
Marulan, — E. cinerea (Argyle Apple), and fine seedlings of
three unidentified species locally known as Yellow Box, Broad
Peppermint, and Snappy or Brittle Gum [collected by Mr. H. H.
Solomon].
Wyalong, N.S.W. — E. Behriana, E. fruticetorum, E. viridis
[collected by Mr. W. J. Mofiat and Mr. Cambage].
Queensland.— Axillary stem-nodules have been recorded by
Mr. Cambage on a seedling of E. pallidifolia-* and he has in-
* Journ. Proe. R. Soc. N. S. Wales, xlix., p. 435 (1916).
230 TUxMOURS OF EUCALVPTS AND ANGOPHOKAS,
formed us that he has a plant of E. pruinosa with stem-nodules,
in his bush-house, raised from seed, which he collected in Tropical
(Queensland.
West Australia. — A pot-plant of E. macrocarpa (PI. x., fig. 6)
which died: we are indebted to Mr. A. G. Hamilton for this
specimen. We have seen a flourishing pot-plant of E. ficifolia,
with two pairs of stem-nodulesj but were unable to see how it
compares with the two Blood woods referred to above, in respect
of the presence of an impaired nodule.
New Zealand. — In reply to a request, addressed to a friend
in New Zealand, for information about Eucalypt seedlings raised
from seed there, we were kindly supplied with five, representing
as many species, from the nursery of T. Horton, Ltd., at Hast-
ings or Pahiatua, we are not quite sure which. They are seed-
lings, from 10-18 inches high, of species that we had not pre-
viously seen. Three are labelled E. Gunnii, E. glohulns. and E.
Stuartiana. The first has two pairs of stem-nodules: the first
pair nearly fused, and both have shoots; the stem was broken off
just above the second pair. The second, though it is the largest
seedling of the lot, has only one pair, unfused. The third has
the first pair only, fused on one side. The fourth is wrongly
labelled E. coriacea, as the leaf-characters are different from
those of specimens of this species, from two localities, given to
us by Mr. Cambage. This has the first pair veiy large and
warty, not completely fused; one of the second pair very small;
and a third pair, very small, and unequal in size. The fifth is
labelled " Ked-flowering Gum,"' with an unfamiliar botanical
name that we cannot trace in any list of Eucalypts, known to
us. This has the first pair large, but not fused, one with two
shoots; a second pair, very small, one with a shoot; and the
third pair still smaller.
Th^se are the first and only seedlings with stem-nodules from
outside Australia that we have seen; or that we know of, except
Mr. Clayton Smith's Californian specimens, already mentioned.
The interesting thing about them is, that they show substantially
what we are accustomed to see in Australian specimens. We
have not seen any Tasmanian seedlings, but we shall expect to
hear that seedlings of E. globulus in Tasmania are liable. We
BY J. ,1. FLETCHER AND C. T. >JUSSON. 23 1
appeal to biologists in other countries, where Eucalypts are
acclimatised, to examine seedlings for the presence or absence of
axillary stem-nodules; and to record their observations.
The problem, in which we hope our illustrations (certainly an
advance upon what has hitherto been attempted, though there
is scope for supplementing them), will arouse some interest, is
not a simple problem, that can be solved by anyone single-handed.
Even when the systematic botanist has done his share, the
problem, in its entirety, requires team-work — the active co-oper-
ation of the field-botanist, the phytopathologist who is an expert
bacteriologist, the morphologist, and the biochemist. The time
is ripe for its consideration. The Mallee Scrubs are steadily
vanishing in the more accessible districts of several of the States.
The investigations of Erwin Smith and some of his colleagues,
on Crown-Gall, reported in detail as to technique, the histology
of the tumours, &c., and well illustrated,"^ are available for the
bacteriologist as a starting-point. In addition, there are Erwin
Smith's " Bacterial Plant-Diseases'" (3 vols, already published),
besides his numerous papers on the subject of plant-tumours, as
well as Clayton Smith's paper; so that there is ample literature
to begin with.
In Coville and Macdougall's " Desert Botanical Laboratory of
the Carnegie Institution," in Hornaday's "Camp-Fires on Desert
and Lava," and in Vols. xiii.,and xvi., of Contributions from the
U. S. National Museum, a number of characteristic, North
American desert plants are described, and in many cases illus-
trated. Some are said to have thickened, underground trunks,
or to be shrubs with numerous stems from a single root, or with
several stems clustered at the top of a thick, black root, or with
numerous stems given off from a thickened root. Some of them,
to us, are suggestive of the appearance and habit of the Austra-
lian Mallees. If the seedlings of the most remarkable of th^m
have not been investigated, we would call the attention of
American botanists to the advisability of examining these, in
order to test the current interpretations of the adult condition;
* "Crown Gall of Plants: its Cause and Remedy," and "The Structure
and Development of Crown-Gall: a Plant-Cancer." Bulletin, Nos.213
(1911) and 25o(1912), Bureau of Plant Industry, U.S. Dept. of Agriculture.
2o2 TUMOURS OF EUCALYPTS AXD ANGOPHORAS,
and to ascertain whether any of them are comparable with the
Australian Mallees.
AVe are greatl}' indebted to our correspondents at a distance,
for their kindness and trouble in sending us such material as
they were able to get. But we are specially indebted to Mr.
Cambage, not only for material, but for his valuable help in
identifying a number of our seedlings, and in other ways. We
have also to thank Principal Potts for the opportunity of getting
samples of seedlings of four species ivom seed-beds at the Hawkes-
bury College which have been of great use to us: and to Mr. A.
A. Lawson for help in completing our series of photographs.
Corrigendum. — Page 191, line 31 - /or length read lengthen.
EXPLANATION OF PLATES IV. -XXVI.
Fig.s.Al, A2. — Nodules in axils of eotj'ledon.s {£\ co)'i/inbo-<a).
Eigs.Bl-B3 — Nodules in leaf-axils {E. hemtphloia).
Fig.s.Cl, C2. — Nodule-shoots after injur}' to growing-points of seedhng-
.stems (E. hemiphloia).
Plate V.
(Upper row, right-left). — Gradational series of live young seedlings, with
from one to three pairs of axillary nocUiles {E. hemiphloia).
(Lower row). — Five similar seedlings of E. sideroxyhn.
Plate vi.
Fig. 1. — Stem -encircling tumour, Mith iihooti^ [Awjophora kmceolata): no
roots incorporated yet.
Fig-2. — Three pairs of axillary nodules [E. ftrtticornia).
Fig. 3. — Second-growth tumour-shoots Mith axillary nodules (E. rest iiif era).
Plate Ml.
Miscellaneous, remarkable t^eedlings. (Left-right), 1-3, E. euguiioidts; -i
and 6, E, piperita; (j and 7, E. ha^maftoina; (nat. size'.
Plate viii.
Four anomalous seedlings (a, E. r€'<in.ifera\ bL b'2. E. /lemiphhia; c, E.
fiideroxyloii); nat. size.
Plate IX.
Refractory seedlings [E. robnda^ with from one to six, futile nodules; (nat.
size).
Plate X.
Another series of refractory seedlings. (Left-right); a, E. robusta; b, E.
macrocarpa; cl, c2, E. longifoiia; (nat. size).
BY J. J. FLETCHER AND C. T. MUSSON. 233
Plate xi.
(Left-right), 1-4, E. corymhosa', 5-8, E. eximia; (a, cotyledonaiy, axillary
nodules; .r, unpaired nodule: /•.??,. root-nodule); nat. size.
Plate xii.
(Left-right), three root-nodules, /-.??. {E, hemiphloia); 2, two tumour-shoots
with axillary nodules (E. jjiperita); three examples of failure, on
the part of the nodules, a', to capture the buds, ax. .■^h., {E.
eugenioides).
Plates xiii.-xvii.
E. .^ideroxi/lon-sevies, continued from the lower figure of Plate v. (See
pp.216-219).
Plate xviii.
E. eugenioides: belated, axillary stem -nodules at nine levels, in addition
to the concrescence (a); ( x |),
Plate xix.
E. eugenioides: a belated stem-nodule (A2) and its main shoot, getting
their chance on the death of the seedling-stem (s.st.).
Plate XX.
E. tereticornis: encircling insect-gall on large tumour-shoot (after the
seedling-stem perished); nat. size.
Plate xxi.
Fig. 1. — E, Behriana (sometimes a tree).
Fig. 2. — 'Non-MaWee {E. tereticoi-nis); last stage.
Fig. 3. —A Mallee (E. viridis).
Plate xxii.
Mallees: la, lb, E. stricta; 2, E. Moorei; 3, E. fruticetorum.
Plate xxiii.
Tumour of E. Behriana-, ( x |).
Plate xxiv.
A small ••Mallee-Koof from Wyalong, N.S.W. {E. sp.).
Plate XXV.
Fig. 1. — Seedling of E. tereticornis, with nodule-shoots, and precocious
branching.
Fig.2. — A Mallee {E. oleosa^); photo taken 50 miles north of Adelaide,
Plate xxvi.
Fig. 1. — A nearer view of the basal portion of the Mallee shown in Plate
xxv., fig.2; the soil has been scraped away from the base.
Fig.2. — Basal portion of a Mallee {E. sp.) at Wyalong; the soil has been
scraped away. White background furnished by handkerchiefs,
234
OE BINARY MONTHLY MEETING.
May 29th, 1918.
Professor H. G. Chapman, M.D., B.S., President, in the Chair.
Letters were communicated from Professor S. J. Johnston,
returning thanks for congratulations on his appointment to the
Chair of Zoology in the University of Sydney; and Dr. A. B.
Walkom, returning thanks for congratulations on his attaining
the Doctorate of Science.
It was resolved that the congratulations of the Members should
be tendered to Dr. T. Griffith Taylor, of Melbourne, on the award,
to him, of the David Syme Prize for 1918.
The Donations and Exchanges received since the previous
Monthly Meeting ■(24th April, 1918), amounting to 15 Vols., 43
Parts or Nos., 7 Bulletins, 2 Reports, and 7 Pamphlets, received
from 42 Societies, and two private donors, were laid upon the
table.
NOTES AND EXHIBITS.
Mr. Fred Turner exhibited a specimen of Peniiisetum longi-
stylum Hochst., an Abyssinian grass now acclimatised in the
neighbourhood of Chats wood. More than twenty-five years ago
the exhibitor found this highly ornamental grass near the old
battery in the Sydney Domain, but the plants were afterwards
destroyed during the levelling of the ground, after the removal
of the cannons.
Mr, Froggatt exhibited an immature specimen of the Spinj^
Stick-Insect, Extafosoma tiaratum Macl., from Rollings Plains,
N.S.W., a remarkable example of protective mimicry. Also
specimens of a common grass in the Holbrook district, N.S.W.,
very thickly infested with the larvse of a Cecidomyia fly, allied
NOTES AND EXHIBITS. 235
to the Hessian Fly. Inspector F. F. Foster, of the Stock Branch,
wlio collected the specimens, reports that it is very abundant.
Mr. Basset Hull showed a small collection of Land and Fresh-
water Molhisks, obtained by Gunner F. B. Hull at various points
in France, on the Western Front.
Mr. E. Cheel exhibited a series of specimens. — (1) Seedling-
plants of Hakea dactyloides and Banksia serraUi from Hill Top,
Main Southern Line, and specimens of roots from full-grown
plants of Grevillea montana from Hill Top, and G. punicea from
the Sydney District, to show their peculiar roots, which exhibit
an excessive production of small, fibrous roots, originating, at
intervals along the main roots, in clusters, somewhat resembling
the "Hairy-root Disease of the Apple-tree"; these are being in-
vestigated for Mycorhiza. — (2) Seedling- plants of Tephrosia
grandijlora^ which have a simple ovate leaf, followed by a tri-
foliate leaf, then a pinnate leaf, although only about 2 inches
high; they have comparatively large, triangularly-shaped nodules
on the rootlets, measuring from 5-7 mm. across. — (3) Two forms
of Oxalis cornicidatus, with the following distinctive characters:
{a I Plants with small, pale green leaves, and small, lemon-yellow
flowers : {b} Plants with larger leaves, which, together with the
stems, are of a purple or bronze-green colour, and each petal
more or less spotted or splashed at the base with four purplish
marks. Corolla slightly larger, and of a deeper yellow than
those of (a). It is interesting to note, that Sigeroka Nohara, in
a paper entitled "Genetical Studies on Oxalis" (Journ. College
of Agric. Imperial University of Tokyo, vi., p. 165, 1915), has
come to the conclusion that 0. cornicidata is a composite species,
which, so far as his investigations, extending over a period of
about six years, have shown, includes at least four different
biotypes. The plants common in the Sydney District agree in
some respects with those in the neighbourhood of Tokyo, but
the small-leaved form seems somewhat different from those men-
tioned by Nohara. -(4) Specimens of Ruhus lacinialus Willd.,
commonly known as the "Cut-leaved or Parsley-leaved Bramble,"
from near Raymond Terrace, collected during a recent visit to
236 NOTES AND KXHIBITS.
Newcastle. The plants are fairly common along the roadside,
about five miles out on the Gloucester Road, and do not appear
to have been collected since January, 1882, when the late Mr.
E. Betche obtained specimens at Gloucester. — (5) Examples of
Rosa hracteata Wendl., from near Raymond Terrace. Specimens
identical with the above were exhibited, and recorded from
Kembla Grange, Cook's River, and Armidale (These Proceedings,
1912, p. 137; and 1916, p.633), under the name Rosa turhinata
Ait.; but further investigation shows that, although they are
identical with specimens in the National Herbarium from Mont-
pellier under the latter name, the determination is erroneous. —
(6) Fresh specimens of Styphelia tubiflora, from Cronulla, show-
ing variation in the colour of the flowers, from creamy-white in
some plants, pink and cream in others, and the normal blood-red.
Although the individual plants showed the above-mentioned
distinctive colours, they were growing within a radius of 15 feet
of each other, and were very marked.
Mr. Hedley showed a photograph of the cenotaph in memory
of Major A. C. Innes (ob. 1857), and Mrs. Innes (nee Margaret
Macleay, daughter of Alexander Macleay : ob. 1858j, in the old
church at Port Macquarie. References to Major and Mrs. Innes,
and to their home at Lake Innes, in 1836, will be found in James
Backhouse's " Narrative of a Visit to the Australian Colonies,'"
p.406(1843).
Mr. A. G. Hamilton communicated some particulars about
so-called "glow-worms," probably larvae of a Tipulid fly, found
on damp, rocky ledges on the banks of a creek near Bundanoon.
They are said to be found also in deserted coal-mines.
Mr. Waite, a visitor from Adelaide, called attention to the
first Part of the liecords of the South Australian Museum, at
Adelaide, recently issued.
tl'iinled oti; June 27tlK lOlS.J
The Origin of Yolk in tl.e ova of Ubrius Idlll Kestv.
t: .9-
(Ipiier rnw. v\^\n to left). Five sec(lliu<;>. with fnnii (Hic to three paii-s of nodules
IE. Id inipliloiaj. (Lower row). Five >iinilar seedlings of E. sidcroxijlon.
P.L.S.N.S.W 1918
1. Steiii-onoircling iimiMiir with shoots (A. lanccolata). -1. Three p;nvs ot ii.xhiles (E. tereticonii.sj.
(Lower fiy.) Seeund-gnAvtli tuniuur-shonis witli axillaiy iiofUiles (E. rrsinifrra).
P.L.S.N.SW. 1919.
llofractury seedlings: a, E. robusta : I., E. inuvrocarini ; cl, c2, E. /■■
/■-'. .y/(/(;v.*.c///o/(-.scric>, Loiiclucled
phiiit sliuwii ill I'l. xvi., t\ix. 1 ; (nat. sizcj.
.L.S.N.S.W. 1918.
/i,'. I ii(/rnt<>idrs: helated, axillary stem-nodiiles at nine levels, irrespective of the concrescence, «.
/•;. <,i,l.ni.n,h
Ix-lated stciM-nodulc (A2) and its main sliuot,
of the seedling-stetii Is. st.)
he .l.atli
P.L.R N.S.W. 1918.
E. teretieornis: encircling insect-gall on large tumour-shoot (after the seedling-stem perished)
nat. size.
P.L.S.N.S.W.
S^Skd^d)
Tumour of E. Behriana; (xf),
P.L.a.N.S.W. 1918.
A small "Mallee-Root" from WyiiVma, X.S.W. {E. s]).).
.L.S.N.SiW. 1918.
Fiji'. 1. Xoarer view of basal portion of the Malk'o in preceding Plate.
Fig. 2. Similar view of another Mallee (white hackgronnd furnished by handkerchiefs).
237
iilA'i'
o:n" the seasonal distribution of some
QUEENSLAND SPECIES OF ARCELLA Ehrenbero-.
By C. D. Gillies, M.Sc, Biology Department, University of
Queensland.
(Co'inmiinirafed hi/ Dr. T. Ifarn^y JiJinslmi. )
(With foiu- Text-fio'ures).
(1.) General
The Rliizopodan genus Arcella Elirenberu-, is represented in
the fauna of freshwater pools of Queensland so far investigated,
l)y four well difterentiated forms, viz., A. viihfaris Ehr., A. dis-
coides^hv., A. mitr<Ua T^eidy, and A. artocrea Leidy. On account
of the present condition of Rliizopodan taxonomy, these terms
are to be interpreted in this paper as follows : —
A. vulc/aris Ehr., small, hemispherical forms in which the test
may be either smooth or gibbose.
A. discoides Ehr., flat, disk-like species, with the alveoli of tlie
test very small.
A. mitrata Leidy, test normally polygonal; alveoli conspicuous;
pylome with a double invagination.
A. artocrea Leidy: under this appellation is included a number
of large forms in which, typically, the basal portion of the test
is rim-like. This feature ranges from being well developed to
absent, but, in a series, there is no questioning the identity of
the latter variant in spite of the loss of the rim. The fundus of
the test is hemispherical, and the alveolar markings are con-
spicuous. These forms may be regarded as the Queensland
representatives of the species described l)y Leidy under this name.
A. dentata Leidy, has not been observed, though it has been
searched for. This may be on account of its rare occurrence, for-
238 QUEENSLAND SPECIES OF ARCELLA,
Leidy [(2) p. 177], Kofoid [iD p. 100], and Wailes [(3) p. 129], all
refer to its scarcity.
In this paper are presented the results of a preliminary inquir}^
into the seasonal distribution of the four forms met with in
Queensland. The various collections examined were preserved
in w^eak formalin, and, for identification and counting, an ocular
of medium powder, and a 2 3 objective, usually w^ere found suffi-
cient. In this investigation, any Arcelln-tes't not containing the
organism was rejected.
Data.
(1). Brisbane Botanical Gardens.
These observations, extending from ^^eptember 29th, 1916, to
29th December, 1917, have been made by the monthly examina-
tion of material from a lagoon near the old Bird-House. Gather-
ings were usually taken about the end of each month, but, on
6th October, 1916, and 12th September, 1917, supplementary
material was obtained, and, to avoid as much as possible the
introduction of disturbing factors, the collections were made
near the northern extremity of the major axis of the lagoon,
which is elliptical in shape.
(b)6th Oct, 1916.
Actual No.
counted. Pei'centage.
0 0
0 0
16 16
84 84
100 100
(d)28th Nov., 1916.
0 0
43 86
7 14
0 0
Total ... 50 100 50 100
A.
A.
(a)
vulgaris
discoides
29th Sept., 1916
Actual No.
counted.
1
3
Percentage.
2-5
7-5
A.
mitrata
2
5
A.
artocrea
34
85
Total
40
100
A.
A.
(c) 30th Oct., 1916.
vulgaris ... 0
discoides ... 1
0
2
A.
mitrata
20
40
A.
artocrea
29
58
BY C. D. GILLIES.
239
(e)
28th Dec, 1916.
(f) 30th Jan.,
1917.
Actual No. 1
Percentage.
Actual No. Percentage
A,
vulgaris
74
71-85
84
84
A.
discoides
5
4-85
0
0
A.
tnitrata
20
19-42
13
13
A.
artocrea
4
3-88
3
3
Total
103
100-00
100
100
(g)
28th Feb., 1917
(h)29th March,
1917.
A
vulgaris
37
37
67
67
A.
discoides
1
1
2
2
A.
mitrata
8
8
19
19
A
artocrea
54
54
12
12
Total
Total
100
100
100
100
(i) 2nd
A. vidgaris ...
A. discoides ...
May, 1917.
71
4
71
4
(J)
31st May, 1917
77 ' 77
5 5
A. mitrata
19
19
10 10
A. artocrea ...
6
6
8 8
100
100
100
100
(k)
23rd June,
1917.
(1)
1st Aug.,
1917.
A. vulgaris
41
40-60
31
62
A. discoides
0
0
3
6
A. mitrata
43
42-57
8
16
A. artocrea
17
16-83
8
16
Total
101
10000
50
100
(m)
31st Aug.,
1917.
(n)
12th Sept.
, 1917
A. vulgaris
82
82
35
70
A. discoides
0
0
0
0
A. mitrata
1
1
4
8
A. artocrea
17
17
11
22
Total
100
100
'^
100
240
QUEENSLAND SPECIES OF AHCKLLA,
(o) 28th Sept.
, 1917.
A.
A.
Actual No.
vulyaris ... 3
discpides ... 1
Percentage
21-43
7-14
A
mifrata
3
21-43
A
artocrea
7
50-00
Total ... 14 100 00
Only 14 Aicella? were counted on tliis occasion, consequently
tlie percentage values are not so significant as in the preceding
tables.
(p) 2nd Nov., 1917.
Actual No. Percentage.
A . vulyaris ... G 18-18
A. discoides ... 3 9-09
A.mitrata ... 15 45 45
A. artocrea ... 9 27-28
(q)30t.h Nov., 1917.
Actual No. Percentage.
1 20
1 20
2 40
1 20
Total
33
100-00
100
In figs. 1-4, the lines connecting the values of this date with
those of 2nd November on the one hand, and 30th December on
the other, are broken to indicate that little importance should
be as.sociated with them, as the total number actually counted is
too small to convey much meaning. On this occasion, Arcella?
for counting were extremely rare.
(r) 29th Dec, 1917.
Actual No.
A. vulgaris ... 26
A. discoides ... 5
A. mitrata ... 14
A artocrea ... 5
Percentage.
52
10
28
JO
Total
(2). Locality, Bribie Island.
A . vulyaris . . .
A. mitrata ...
50
100
Date, early in April, 1915.
Actual No. Percentage.
23 60-53
15 39-47
Total
38
10000
6Y C. t». GlLLtES.
241
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24:
QUEENSLAND SPECIES OF ARCELLA,
(3). Locality, Beerburrum. Date, 29th June, 1917.
Two living Arcellae observed, both A. vulgaris.
(4). Locality, Chelmer. Dale, 29th Aug., 1917.
Eight living Arcellte observ^ed, all A. vulgaris.
(5). Locality, Graceville. Date, 29th Aug., 1917.
Twenty-three living Arcellte observed, all A. vulgaris.
(6). Locality, Cooroy. Date, 6th Oct., 1917.
Fifty living Arcellee observed, all A. artocrea.
(7). Locality, West Burleigh. Date, October, 1917.
Twenty-three living Arcelhe observed, all A. artocrea.
(8). Locality, Maryborough. Date, 24th Nov., 1917.
Twenty-two living Arcellffi counted, all A. vulgaris.
/
7-5
A
\
I
i
-\
C5
(,0
\
/
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f<i
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ss
I
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i
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'•-■■•.■•
A
k
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A
I'd
r
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:k
h
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tK^:
'Ms
■■ •■■•. i
>'■'.■ '.■
30
F
!■■* .•'
a
■ I- ■'■ ■
•'.••'..:•
' .'-.• ' i"
■■'■''.'■:
, :> \
(■■':
':.'■:.
IS
h
•.'■.■;'■■•
V i 'J'', I
y -'.-.'
''•■'.".■
K:-:
•.'*:. V'.l
1 ■'.■■['.
IS
/e
k
■■■■]■.:'.
*■■;_- ,•:
.xi-"
:■-'-:■:'
■■.■■'.■■
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y^
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m
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\:--^,:
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:. j ' ;•
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Oct. Nov. Dec Ja-r). Fet. Mir Apl. Miy Jun. Jul. /lu§. Sep. Oct. Nov. Dee.
Fi^. 1. — Percentage -t'luqueuoy of ArcdJa ru/gari.shi Arcellaii fauna. Brisbane
Botanical (hardens: Sept. 29th, 1916, to Dec. 29th, 1917.
(3.) Conclusions.
Figs. 1-4 are graphical representations of the data given in
Table i., and each species is graphed separately. It will be seen
that the polygons are constructed on percentages. In taking
BY C. D. GILLIKS.
243
the observations, when the Arcellse were sufficiently abundant,
100 counts were made; but, on a number of occasions, this was
not done through the scarcity of the genus, the extreme case
being in connection with the gathering of 30th November, 1917,
when only five counts were recorded.
Fig. 1, A. vulgaris. The graph shows two primary minimum
periods (29th Sept. -28th Nov., 1916, and 28th 8ept.-1 Nov., 1917)^
and a long maximum period (28th Dec, 1916-12th Sept., 1917)
with three secondary maxima (30th Jan., 31st May, 31st Aug.,
1917), and two secondary minima (28th Feb. and 28th June,
1917). The highest point of the curve is 84% (28th Jan., 1917),
fOO.
m<l 1
-he 1
owes
>t zero (b
th O
ct., ,
Wth
Oct.
, 28
th ^
ov..
191fc
>)•
qo
%0
1
10.
\
\
.10
r
\
10
/ r
A
/ ^
\
10.
10
/ '
A
A
<:'■
0.
l^
'. ^
:^
^
-v^
x^^
^
^
[s^
Ji
Ox-j-v-;
•'■■■:■:':'.
; i- ■ '•"■
Oct. Nov. Dec. Ja;n. Feb. Ma.r Apl. Ma.y Jun, Jul. AuJ. Sep. Oct, Nov. Dee.
Fig.2. — Percentage-frequency of A. diacoides in Arcellan fauna. Ihisbane
Botanical Gardens : Sept. 29th, 191(3, to Dec. 291 h, 1917.
It seems probable that the primary maximum and minimum
periods are typical for the species in Southern Queensland,
because, in the occasional material collected in the maximum
period Dec. -Sept., the dominant form is A. vulgaris {see Bribie
244
QUEENSLAND SPECIES OF ARCELLA,
Island, CheJiner, Graceville, Beerl)urrum, and Maryborough).*
Fuithermoie, material gatheied during the minimum period
Sept.- Dec, sliows a different dominant form (see Cooroy and
West Burleigh). A feature of considerable importance in this
connection is, that the most southerly and .the most northerly
locality are ai)Out 190 miles apart, viz., West Burleigh and
Maryboi'ough
Fig.2, A. discoides. Tlie values in this graph are mostly small,
a!id, out of eighteen observations, eleven range from 0 5% (6th
Oct., 30th Oct., 28th Dec, 1916; 30th Jan., 28th Feb., 29th
March, 2nd May, 31st May, 28th June, 31st Aug., 12th Sept..
1917). There is only one prominent maximum (86%, 2Sth Nov.,
1916).
Until more data arc (collected in the case of this species, aiid
of A. mitnUa. it is considered premature to draw conclusions,
though there is reason to believe November is a maximuni 2)eriod.
SO
^0
30
to.
10.
Oct. Nov. Dec. Ja/n. Feb. Ma-r. Apl. Miy Jan. Jul. Auj. Sep. Oct. Nov. Dec.
Fi^'.o. — Percentage-frequency (if A. mitrnta in Arcellan fauna. IJrisbanc
IJotanical (Jardens : Sept. -illth. lUlC, to Dec. 29th, 1917.
Fig. 3, A. mitrafa. Three maxima are present (30th Oct., 191G:
28th June, 2nd Nov., 1917) and of these, two (30th Oct., 1916,
and 2nd Nov., 1917) occur during the maximum period of J.
vulgaris. On only one occasion did the percentage fall belo^^
0% (1% 31st Aug., 1917). It is the most consistent form of the
* '24tli Nov. is sntticientlj' near December to V)c included in the maxiunuii
period.
6Y CD. GILLIES.
245
four, as it occurs througliout the year, and its numerical range
is tlie lea.st {ser Table ii.).
Table ii.
fOO
Highest values.
Lowest values.
Range.
A. ruhjariK ...
A. discoidis ...
A. mitrata ...
A. arfocrea
84% (30th Jan.. 1917)
86% (28th Nov.. 1917)
46% (2nd Nov., 1917)
85% (29th Sept., 1916)
0%
0%
• /c
0%
84
86
4o
80
p
%0
\
70
\
CO
A
so
I
t^O
• ■
\
\
\
,
\
A
\
^o
*
\
■•\
• \
?n
-
:i|.
,.:S
\. >
\
/ • • • .'
M
/=
_
>-■'. '■ ■■
■:•.'.•,;•
;■ V
'0
c:^"'
-^'^■V/
0
J
^
^B
■:'■■ '
^^
M
-••
■''.'■■ •'•
:V'. . "',
-:-■,
': '^ ; '\
'':'^'
■jVv
Oct, Nov. Dec. Ja.n. Feb. Mtr, Apl. Miy. Jun. Jul. Au6. Jep. Oct. Nov. Dec.
Fig.4. ^Percentage-frequency of A. arlocrca in Aicellan fauna. liiisljanc
BotanicaKiardens: Sept. 29th. UlKi, to Dec. 29th, 1917.
Fig.4, A. (trtocrea.. Tliree maxima are present ('l^i\\ 8ept:.,
1916, 28tli Feb., 28th .Sept., 1017), the liigliesl point of the
curve being 85% (29t]i Sept., I 916), wliiie the lowest is zero (28th
Nov., 1916) The period September-October seems to be a
dominant one foi- this foi-m in Southern Queenshmd, as it includes
two of the maxima, viz., 29th Sejit., 1916, and 2Sth Sept., 1917;
and in niaterial from Cooroy and West Burleigh (.vee 6, 1 ) gath-
246 QUEENSLAND SPECIES OF AKCELLA.
ered in October, 1917, the only living Arcellm present were A.
(irtocrea. Furthermore, the two localities are about 130 miles
apart. Again in the occasional material collected outside the
period September-October \see (2), (3), (4), (5j, (8), the dominant
form is A. vulgaris, not A. artocrea.
It is hoped by the collection of more data that deductions may
be made in connection with A. discoides and A. mitrata, as well
as in supplying confirmatory evidence in regard to A. vulgaris
and A. artocrea.
BIBLIOGRAPHY.
1. KoFoiD, C. A.— "The Plankton of the lUinois River, 1894-1899/' Bull.
Illinois State Lab. Nat. Hist., viii., Art. 1, 1908.
2. Leidy, J. — "The Freshwater Rhizopoda of N. America." 1879.
3. Wailes, G. H. — "Freshwater Rhizopoda and Heliozoa f rom the States
of New York, New Jersej', and Georgia, U.S.A." Journ. Linn.
Soc. Lond., Zool. xxxii., 1912, pp. 121-161.
247
ON THE LEAF-ANATOMY OF SC.EVOLA CRASSIFOLIA,
AVITH SPECIAL REFERENCE TO THE EPIDERMAL
8ECRETI0N.
By Mahjokie Isabel Collins, B.Sc, (•Syd.), Demonstkatok in
Botany, the Uniyeksity of Adelaide.
(Plates xxvii.-xxviii., and six Text-figures.)
Introduction.
Sccevola crassifolia forms a prominent element in the sand-dune
Hora near Adelaide, where it occurs in association with Sjnnifex
hirsutus, with such shrubs as Olearia axillaris, Leucopogon
Richei, etc., and with smaller plants such as Pelargonium aua-
trale, Lotus australis, (Eiiothera 6^e7^n^s (introduced), and Senecio
lautus (3, p. 584).
In habit, .S'. crassifolia is a low, spreading shrub, which is
able, by lateral growth, to occupy large areas of the dune. Its
older woody stems form a dense undergrowth, in which a quantity
of blown sand and dried leaves of Posidonia are accumulated,
thus helping to build up the dunes. 6'. crassi/olia is able to
survive burial in sand by an upward elongation of the stem, and
by the development of adventitious roots.
Owing to the extreme stickiness of the buds and young leaves,
the varnishing or lacquering of the surface of older leaves, the
dullness of surface, and the succulence of mature leaves, it was
thought that an examination of the leaf-anatomy would prove
interesting. The investigation was carried out in the Botanical
Laboratory of the University of Adelaide, at the suggestion of
Professor Osborn, to whom I am much indebted for his constant
help and encouragement.
Summary of Previous Investigations.
Despite its particular interest, the N.O. Goodeniacea^ has
received but scant attention at the hands of botanists. As early
248 LEAF-ANATOMY OF SdiVOLA CRASSIFOF.t A.
as 1876, Vesque{5) published an account of the anatomy of
Goodenia ovata, which, however, was restricted to an investiga-
tion of the anomalous stem-structure. With this, he compared
the stems of Scievola crassi folio,, S. Pfuniieri, S. microca7'pa, and
S. spinescens, but there is no investigation of leaf-anatomy in
any of these species.
In the Systematic Anatomy of the Dicotyledons (i., p. 471 ),
Solereder gives a Ijrief account of the leaf-structure of the
Goodeniace*, based upon his own observations of Goodenia
ovata, Selliera radicans^ Screvola Plumieri, and Damjnera
Brownii, and upon those of Briquet in the case of Bninoiiia.
He records that the stomata may occui* upon both surfaces, or
upon the lower surface only, and that the number of subsidiary
cells, when these are present, is four, two of which are placed
parallel to the pore, the other two being applied to the narrow
ends of the guard -cells. In form, the leaf is found to be typically
bifacial, as exemplified by Daniprera Brotvnii^ but is sometimes
centric, as in SccEvola Vluinieri. 8ilicified groups of cells are
common, in the formation of which certain epidermal and the
adjacent mesophyll-cells take part. As well as the ordinary
clothing-hairs, which are characteristic of most members of the
Order, peltate, glandular hairs have been (observed in Goodenia
ovata and Sccevola Fhtmierl. These are described as being
" shortly stalked, peltate glands, the shield of which consists of
a few cells separated by radial walls."
The above account is a summary of our present knowledge of
leaf-anatomy in the Goodeniacea'. It is interesting to note that,
in none of the plants hitherto investigated, has any copious,
epidermal secretion been described. In *S'. crassij'olia, however,
the buds and young leaves are coated on both surfaces with a
thick layer of sticky resin, which is secreted by numerous,
stalked, glandular hairs.
The secretion of mucilage or resin, or both together, has long
been recognised as a method of bud-protection. Groom (2) has
given an interesting account of the mucilage- and resin-secreting
organs of buds. 'iliis work is restricted to those plants whicli
bear "colleters" or "villi" on the stipules. These " colleters,"'
BY MARJORIE ISABEL COLF.TNS. 249
wliich are almost invariably pear-shaped, raiilticelliilar bodies, in
Nvliich a peripheral palisade-layer secretes the miicila_<ie or i-esin,
are only functional in the bud, and fall off with the stipules.
Only in one case— that of Woi-mia Bvrhidyei (Dilleniacese)— does
Groom record an instance of the secretion being derived from
glandular hairs on the leaves themselves, but, of this, no descrip-
tion is given. Volkens(6), however, has described many plants
in which the secretion is not only derived from glandular haiis
on the leaves themselves, but is also produced for a long period
of time, sometimes till maturity of the leaf. It is to Volkens
that we owe what knowledge we have of "leaf-lacquering ' as a
xerophytic adaptation.
In the present investigation, the development of the leaf is
traced from the earliest stages to the mature condition, with
special reference to the glandular covering in the juvenile state,
the function, and fate of its secretion.
MOKPHOLOm OF THK ShOOT.
A shoot of Srcevola crassifolia is characterised by the vertical
arrangement of the leaves, which is consistent from the leaves in
the bud to those farthest away from the growing apex. I he
leaves are all ovate, moi-e or less toothed, petiolate, the base
being closely applied to the stem on its upper side. The leaf-
teeth are more marked in the bud and young lea\'es than in the
mature leaves, in whicli they are often obliterated by secondary
increase in thickness. The terminal bud is small, and, durint/
the growing season (August toC)ctober, the Spring months about
Adelaide) it is hidden by the rapidly expanding young leaves.
Later, when this activity ceases, the bud is hidden at the base
of the most distal leaf. The leaves in the bud are not folded
about one another; they are opposite and vertical, the morpho-
logical upper surface of one leaf being closely glued to the upper
surface of the next youngest leaf.
The whole growing region of the shoot in S. crassifolia is
covered with a sticky secretion, which gradually dries up farther
away from the stem-apex. As the leaves become expanded from
the bud, they increase rapidly in surface-area, and later lose
250 LEAF-ANATOMY OF SC.EVOLA CRASSIFOLIA,
their extreme stickiness. 'J'hey are, at this stage, cliaracterised
by glistening patches of drying secretion, which give them their
varnished or lacquered appearance.
The mature leaves are large, thick, and succulent, and present
a dull surface. Despite the fact that the secretion tends to dry
up on the expanded portion of the older leaves, the leaf-base is
always distinctly sticky, even in the mature leaves. In this
manner, the axillary buds are covered by secretion.
The voung flower-buds are borne in dense, terminal spikes.
As is usual in the Goodeniacese, the cah'x is inconspicuous, and
the corolla protects the essential parts of the flower. In S.
crassifolia, each flower is situated between two bracteoles, and
is subtended by a bract, which is glued to the exposed surface of
the bud by means of its sticky secretion. The glandular hairs,
from which the secretion is derived, are of the same nature as
those on the leaves.
DiSTKI BUTTON OF THE ACTIVELY SECRETING GlANDS.
It is in the buds that the glandular hairs reach their maximum
development. Upon examination of a section through the apex
of a shoot, it will be seen that the rudimentary leaves, even
before they are completely differentiated from the stem-apex, are
provided with numerous, closely-set, mature, actively secreting,
glandular hairs. These hairs are more abundantly developed on
those surfaces which are exposed earliest. In the bud, there is
a continuous production of glandular hairs, the various stages in
development being found mingled with the mature glands. These
developmental stages are not restricted to the youngest portions
of the bud, nor to any particular part of the leaf, but, as will be
seen later, apparently any epidermal cell up to a certain stage in
development, may form a gland (PI. xxvii., fig.l). In this
manner, new glands are provided to keep pace with the increase
in surface of the leaf, the secretion of which will prevent the
sticky coat from being broken at any point.
Once the young leaves are freed from the bud, glandular hairs
are no longer formed; upon such leaves only, the mature con-
dition is found. The distance between these elands tends to
BY MAR.TORIE ISABKL COLLINS.
251
increase as the leaf-surface increases in area by the differentia-
tion of the leaf-tissues. Tn the older leaves, the glands lose
their activity, except as mentioned above, in that region where
the leaf-base clasps the stem. There, the glands remain active
throughout the life of the leaf. A section through this region
shows an abundant development of glandular hairs of the same
nature as those on the young leaves. By the activity of these
glands, the axils of the leaves are filled with secretion, and pro-
vision is made for the protection of the axillary buds. At the
junction of the stem and leaf-base is a circlet of long, multi-
cellular clothing-hairs (PI. xxvii., fig 2;. These were tested for
resin and mucilage, with negative result.
The Development of the Glands.
That the glands are hairs, is evinced in the manner of their
origin from a single epidermal cell. Any epidermal cell may
form a gland. The- cell enlarges, the nucleus divides, and a
horizontal wall is formed separating two cells, the uppermost of
which becomes considerably enlarged and spherical in outline
(Text-fig. 1, a). A second, horizontal wall cuts off the rudiment
Text-fig. 1. — Stages (a-g) in the development of the glandular hairs; ( x 530).
of the .stalk at the base of the spherical cell, which latter is
destined to become the head-cell of the gland (Text-fig. 1,6). At
this stage, either a second stalk-cell is formed (Text-fig. l,c>, fol-
252
LEAF-ANATOMY OF SC.F.VOLA CHASSIFOLIA,
lowed by a median, vertical division in the head-cell (Text-fig. !/•),
or the latter division takes place before the formation of the
second stalk-cell (Text-fig. \,d). A third stalk-cell is now formed,
either preceded or followed by the ultimate divisions in the head,
ceils ( Text-fig. 1,/, y).
a b
Text-Hg.*2 : II, surt'aee-view of a mature ^laiid; l>, pait of section through
yount>; leaf, to show crowdinu' of niatuie glands, and oveilappinsj; of
shields: ( x -)8()).
The head of the matuie gland forms a shield of eight cells,
only four of which are .seen in transverse section (Text-fig. 2, n).
The shield, which is ultimately reflexed at the sides, is borne
upon a stalk of three tiers of cells, the two lower tiers being
composed each of a single cell, while the upper is composed of
two cells, which grow out laterallv to support the head-cells
(Text-fig.2,6).
The Nature of the Skcretion.
The secretion, which covers both upper and lower surfaces of
the leaf, is a yellowish, translucent, homogeneous substance,
which is soluble in alcohol. It is very sticky when fresh, and
gives the characteristic red colouration of resin when tested with
Alkanna root-extract. A test for mucilage, carried out with
corallin-soda, gives a negative result. The secretion reaches its
maximum thickness in the bud, every available space between
the young leaves being filled up. This fact, which is probably a
result of the continuous production of glands in the bud, indicates
BY MAR.TORIK ISABEL COLLINS. '2i).^
that tlie secretion of resin in S. crassifoHah primarily concerned
with bud-protection.
Tlie secretion is still mobile upon the X'oung leaves outside the
bud, but since no new glands are being formed to keep pace with
the increase of leaf-surface, it becomes spread out over a greater
area and, consequently, more attenuated. The thickness of the
film of secretion varies in different parts of the same leaf, in
leaves of different age, and in diffeient plants. It often reaches
a thickness nearly twice the height of the glands, i.e., ab.mt
7G/X. (PI. xxviii., fig. 3).
In the older leaves, the glands lose their activity, the secretion
becomes less mobile, and, in drying up, oacks along radiating
lines. Tt is at this stage that
the leaves present a lacquered
appearance. The glands, being-
no longer functional, cease to
grow, whilst the neigh homing
epidermal cells enlarge consider-
ably, and, as a result, the glands
ultimately appear to be sunken
(Text-fig. 3). The dried secre-
tion falls off" when the leaf
0
^/
& 0
0
^0
(5 t)
0
0
0 0
0
0
ol K 1)
0 (^
reaches its succulent condition, Text-fi-.8.-Sunken, olandular hair in
,, . 1 p TIT '^ mature leaf: ( xoRO).
small patches, oi negligible
quantity, being restricted to the region of the glands.
The Development of the Stomata.
Tt is obvious that any stomata which are present on the young
leaf coated w^ith its sticky secretion, cannot be functional. Tn
correlation with the rapid growth of the leaf-area, new stomata
are continually being formed. Thus, in a section parallel to the
surface of a young leaf, all stages in development of the stomata
may be studied, as well as a small percentage of stomata which
have already reached maturity (Text-fig. 4).
Solereder (4, Vol. ii., pp. 1079-1080), wlien classifying stomata,
refers the Goodeniacete to two groups; (1) where there are no
subsidiary cells, and ('2) where subsidiary cells are formed par-
18
254
LEAF-ANATOMV OF SCEVOI.A CRASSIFOLIA,
allel to the pore. In S crassifolia there are no subsidiary cells.
A single epidermal cell divides by an oblique wall into two cells
of unequal size. The larger of these cells remains an epidermal
cell, while the smaller is the mother-cell of the guard-cells. The
only further division which takes place is that which forms the
two guard-cells. The mature stoma is surrounded by a variable
number of epidermal cells ( Text. fig. 4 o?). The stomata reach
Text-fig. 4. — Stages {a-c) in the development of the stomata, as seen in
transverse section of young leaf : a, initial stage, showing mother-
cell of the guard-cells; Ji-r, showing development of two guard-cells;
no pore is formed; d, sui'f ace-view of young leaf, showing stages
in development of stomata; ( x 300).
their full development when the flow of secretion has ceased.
Those which mature first, are lendered functionless, for a time
at least, owing to the blocking of their pores with resin. It
seems highly probable that many of those stomata, in which a
plug of resin is found enclosed in the stomatal aperture, will
never function (PI. xxviii., fig. 4). Only those stomata which
are belated in development, and are freed before reaching matu-
rity, by the drying-up and cracking of the resin, are entirely
unblocked by the secretion.
In those plants with lacquered leaves, which Yolkens(6) de-
BY MAHJORIE ISABEL COLLINS.
2oo
scribed, such as species of BrachylcEua, Baccharis, EscallonKt, etc.,
the stomata are belated in development: and, in a few cases in
which the secretion remains active until maturity of the leaf, the
stomata become elevated until they project above the surface of
the secretion. The mature stoma of .V. crassifolia remains at
the surface-level of the leaf. There is little of interest to remark
in the structure of the mature stoma, a small, outer chamber
being formed in the usual way, by an overhanging lip of cuticle.
Internal Development of the Leaf.
As in the case of the stomata, the development of the leaf-
tissues is belated. In a young leaf just freed from the bud, there
is no internal differentiation
except that of the vascular sys-
tem. The leaf, at this stage, is
made up of a compact mass of
rounded cells, the vascular
strands occupying a median
position. It is only when the
secretion becomes less active,
that differentiation of the leaf-
tissues takes place.
The fully-grown leaf of .S'.
crafisi/olia is very thick and
succulent, owing to the devel-
opment of secondary, water-
storage parenchyma. The epi-
dermal cell-walls show a great
development of cellulose, but
are only moderately cuticular-
ised. There is no si^ongy meso"
phyll, the greater part of the
leaf being made up of several
rows of palisade-cells. A tis-
sue of rounded cells, amongst
which are situated the vascular
strands, occupies a median posi-
tion in the leaf, and is bounded on either side by, sometimes, as
Text -tig. o. — Section of mature leaf
showing occurrence of branched
nnicilage-cells in the palisade-
tissue; ( X 300).
2o6
LEAF-AKATOMY OF SCiEVOLA CKASSIFOLIA.
many as five rows of palisade-celis. This central tissue is re-
markable in the ver\' old leaves, as its cells become enlarged,
lose the greater part or all of their chlorophyll, and take on the
function of water-storage. Those cells immediate) v surrounding
the vascular strands are the first to assume the new function
(PI. XX
tig. 5). From this central, water-storage tissue and
from the palisade tissue, special water-storage organs, such as
mucilage cells and water-storage tracheides are differentiated.
Miic'daye cells. — lu the early stages of differentiation, certain
cells, particularly in the palisade-region, develop mucilaginous
walls. These cells keep pace with the growth in thickness of
the leaf by branching in all directions, producing, thus, a peculiar
and striking result (Text-fig.5). Text-fig. Gti shows one of tliese
branched mucilage-cells swollen after treatment with potash, and
stained with corallin soda to show successive deposits uf mucilage
in the cell-wall.
a. b
'IV-xt-H^^.d : rt, single luucila^e-cell cilter tit'atiuent with potash, and stained
with curallin-soda, to show strati tication of luuoilage; ( x 230) : /', a
group of water- storajfe tracheides from the central tissue of the
mature leaf; ( x 400).
Water star (iqe tracheides. — llestricted to the central tissue are
special water-storage tracheides occurring both at the bundle ends,
and in groups, quite independent of the bundles. They are of
BY MAR.TORIK ISABEI. COLLINS. 257
the type uitli thickened walls bearing transversely elongated
pits (Text-fig. 6,6'.
At the midrib, the tissues of the leaf merge into a compact
chlorenchymaof rounded cells. Tlie vascular strands are strengtli-
ened on both the upper and lower surface Ijy a bundle of fibres.
Crystals of calcium oxalate are found in the cells of the leaf in
very small quantity.
Conclusion.
The secietion of resin by glandular hairs in the bud and young
leaves, and the lacquering of older leaves by the ultimate drying-
up of this secretion, have been shown by Volkens(6) to be peculiar
to some xerophytic plants. Sccevola crdsslfoiin, being a sand-
dune [)lant, may be regarded as a xerophyte.
The chief ecological factors with which a dune-plant has to
contend, have been summarised byCowles(2, pp. 107-111). They
are, (1), intense illumination, both direct and leHected; (2), the
great divergence in the temperature-extremes, which is still
further increased by the low specific heat of sand; (3), the drying
action of the wind, and the injurious effects of wind-blown sand;
(4), the porous nature of the sand, and its consequent low water-
capacity. The third and fourth of these factors are, perhaps,
the most potent. Tn South Australia, however, the first factor
may have some considerable importance, since there are often
periods of several days with cloudless sky, while the glare from
the sun is still further intensified by the whiteness of the coastal
sands.
As we have seen, S. crassij'olia is able to survive burial in sand
by an upward elongation of the shoot, and by the development
of adventitious roots. By virtue of tlieir thick, sticky secretion,
the buds and young leaves are protected from desiccation, as are
also the older leaves, owing to their succulence. A further pro-
tection from intense illumination is suggested by the yellow
colouration of the secretion. The polished surface of older leaves,
caused by the partially dried secretion, also serves to refiect
certain rays of light.
The low water-capacity of the sand is compensated, to some
extent, by the water-storage tissues of the older leaves.
258 leaf-anatomy op sc^vola crassifolia,
Summary.
i. Sccevola crassifolia is characterised by the development of
peltate, glandular hairs, which secrete resin in great quantity.
The-^e hairs reach tlieir maximum activity in the buds and young
leaves.
ii. As the leaves become older, the glands become less active,
the secretion loses its mobility, dries-up, and gives the surface
of the leaf a lacquered appearance.
iii. The mature leaves of .S'. crassifolia are thick and succulent.
The glands are sunken, and no longer functional on the blade of
the leaf. They retain their activity only in the region of the
leaf-base, protecting the axillary buds. Si^ecial xerophilous
adaptations are found in :
[a). The secondary increase in size of the epidermal cells.
(6). The massive development of palisade-tissue.
(c). The development of special water-storage cells, such as
mucilage-cells, water-storage tracheides, and ordinary, thin-
walled, water-storage cells.
EXPLANATION OF PLATES XX VII. -XXVIII.
All tigiues were made at table-level, with the aid of Zeiss* camera lucida,
with mirror at its j^reatest inclination, and with tube at 160 mm. Leitz
objectives 3 and (J, and oculars 2 and 4 were used.
Plate xxvii.
Firf. 1. — A slit^htly oblique section throu^fli a bud, to show development of
•glandular hairs : a, stem-apex; h, rudimentary- leaf; c, leaf-base; r/,
petiole of slighth' older leaf; c, glandular hair; ( > 160).
Fig. 'J. —Section across sheathing base of a mature leaf, to show develo})-
ment of glandular and clothing hairs; ( x 1(50).
Plate xxviii.
Fig.o. — -Part of .section through young leaf, to sliow depth of secretion and
outline of glandular hairs; ( x o30).
Fig.4, — Section of a lacquered leaf, to show patch of secretion blocking
mature stoma; ( x 300).
Fig. 5. — Section of a mature leaf : p, palisade-tissue: c, central water-
storage tissue; m, mucilage-cell; f, water-storage tracheides; ( x 100).
BY MAK.TORIK ISABEL COLLINS. 209
LITERATURE CITED.
1. CdWLES, H. C. — "The Ecoloifieal Relations of the Vegetation on the
Sand Dunes <»f I^ake Michigan. "' iiotanical (ia/ctte. xxvii.. Nos. "2,
3, 4. 5, 1899.
2. (Jroo.m. p. — "On Hiul-]^'otection in the Dicotyledons. '" Trans. Liim.
Soc. (2) iii.. 1S9-2.
3. OsBORN. T. (r. B. — " Types of Vegetation on the Coast in the Neicfh-
hourhood of Adelaide. S.A." Report Brit. Assoc. Adv. Sci., 1014.
i. SoLEREOEK. — Sj'stematic Anatomy of the l)ioot3'ledons. Vols, i.-ii.,
1908.
5. Vesqfe. — "The Anatomy of (i'oofh)ii(i omla." Ann. Sci. Xat., Hot.,
Ser. 6. iii., 1870.
6. VoLKENS, Ct. -" Ueber Pflanzen mit lackirten IJliittern." Her. der.
Deutsch. Hot. Resell., 1890.
260
A F0S8IL INSECT- AVTNG FROM THE ROOF OF THE
COAE 8EAM TX THE SYDNEY HARBOUR
COLLIERY.
Ry R. J. TiLLYAHD, M.A., D.Sc, F.L.S.. F.E.S, Linnean
Maclkay Fellow of the Society in Zoology
(One toxt-fiiiiire.)
In tlie Journal and Proceed ins^s of tlie Royal Society of New
South Wales, Vol. xlv., 1911, p.r)5I, Mr. AY. S. Dun records the
occurrence of a species of Tceniopteris in the roof of the Coal-
seam in the Sydney Harbour Colliery at Balmain (Copper Per-
mian). An excellent figure of the actual specimen, reproduced
from a photograph, is given in Plate xli., in the same paper. The
writer concludes with the following note : — "Associated with
the leaf is the impression of an insect's wing, a description of
which will be submitted later."
Recently Mr. Dun has very kindly passed this specimen over
to me for description, and I desire to thank him for the oppor-
tunity of studying it.
The wing lies upon the smooth surface of a very dark, hard
shale. Its position is best located by the following note.
Measure 18 mm. from the distal preserved end along the mid-rib
of the frond of Tcfiiiopteris, and, from the point .so obtained, erect
a perpendiculai- on the side on which the centre of curvature of
the frond lies. The distal end of the insect-wing lies at a dis-
tance of 20 mm. along this perpendicular.
The wing can be actually seen in the reproduction mentioned
above, but it is so faint in outline that it would not be noticed
unless some measurements were given to assist one in locating
it. The same may be said to be the case with the actual speci-
men. Once it has been located, a careful examination with a
hand-lens will show the presence of the main veins, and, in
certain lights, of cross-veins also. All of these, except only the
BY R. J. TILLYARD. 261
radius, are exceedingly delicate in outline, so that it is by no
means easy to make an accurate drawing of the fossil.
The portion preserved consists of a considerable part (probably
about three-fifths) of a slender wing, with the costal margin well
marked right up to the apex. The posterior border is missing,
except for a short piece lying towards the base of the fossil.
The space between the radius and the costal margin is distinctly
coloured brownish. This is almost certainly due to the preser-
vation of brown pigment originally located in this area of the
wing.
In attempting to place this fossil correctly, I have had to rely
upon only two or three important characters, since the base of
the wing, which may be presumed to have held the most definite
venational structures, is missing. First of all, I have relied
upon the absence of the subcostal vein from all the distal portion
of the costal space. Secondly, the well marked radius, evidently
very strongly built, has running below and parallel to it a well-
developed radial sector, giving off obliquely descending branches
at fairly wide intervals Combining these characters with a
third, viz., that the cross-veins are fairly wide apart, regular,
and somewhat oblique, we should have little difficulty in making
a very close comparison between this fossil and the interesting
family Elcanidce, of the Order Orthoptera, from the Mesozoic
strata of England and Germany.
Handlirsch defines the ElcanidcB as follows^ (his text is in
German) : — This family is characterised by the possession
of long, typical Locustoid antennsCj well-developed jumping legs,
and also, in the female, a long ovipositor. On the other hand,
up to the present, no wing has been found with a stridulating
organ.
The wing-venation resembles in many respects that of the
Acridioidea more than that of the Locustoidea existing to-day.
The forewing is characterised by a costal vein slightly removed
from the border, and, therefore, by a precostal area. The sub-
costa is greatly shortened; the radius is free, and sends branches
towards the free anterior border; its sector arises near the base,
* Die Fossilen Insekten, p. 412.
19
262 FOSSIL INSKCT-WING FROM SYDXKY HARBOUR COLLIERY,
and runs nearly parallel with the main stem towards the tip: it
sends a variable large number of branches sharply from below
towards the apical Imrder. Media and cubitus strongly reduced,
consisting of small branches. Anal field somewhat small, with
few veins.
The hind wing is similar to the fore in respect of the structure
of the radius, media, and cubitus, but appears to lia\e undergone
a vast reduction of the subcosta, and to possess no precostal area.
Its anal field is large and fan-like.
Stiff cross-veins divide the wing-areas into quadrilateral cells.
Most of the wings are spotted.
It will be seen that, owing to the fragmentary nature of the
wing, many of the above characters are not available as tests.
On tlie other hand, the part preserved agrees very closely with
the definition. The small portion of the posterior border pre-
served is of some value to us, since it points to the fossil repre-
senting a forewing. For, in the hindwing, we should have ex-
pected to see, in this position, part of the anal fan which is so
conspicuous in this family, but which is evidently absent here.
Handlirsch lists forty species of the family Elcanidce from the
Lias, and twelve from the Upper Jurassic. All but one of these
are placed in the genus EJcana Giebel, the only other genus
being Parelcana Handl I propose to place the fossil here de-
scribed in a new genus, Elcaiiopsis, with the characters given
below.
Order ORTHOPTERA.
Family ELCANID.^.
Genus Elcanopsis, n.g. (Text-fig. 1).
Characters of forewing as in Elcana Giebel, except the fol-
lowing : — Sc towards its distal end lies very close to C, not
nearer to R as in Elcana. Wing not spotted, but marked with
brown pigment along the costal space. Branches of Rs few in
number, and placed fairly wide apart. Cross-veins not very
numerous or close together.
Genotype, Elcanopsis sydneiensis, n.sp. (Upper Permian :
Balmain Colliery, Sydney Harbour).
The occuri^ence of an Elcanid genus in the Upper Permian is
BY R. J TILLYARD.
263
of considerable interest, since the insect-fauna already known
from the Upper Permian of Newcastle, New South Wales, con-
tains only types which would be regarded as of Mesozoic age if
they had been found in the Northern Hemisphere. It is also
interesting to note that the Upper Jurassic members of the
family approach, in their venational type, the Acridiiihe of the
present day, whereas the Liassic species differ more widely from
these last, in that the number of branches of the radial sector,
and the number of cross veins, is considerably less, and the species
are of a smaller size In these characters, the new genus is the
most reduced of all; so that the supposition now naturally arises
that the modern Acridioid type of venation may have been
formed by addition of new elements to what was originally a
much >-impler and moi'e open type of venation.
Text-fig. 1.
Elcanopsis s?/fZ??e /ensjX n.g. et sp., portion of forewing; ( x 18*5). Upper
Permian Coal-Measures : Balmain Colliery, S3'dne5' Harbour. C,
costal border; R, radius; Rs, radial sector; Sc, subcosta; the area
covered bj- the brown pigment is shaded.
Elcanopsis sydneiensis, n.sp. (Text-fig. 1).
Total length of fragment, 7 mm. Greatest breadth, 2*7 mm.
Foreiving with the distal end of Sc preserved, and apparently
ending up on ^) at about the middle of the costal border. R
very strongly built, very gently curved, the tip slightly turned
upwards. Apparently six very poorly preserved cross-veins
between R and C, beyond the end of Sc, and two oblique ones
between Sc and R basad from these. Nine or ten cross- veins
preserved between Pt and Rs; some of these fairly distinct. Rs
264 FOSSIL INSECT-WING FROM SYDNEY HARBOUR COLLIERY.
a slender but well-preserved vein, parallel to R, and having four
descending branches placed somewhat wide apart. The cross-
veins between these branches mostly not visible, but indications
of them here and there in a good light. A portion of a fairly
strong vein is preserved near the posterior border, but this vein
cannot be named with certainty. It appears to be connected
with the border by means of a series of oblique cross-veins.
Type, in Coll. Geological Survey of New South Wales,
Sydney.
Postsa'i}>t (added June 14th, 1918).^In my paper on the
" Permian and Triassic Insects from New South Wales, in the
Collection of Mr. John Mitchell," These Proceedings, 1917, xlii.,
Pt. iv., p. 7 25, I named Schizoneiira as one of the genera of the
fossil flora of the Upper Coal-Measures of the Newcastle District
The name was taken from the list given in Siissmilch's "Geology
of New South Wales," on p. 135, where also the genus Tceni-
opteris is included. As several geologists have taken exception
to the inclusion of this genus in the Permian Flora, I take this
opportunity of pointing out that I did not state that the genus
Schizoneura occurred at the horizon of either the Belmont or
Newcastle Insect-Beds. I have to thank Mr. W. S. Dun for
explaining the exact state of the case to me. It appears that
Schizoneura is found only at the very top of the Permian Beds,
in beds that might w^ell be considered as the passage-beds between
the Permian and Triassic, but which should be classed as Per-
mian as long as we define the presence of Glossopteris as the
criterion by which to separate the Pala3ozoic from the Mesozoic
in Australia. The same is true of 7'ce7iiopteris, which is asso-
ciated with the insect-wing described in this paper; it should be
noted especially that the species, though unnamed, is certainly
not T. Baintreei McCoy, ( = T. spatulata McClelland), which is
characteristic of the Jurassic. The horizon of the wing described
in this paper is, therefore, some 600 feet vertically above that
of the Belmont Insects previously described, and more than 800
feet above that of the Newcastle Insects. Whether, therefore,
Elcanopsis be considered to be a Permian genus or not, there
can be no question that these other insects are such.— R.J.T.
265
THE PANORPOID COMPLEX.
A Study of the Phylogeny of the Holometabolous Insects,
WITH Special Reference to the Subclasses Panorpoidea
AND NeUROPTEROIDEA. [INTRODUCTION].
By R. J. TiLLYARD, M.A., D.Sc, F.L.S.. F.E.S., Linnean
Macleay Fellow of the Society in Zoology.
Introduction.
Section i. — Definition of the Complex.
In his great work upon the Phylogeny of the Orders of Insects.
Handlirsch (2) divides the whole of the Holometabola into four
Sub-classes, as follows —
Sub-class.
Orders.
Hymenopteroidea
coleopteroidea
Neukopteroidea
Panorpoidea
Hymenoptera.
Coleoptera.
Megaloptera, Raphidioidea, Neuroptera.
PanorpatxV, Phryganoidea, Lepidoptera, Diptera
Aphaniptera.
In making this division, Handlirsch widely separates the first
two of these Sub-classes from the last two, placing between them
not only what we may, for convenience, term the three Amplii-
biotic Orders (Odonata, Plectoptera, and Perlaria), each of
which he elevates to the rank of a Sub-class, but also the Embi-
daria, which he also considers to be a Sub-class. For the
Hymenopteroidea and Coleopteroidea, Handlirsch indicates a
descent from a JJlattoid type of ancestor. For the JMeuropter-
oidea and Panorpoidea, he is more inclined to a Palseodictyopt-
erous type of ancestor; indeed, he definitely states his belief that
the Neuropteroidea are descended directly from the Palaiodicty-
optera, while he considers the Panorpatre, and hence the Panor-
266 THK PANORPOiD coMPlp:x (Introduction),
poidea also, to have been derived from the obscure Megasecoptera
of the Upper Carboniferous- an Order which, at the best, can
be regarded only as a specialised side-branch of the Pal?eo-
dictyoptera.
AVe are thus faced with the fact that, in Handlirsch's opinion,
the Holometabola are a composite group of polyphyletic origin.
Such a view is, indeed, fairly generally held amongst entomolo-
gists; for no satisfactor}^ reason seems to have been brought
forward why the evolution of the resting-pupa should not have
been accomplished more than once, along several quite distinct
lines of ascent.
Having for a very long time carefully read and studied Hand-
lirsch's views, I have to confess that, in spite of many excellent
arguments brought forward, there seems to me to be much that
is unsatisfactory in them. As examples, I might refer to his
discussion of the origin of the Hymenoptera, which fails to con-
vince me in a single point. The same may be said of the sug-
gestion that the Panorpatse are derived from the Megasecoptera.
It seems to me that any unbiassed student of the Phylogeny of
the Holometabola must come to the following conclusions : —
(1) That the origin of the Hymenoptera is still about as great
a m\^fetery as it well can be; and that Handlirsch's solution of
this problem has many inherent defects that make it unaccept-
able.
(2) That the origin of the Coleoptera from a Blattoid type of
ancestor is a fair prolmbilit}-; but that much more research of
an intensive character must be carried out before this theory can
be either proved or disproved.
(3) That the origin of the Neuropteroidea from Palseodicty-
opterous ancestors is extremely probable: nevertheless here also
any researches that tend to strengthen the argument would be
very welcome.
(4) That the origin of the Pauorpoidea from the Megasecoptera
is not supported by a single piece of evidence worth considering;
but that the possibility of the relationship between Keuropter-
oidea and Panorpoidea being exceedingly close requires recon-
sideration, and the coincident possibility of the Panorpoidea
BY R. J. TlLLYARt). 267
having had Palifiodictyopterous ancestors also requires careful
investigation.
My own researches upon the JSfeuropteroidea and Fanorpoidea,
as far as they have gone, have convinced me that these two Sub-
classes have a great deal in common. They have also revealed
the possibility of the Hymenoptera, and even the Coleoptera,
having a closer relationship with the Fanorpoidea than is gener-
ally suspected. Not only do many signs point to the Mecoptera
(Fanorpatae) as being a central Order round which all the rest of
the Holometabola may be more or less closely grouped, but the
Palseontological evidence also points unmistakably in the same
direction. For fossil Mecoptera of the genus Pennochorista,
closely allied to the existing Australian genus TfEiiiochorista^
have now been proved to exist in the Fermian of Newcastle, New
South Wales(6),; whereas no other Holometabolous insects are
known from Falseozoic strata at all. Even admitting the incom-
pleteness of the fossil record, we must be immediately struck
with the fact that the Mecoptera existed in Fermian times in
Australia, in a form very similar to that of to-day. This points
to the Order having arisen well before Fermian times. The
earliest known Neuropteroidea are Upper Triassic, the earliest
Coleoptera also Upper Triassic, the earliest Trichoptera the
same, and the earliest Hymenoptera Upper Jurassic. Even if
we grant that it is a reasonable expectation that all these Orders
will one day be found to have had representatives in earlier
strata, yet the same probability holds for the Mecoptera.
Reviewing the whole case, it seems to me that the time is ripe
for a careful study of the whole problem, as far as the evidence
will admit, from the point of view of the Mecoptera as the
central Order; that is to say, the Order which has preserved,
both in its larval, pupal, and imaginal structures, the largest
number of archaic characters derived from the original ancestor
or ancestors of the Holometabola, whatever they may have been.
It is to suggest this point of view that 1 have selected as title
the somewhat elastic term " The Fanorpoid Complex." By this
title, I intend to convey that the research entered upon in this
paper has, for its main object, the complete working out of the
268 THE PANORPOID COMPLEX (Introduction),
relationships of the Order Mecoptera to those other Orders which
stand in closest relationship to it. In accepting Handlirsch's
division of the Holometabola into four Sub-classes, it is to be
understood that I do so only provisionally, for the convenience
of sifting and allocating the evidence that this arrangement
naturally offers. And, as it is obvious that the main weight of
the argument must centre around those Orders which are clearly
most closely associated with the Mecoptera, so it will be clear
that the Orders comprising the Panorpoidea must be most fully
reviewed. There is, however, a great deal of evidence to be ob-
tained from a study of the Neuropteroidea; and this Sub-class
can by no means be denied full consideration in the argument.
With respect to the Coleoptera and the Hymenoptera, the evi-
dence to be obtained from them at the present time is compara-
tively scanty, and has little weight compared with that derived
from the other Orders, though it may help to throw some light
upon the larger problem of the origin of the Holometabola as a
whole. These ideas I have tried to convey in the sub-title
selected for this paper.
If, then, we accept provisionally, for clarity of argument, the
two Sub classes Neuropteroidea and Panorpoidea, in the sense
that Handlirsch defined them (but with some modiHcations in
the nomenclature of the Orders composing them), we may then
proceed to define the " Panorpoid Complex" as that assemblage
of Orders ichose ancestral characters can be shown to possess close
ajfiniti/ with the characters preserved to a yreat measure itt the
Order Mecojdera, without in any way binding ourselves to the
inclusion of any particular Order in the Complex. The Complex
itself is an elastic assemblage of Orders; and any particular
Order may be included in it, or removed from it, according as
the weight of evidence may determine.
Throughout this paper, I shall use the name Mecoptera for
the Panorpatte of Handlirsch, and Trichoptera for his Phrygan-
oidea. The Rapliidioidea I hold to be not sufficiently distinct
from the Megaloptera to deserve ordinal rank, since van der
Weele's work (7) indicates the strong probability of their origin
from the more ancient Sialoid stem. Hence I shall merge them
BY R. J. TILLYARD. 269
into the Order Megaloptera, which will then consist of two Sub-
orders, viz , the Sialoidea (aquatic) and the Raphidioidea (terres-
trial). The term Neuroptera is now so ambiguous, being still
used in present-day writings to indicate so many different and
heterogeneous groups of Orders, that I have no hesitation, for
the sake of clearness of argument, in substituting the name
Planipennia for it, although I do definitely subscribe to the
opinion that the name Neuroptera ought by now to be strictly
confined to this Order only.
For tlie purposes of this paper, I shall definitely include my
new fossil Order, Protomecoptera, within the Sub-class Panor-
poidea, to which it clearly belongs. The type (and so far the only
representative) of this Order is Archipanorpa ma<j7iijica Tillyard,
from the Upper Trias of Ipswich, Queensland. Whether this
type should constitute a new Order, or only a Sub-order within
the Order Mecoptera, it will undoubtedly conduce to claritv of
argument if we employ the term Protomecoptera in the sense in
which I originally defined it (5).
The scheme adopted in this paper may now be exhibited as
follows : —
Sub class Panorpoidea :
Orders Protomecoptera (fossil only), Mecoptera, Trichoptera,
Lepidoptera, Diptera, and Aphaniptera.
Sub-class Neuroptekoidea :
Orders Megaloptera (including the aquatic Sub-order Sialoidea,
and the terrestrial Sub-order Ptaphidioidea) and Planipennia
( = Neuroptera, s.str.).
Section ii. — General Principles.
Most of the work that has been done in Entomology upon the
Phylogeny of any given Order has naturally been carried out by
experts upon the Order in question, with litlle reference to out-
side Orders. As an example of this, we may cite Meyrick's
numerous works on the Lepidoptera, which, apart from their
systematic aims, have also attempted to show the origin of the
Order, as a whole, from the Trichoptera. In ihe whole of this
work, Meyrick makes allusion to only one genus [lihyacophila)
270 THE PANORtoiD COMPLEX (Introduction),
of the supposedly ancestral Order, and, indeed, suggests, in more
than one place, that the whole of the Lepidoptera are descended
from this still existing and abundant genus of Caddis-flies.
Throughout the work, the characters of the Lepidoptera are
subjected to a searching scrutiny and criticism, but the same
tests are not applied to the Trichoptera, which are, nevertheless,
an equally important factor in the problem [3, i\.
Tt has, for long, seemed to me that a problem of this kind,
attacked in this manner, i.e., by intensive study of the internal
differences within the Order in question, without an equally
intensive study of the differences existing within the Oiders
supposedly ancestral to it, could not possibly yield a complete
and accurate solution. The alternative, chosen by some few
authors in recent years, has been to pass in review the sup-
posedly archaic characters of related Orders, and to attempt to
derive from this evidence some idea as to the standing of one
Order to another. As an example of this, I need only mention
again Handlirsch's famous attempt to give us a complete Phylo-
geny of the whole of the Orders of Insects, as well as a Phylo-
geny of the Class Insecta itself(2).
It will be at once admitted that this second method is the
right one. It has, however, one obvious disadvantage. In
order to carry it out successfully, the author should be equally
expert upon all the Orders that he reviews. But no man could
hope to attain the knowledge in half-a-dozen or more Orders
that can be attained in one by a life-study of it. Hence we see
that, while the outlook of such an author will be a broader one
than that of the expert in one Order only, yet the evidence
brought forward cannot all be accepted at the same face- value;
and the chances of misinterpretations at vital points is evidently
very much greater.
As an example of this, we might contrast the treatment of
the Phylogeny of the Lepidoptera by Meyrick on the one hand,
and by Handlirsch, on the other. First of all, Meyrick confines
himself almost entirely to wing-venation. By an exceedingly
full analysis of the different types within the Order, he arrives
at the conclusion that the homonomously -winged Jugatse repre-
BY R. J. TILLYAPr). 271
sent the oldest type, within the Order. Having then discovered,
in the living genus RhyacopJula of the Order Trichoptera, a
venation with apparently all the essential characters of the
Jugatse, he claims that the Jugatie, and therefore all the Lepi-
doptera, are to be regarded as having had a Rhyacophilous
ancestor.
If the characters considered by Meyrick were the only ones
that concerned the question, and if the Orders Trichoptera and
Lepidoptera were so isolated from all the other Orders of Insects
that there could be no question of affinities in any other direc-
tion, Meyrick's solution might be accepted as correct, in spite o'
its having been based mainly upon the wing-venation only. But
this is not the case. As we shall see, the Order Mecoptera has
quite equal, if not superior, claims to be regarded as the ancestral
type from which the Lepidoptera sprang, while the claims of the
Planipennia, in certain directions, may by no means be over-
looked. Moreover, no attempt was made to test the claim, which
surely can legitimately )je made, that the Trichoptera themselves
are a by no means unspecialised Order, with almost as much
right to being considered an end-term in a Phylogenetic Series
as the Lepidoptera themselves.
Many of these objections to the method of procedure adopted
by Meyrick are overcome by the method which Handlirsch fol-
lowed. This author, first of all, considers the relationships of
the Trichoptera with the Mecoptera, and concludes that the
former are an offshoot of the latter. He then propounds the
question as to whether the Lepidoptera are to be legitimately
regarded as the derivatives of the Trichoptera themselves, or
whether we are compelled to go further back, to the older
Mecoptera, in order to indicate their ancestors. His review of
this question is, in its way, a masterly exposition of the facts, as
far as he knew them; and his conclusion, that the Lepidoptera
are not descended from the Trichoptera, but directly from tlie
older Mecoptera, cannot fail to commend itself to all biologists,
if the facts that he quotes are really correct.
Here, then, comes in the question which I have already men-
tioned, viz., the amount of equipment of specialised knowledge
272 THE PANORPOID COMPLEX (Introduction),
of all the Orders taken into consideration, necessary for an
author who would decide this problem. Having carefully read
through the whole of Handlirsch's work, I was at once struck
with his evident lack of specialised knowledge of the two Orders
most fundamental to the whole question, viz., the Triclioptera
and the Mecoptera. The characters of these two Orders are
reviewed very superficially in comparison with those of the
Lepidoptera and Diptera: in fact, the whole of their phylogeny
is decided within the limits of a single page of print. And, on
that one page, I find statements made which, as far as my know-
ledge of these Orders goes, are incorrect. Such, for example, is
the statement on p 1254, that the jugum appears in the Meco-
ptera as well as in the Lepidoptera, and that on p. 1253, that the
Mecoptera retain the archaic homonomous wings and archaic
mouth-parts. In my studies of the Mecoptera, I have found
that none of these statements are justified, and I shall have to
deal with them more fully in the part of this paper devoted to
that Order.
We are thus faced with the situation of having to choose
between the limited outlook, but greater accuracy in detail, of
tlie solution offered us by the specialist in one Order, of whom I
take Meyrick as a conspicuous example, since he shows in a most
remarkable degree the combination of both these characteristics;
and the wider outlook, but occasional inaccuracy in detail, of the
solution offered us by the non-specialist, whose aim should be to
regard all the Orders coming under his review as equally entitled
to careful examination and consideration.
It will he at once obvious, without any further argument, that
neither of these alternatives is entirely satisfactory. Before we
can deal satisfactorily with the phylogeny of a group of Orders,
we must attain, as far as possible, to the specialist's knowledge
in every one of them. Such knowledge, in the present advanced
state of Entomology, is unattainable within the lifetime of any
single man. We must, therefore, ask ourselves, firstly, whether
we may legitimately attack a problem of such magnitude,
equipped with anything less than the maximum of attainable
knowledge on all Orders; and, secondly, whethei-, if this first
BY R. J. TILLYARD. 273
question be answered affirmatively, we can indicate with any
certainty which portions of that knowledge are essential, and
which are not.
Now there will be, in any group of Orders, some which show
definite specialisation above the others: as, for instance, the
Lepidoptera and Diptera above the Trichoptera and Mecoptera.
All entomologists are agreed upon this. Is it more necessary to
attain to the specialised knowledge of the lower groups, or of the
higher groups ? It will be seen that Meyrick was a specialist in
the higher of the two Orders which he discussed, while the
same is true of Handlirsch, in a more general way, since he,
everywhere, shows a far more intimate knowledge of the Lepi-
doptera and Diptera, but especially of the latter, than he does
of the Trichoptera and Mecoptera. Would it not be preferable
for the author, who is to attempt a satisfactory solution of this
problem, to throw the main weight of his studies on to the side
of the more archaic Orders, which, unfortunately for the progress
of Entomology, have been so neglected for many years, and to
be content to attain to a first-hand knowledge of only the more
archaic types within the more highly specialised Orders?
I believe that this question must be answered in the affirma-
tive, for some very good reasons. Firstly, the knowledge of the
more archaic Orders is the only sure foundation upon which the
Phylogeny of the higher Orders can be built. Any attempt
without this knowledge is merely building up a house without
la3'ing secure foundations. Secondly, the connections sought for
in the tracking out of the ancestry- of the higher Orders must be
those between some members (either existing or extinct) of the
more archaic Orders and the least specialised families of the
higher Orders. Thirdly, if at any time the problem transcends
the attainable bounds of knowledge of the investigator who has
adopted the course here advocated, he can always call in the
advice of "the specialist, who possesses just that very type of
knowledge, in his own group, which would render the Phylo-
genist's task too overwhelming for him, on the principle of "not
being able to see the wood for the trees.'' And, finally, there
are already, in Entomology, specialists enough in all the higher
274 THE PANORPOID COMPLEX (Introduction),
Orders; so that the chance of being brought to a full-stop,
because nobody in the world can supply the gap required, is so
unlikely a possibility that it may be dismissed without further
thought.
That being so, T decided that the investigation into the Phylo-
geny of the Panorpoid Complex was a task that I might venture
to undertake, and that its difficulty and immensity would be
more than compensated for b\' the advantage to Entomology in
general, if a satisfactory solution could be found. Having, then,
decided to confine mvself to researches within the limits indi-
cated in the previous paragraph, I have carried out an exhaustive
surve}" of the older Orders under review, viz., the Planipennia,
Megaloptera, Mecopteia, and Trichoptera, and have been content
to study, in the immense Orders Diptera and Lepidoptera, mainly
the older families only. The survey has been extended far
enough to give me some idea of the position and inter-relation-
ships of all the important families of each of these Orders; but
intensive study has l)een chiefly confined to the older families,
together with other more specialised types in which I thought I
could recognise characters of value as evidence on the question
at issue. Further, it will be at once evident that the equipment
of the author who attempts to solve this immense problem will
not be complete unless he masters what there is to know about
the Fossil Record of the Orders in question, and of their possible
ancestral Orders. This also I have endeavoured to do. The
fortunate circumstance of my having in my hands a large amount
of new and unique material of this kind from the Permian and
Triassic strata of Eastern Australia has been one of the chief
factors in my decision to undertake this task; indeed, it has
almost imposed the obligation upon me, since nobody who has
not studied these fossils could possibly be so favourably placed
for discussing this question as I happen to be, simply through
this great good fortune.
It will, I think, be readily granted that the Order is the only
satisfactory unit upon which a study of this kind can be based.
We have, therefore, to consider what view of an Order we must
take, in dealing with it as a separate unit in a paper such as
this.
BY R. J. TILLYARD. 275
Now there are, in any given Order, many diverse groups.
Some of tliCvSe will exhibit very high specialisations in one or
more directions: others wdll show unexpected archaisms, which
may make it exceedingly awkward to frame a comprehensive
definition of the Order. As an example of this, we may take
the evolution of the mouth-parts within the Order Lepidoptera.
In the great majority of the families within this Order, the
mandibles are absent, the maxillary palps absent, the galeae
produced into a long sucking-tube or haustellum, and the labial
palps present and more or less highly specialised. But, in the
Alirropterygidce, the mandibles are present, as are also the typical
maxillae of older Holometabolous Orders, with five-jointed palpi,
and unspecialised galeae and lacinise There is, therefore, no
reason, from the Phylogenetic view-point, why we should not
consider the Lepidoptera as, ot bottom, essentially a mandibulate
Order, when we come to discuss its relationship with the other
Orders of the Panorpoid Complex.
It must, therefore, be evident that, for the purposes of this
paper, the usual definitions of Orders to be found in text-books
not only will not serve our purpose, but may actually be mislead-
ing. I propose to overcome this difficulty by the use of Arche-
types (German, Ur-typns). For each Order that comes under
review, we must define an archetype, which shall include in itself
all the most archaic characters found within the Order. Having
done this, we may reasonably discuss the relationships of the
archetypes of the various Orders, wuth some prospect of a suc-
cessful issue.
In following out this line of argument, we can lav down two
guiding principles : —
(1) The Phylogeny must not be determined from one set of
characters only (e.g., wing-venation), however important that set
of characters may be. But it must be determined by a review
of as many characters as possible.
(2) The greatest care must be exercised in the determination
of the characters of the Archetype. For instance, in the case
of the jugum and frenulum in Lepidoptera, it is not suflBcient to
adopt the jugum as the more archaic character, merely because
276 THE PANORPOID COMPLEX (Introduction),
it is found in those Lepidoptera which have the most archaic
wing-venation. If a mistake is made here, the whole Phylogeny
is bound to go wrong. As a corollary, it should follow that
characters in which the evidence of archaism or specialisation
may be uncertain, with a division of opinion upon the point
amongst those entomologists who have studied it, must be either
definitely cleared up by means of new evidence, or they must be
entirely omitted from the argument.
The Detej'mination of Archetypes.
We ma)^ exhibit this principle as follows : —
Let a, b, c, d, etc., represent a series of characters which
are constant throughout the Order in question.
Let p, q, r, s, . . . . etc., represent a further series of characters
which are not constant throughout the Order, but show definite
lines of evolutionary advance in one or more directions.
Taking any one of these characters ]>, after examination of its
structure in various families, let F represent the most archaic
condition, while p', ]"■> ^t^., represent various specialised condi-
tions of the same.
Any family within the Order can now be defined by the
characters
a, b, c, d, etc., p, q, r, s, etc.,
where p can have any of the values F, p', p'\ etc., q any of the
values (j>>, 5', q\ etc., and so on.
For instance, one family may be defined by
«, ^, c, f/, ;y, q\ R, s"\
The Archetype of the Order, which may or may not still be
existing (the greater the number of characters taken, the less
likely is it to be still existing) will be defined by
«, 6, c, c/, F, Q, F, 'S,
77te Determination of the Fhylogeuy of separate Orders.
The same principle as above may be followed in this case, pro-
vided we take the characters of the ArcJietype as the characters of
the Order it represents, for the purposes of the Phylogeny.
In this case, the characters a, b, c, d, etc., which were
constants throughout any single Order, will now be variables for
BY R. J. TILLYARD. 277
the d liferent Archetj^pes; also P, Q, B, S, not having the
same values for different Orders, we may omit these letters, and
consider that the whole series of characters is comprised in a
single (longer) series of variables a, b, c, d, e, J] etc.
Taking a group of Orders, we may now represent the most
archaic value of the character a hy A, while «', a\ etc., represent
various specialised conditions of the same. Then we shall be
able to write the Archetypes of different Orders in this way
Archetype of Order 1: - ^4, h", C, d', e, F, g" . . . (say).
Archetype of Order 2: -a, h\ C, d", EJ'\ G . . . (say).
Archetype of Order 3:—^, B, c', Z>, e",f\ g' . . . (say); and
so on.
The condition that any one Order may be ancestral to another
can now be determined by comparing the Archetype of the
supposedly higher Order with any supposedly ancestral group
lying within the bounds of the Lower Order. But, as the
characters of the Archetype of the supposedly ancestral Order
are even more archaic than those of the supposedly ancestral
group lying within that Order, the determination may be made,
once for all, by comparing the Archetypes of the two Orders.
The following rule may be laid down : —
The condition that one Order may be truly considered ances-
tral to another, is that there must not exist a single character in
the Archetype of the former, ivhich is more highly specialised than
the corresponding character in that of the latter.
For example, consider the case of Orders 1 and 3 above. The
character A is common to both Archetypes. For the character
6, the Archetype of Order 3 is the older {B against h"). But for
the character c, the Archetype of Order 1 is the older (C against
c). Hence, clearly, neither of these two Orders can be con-
sidered as ancestral to the other; and we must go back, for their
common ancestor, to an Order (probably no longer existing)
whose Archetype shows the characters A, Jj, C, . . . .
The Recognition of Embryonic ^Structures.
It would seem necessary here to call attention to the fact,
which seems to have been quite ignored by most of those ento-
20
278 THE PANORPOID COMPLEX (Tntroduction),
mologists who have not studied other Classes of Animals, that it
is not necessary to be able to trace back a structure to the
embryo without a break, in order to prove that it is derived
from an embryonic structure. Particularly does this apply to
appendages within the Phylum Arthropoda. Examples of this
are the well-known observations on the re-development of limbs
from latent embryonic rudiments in Crustacea, e.g., the man-
dibular palp in Decapod larvae, and the maxillipedes of the
Stomatopoda. In fact, the rudiments may appear in the embryo,
disappear later on, and then reappear again in the imago, as in
the case of the thoracic legs of the genus Chalicodoma and other
genera in the Order Hymenoptera.
Thus we see that it is not sufficient proof that a structure is
not derived from an embryonic rudiment to show that it does
not develop directly from that rudiment; whereas, it is, most
certainly, quite sufficient proof that it is so derived, if we can
trace it up from the embrj^o, through the larva and pupa, with-
out a break, to the imago (or, if the structure in question is only
a larval one, then it need only be traced from the embryo to the
larva).
We may take, as an example of this, Lubbock's observation
upon the development of the lateral gills in Chloeon, which, he
proved, did not begin to develop until the third instar of the
larva. Many entomologists have used this as proof that these
gills cannot be homologous with the original segmental abdominal
appendages, which are represented in the embryo, and then dis-
appear. This argument is quite fallacious, and must be care-
fully avoided, since it is a frequent temptation to use it. Further,
there is another potential fallacy in this kind of argument.
Chloeon is a highly specialised member of the Order Plectoptera.
May it not be quite possible that the break between the embry-
onic appendages and the formation of the gills, which is so con-
spicuous in this genus, may be very much reduced, if not com-
pletely removed, in the case of the development of the larva of
some much more archaic type, such as Onisciy aster, not yet
studied ?
13Y R. j. TILLYARD. 279
Meyrick's Law.
In 1893, Comstock(l) gave the name " Meyrick's Law" to the
following statement, which was first enunciated by Meyrick in
1884: — "When an organ has wholly disappeared in a genus,
other genera which originate as oftshoots from this genus cannot
regain the organ, although they might develop a substitute
for it."
Stated in this manner, this law may be accepted, provided it
is clearly understood what its limitations are. These mav be
stated as follows : —
(1) It must be capable of proof that the organ in question has
ivholly disappeared from the assumed ancestral genus. For in-
stance, a concealed rudiment of an organ may reappear, after a
more or less lengthy period of suppression during the course of
evolution of the group. It will be clear that this fact alone
must make the application of the law exceedingly problematical,
except in the case of very simple and easily examined structures
or organs.
(2) Great care must be exercised in determining what is to be
regarded as a substitute ior an organ, as contrasted with an actual
redevelopment of the original organ from a concealed rudiment.
(3) The converse of the law is not necessarily true; i.e., if a
genus B shows the loss of certain organs in comparison with
another genus A, we are not entitled to state, without further
evidence, that B is descended from A; we may only state that A
is more archaic than B in respect of the characters in question.
How little the importance of the safeguards in the original
statement of this law has been regarded may be gathered from
Meyrick's own later work. In 1895(3), we find him enunciating
three "laws" as his guiding principles in the determination of the
Phylogeny of the Lepidoptera. These are stated as follows : —
"(1) A'o new organ can be produced except as a modification
o/ some previously eoiisti'ug sti'ucture."
" (2) A Jost organ cannot be regained.'^
" (3) A rudimentary organ is rarely redeveloped.^' (p. 10).
Of these. No. 2 is Meyrick's Law restated without its original
safeguards. This, taken in conjunction with the other two,
^80 THE PAXORPOiD COMPLEX (Introduction),
forms a very generalised statement covering a very much wider
field than the original one, and open to far graver objections.
As regards Law No. 1, this can only be accepted as true in its
very widest sense, i.e., that there must be pre-existing living
material, or cells, as a basis from which any new structure is to
be developed; and, in that sense, it ceases to have any value,
and becomes a truism. With respect to Law No. 2, this ceases
to be true when the restrictions contained in the original state-
ment of the law are removed. To mention only one obvious
example. The thoracic legs of certain Hymenoptera (e^g , Chali-
codoma) appear in the embryo; they then disappear throughout
larval life, but reappear again in their complete form in the
imago. Thus, in this case, as in many others, "a lost organ is
regained"; and a bald statement to the contrary is not only not
a law, but it is not true. Moreover, it may not be argued that
the law is true when applied to Phylogenetic problems only, and
not to Ontogeny. For, if there is any truth in the Biogenetic
Law at all, it is certain that what occurs in Ontogeny is of the
same nature, and governed by many of the same laws, as what
occurs in Phylogeny; and if we frequently find that lost organs
are regained in the course of Ontogeny, then it follows that we
may by no means assert that the same possilnlity cannot hold
for Phylogeny. With respect to Law No. 3, it is only necessary
to remark that there is probably no Phylum in which rudiment-
ary organs are so often redeveloped as in the Arthropoda, and
that this is particularly true of appendages.
It would seem much wiser to do without any attempt to formu-
late laws (so-called) for the solving of Phylogenetic problems, and
to treat each case, as it presents itself, upon its own merits. In
most cases, for instance, it is quite capable of definite proof that,
in a given phyletic series, an organ originally present has been
lost, and never regained in any of the descendant genera. In
other cases, not so numerous it is true, it is capable of proof, by
reference to palaiontological evidence, that lost organs have been
regained, or, perhaps, that organs which, on the strict applica-
tion of Law No. 2, would have to be regarded as originally pre-
sent and subsequently lost, are really new developments not
BY R. J. TILLYARD. 281
present in the ancestral form. In such cases, a rigid application
of the law would result in a complete inversion of the phylo-
genetic conclusion.
Meyrick liimself applied his law chiefly to the problem of
wing-venation in the Lepidoptera. In a case like this, where
almost the whole course of evolution is towards reduction and
simplification, tliere is not much danger in its use. Yet even
here caution is necessary. It is not true, for instance, that
veins which have once been lost cannot be re-developed; nor is
it true that new cross-veins cannot be formed in a wing-area in
which no such structures existed in the ancestral form. One has
only to examine the tracheation of the pupal wing in the Lepi-
doptera to see that there is an immense field of possibilities in
both these directions, owing to the persistence of the finer
tracheae, ramifying in all directions beneath the wing-membrane.
As long as these tracheae persist, so long must the possibility of
an imaginal venational meshwork reappearing be held to exist.
Moreover, in a case in which the wing is changing its shape, so
that a certain part of it may become broadened, it is exceedingly
likely that one or more of the tracheae underlying the veinlets
of this area will become lengthened and strengthened, so that it
may eventually lead to the replacement of an original short vein-
let by a longer oblique branch, which, on Meyrick's interpreta-
tion, would have to be regarded as an original archaic branch of
the main vein from which it springs One of the best examples
of this is the effect of the widening of the costal area in the
Psychopsidce, in which an originally fairly simple series of
veinlets has become greatly lengthened and enlarged, most of its
units branching many times; and all of them connected together
by newly developed series of cross veins, which were certainlv
not present in the ancestral form.
The above example shows us that, even in so restricted a study
as that of Wing- Venation, Meyrick's Law can only be used with
great caution. It must be restricted to areas of the ivirtg that
are undergoing reduction. In the present state of evolution of
the Insect-Wing, it is certainly true that the tendency is towards
reduction in the great majority of cases. But this cannot always
282 THE PANORPOID COMPLEX (Introduction),
have been the case. Bearing in mind the complex nature of the
win'^-venation in the most archaic types of insects, we are bound
Xfo conclude that, at the beginning of the evolution of the wing,
there must have been a period of great enlargement, with a rapid
and abundant production of veins in all parts (following upon
the rapid increase in the tracheation of the developing organ).
After this had reached its maximum, there began a period of
arrangement and reduction, during which the very beautiful and
perfect wing-types at present existing were evolved. We may
term the process by which any part of a wing becomes broadened,
with consequent production of further venation in that area,
Platygenesis (Gr. TrAan's, broad); while the opposite process, in
which the wing-area in question becomes reduced or narrowed,
with consequent reduction or elimination of some of its existing
venation, may be termed Stenogenesis (Gr. (rTev6<;, narrow). The
development of the costal space in the Psj/chopsidce, of the
enlarcred anal area of the hindwing in Anisopterid Odonata, and
of the wide anal fan in the Orthoptera, Perlaria, and other
insects, are good examples of Platygenesis; while the process of
Stenogenesis can l)e followed out very fully in such an evolu-
tionary series as the Diptera Nemocera.
Scheme of the Work.
We have already stated that phylogenetic conclusions, to be
acceptable, must be based upon an examination of as many char-
acters as possible. In undertaking an analysis of a large number
of characters, two methods of procedure are possible. We may
either select each Order in turn and study its characteristics
fullv, with a view to the determination of its Archetype: or we
may select any given set of characters, as, for instance, those
offered by the Wing-venation, and study them as they are ex-
hibited throughout the whole of the Orders under discussion;
proceeding to deal with other sets of characters in subsequent
parts of the work.
This second method must obviously be the one chosen, for
only by it can the necessary comparisons be made between the
same sets of characters in different Orders. Thus we have to
BY R. J. TILLYARD. 283
postpone the definition of Archetypes until the whole of the
characters selected have been surveyed; and the final decision
as to the phylogenetic relationships of the various Orders will
not become fully apparent until the work is completed. Mean-
while, it seems advisable to select the characters for study in
such a way that the work can be subdivided up into a series of
parts, each complete in itself, and suitable for separate publica-
tion. Working on these lines, we may conveniently begin with
those characters which have been most used in phylogenetic
discussions, and proceed to deal with the rest in a definite order.
Thus we shall at first confine our attention to a study of the
Wings alone, and these will occupy two or more parts of the
work. Next to these, we shall take the Mouth-parts. Sub-
sequent parts will deal with other imaginal structures; and,
lastly, the structure of the larva and pupa will be reviewed. As
regards the structure of the egg, and the Embr^^ology, it is to be
regretted that so little is known about the latter in the case of
the more archaic Orders. As our knowledge stands at present,
the evidence available on this point, for the purposes of this
paper, is so incomplete, that it can have little bearing upon the
main result.
The investigations into this problem were actually begun
about two years ago, and are still going on. I feel that I can
safely leave the completion of the more specialiseci parts of the
work to a later date; because, if I waited until they were all
finished, the main conclusions of this work would not, perhaps,
see the light of publicity for some years. The results already
attained are, to my mind, of such importance that I have no
hesitation in placing them in the hands of the scientific; public,
and their value will not be lessened )jy delay in the completion
of the whole fabric of my woik.
BIBLIOGRAPHY.
1. CoMSTOCK, J. H., 1893. — Evolution and Taxonomy, in "The Wilder
Quarter Centenaiy Book." Ithaca, N.Y., ISl).'). pp..')?-! 14. iMej--
rick's Law, p. 47).
2. Handlirsch, a., 1908.— Die Fossilen Insekten. Leipzig, 1908. (Die
Phylogenie der Pterj'gogenea, viii. Abschnitt, pp. 1228-1318).
284 THE PANORPOID COMPLEX (Introduction).
3. Meybick, E., 1895. — Handbook of British Lepidoptera. London, Mac-
niillan and Co., 1895. (See p. 10).
i. , 1912. — Micropteryrjifhv in "(lenera Insectonim," Fasc.
132. Brussels, M. P. Wytsman, 1912. (See pp. 1-2).
5. TiJ.LYARD, R. J., 1917. — " Mesozoie Insects of Queensland. No. 1.
Planipennia, Triehoptera, and tlie new Order Protomecoptera."
Proe. Linn. 8oc. N. S. Wales, 1917, xlii., Pt. i., pp. 175-200. (See
p. 188).
6. . 1918. — "Permian and Triassic Insects from New South
Wales in the Collection of Mr. John Mitchell." I.e., 1917, xlii.,
Pt. iv., pp. 720-756.
7. Weele, H. W. van der, 1910. — Megaloptera, in Coll. de Selj^s, Fasc.
V. Brussels, 1910.
Note. — A bibliographj' will be supplied for each part, with the reference
numbers running consecutivelj'; but those papers which are not referred
to in any given part will be omitted from the bibiiographj' for that part.
285
ORDINARY MONTHLY MEETING.
June 26th, 1918.
Professor H. G. Cliapman, M.D., B.S , President, in the Chair.
A letter from Dr. T. Gritiitli Taylor, Melbourne, returning
thanks iov congratulations on the award to him of the David
8yme Prize for 1918, was communicated to the Meeting.
The President communicated some news of Lieutenant Gold-
finch, a Soldier-Member in hospital in England, forwarded by
Dr. A. J. Turner. It was resolved that a friendly message of
sympathy should be sent.
"o"
The Donations and Exchanges received since the previous
Monthly Meeting (29th May, 1918), amounting to 42 Parts or
Nos., 8 Bulletins, 3 Reports, and 12 Pamphlets, received from 39
Societies, Institutions, etc., were laid upon the table.
21
286
THE PANORPOID COMPLEX.
Part i. — The Wing-coupling Apparatus, with special
REFERENCE TO THE LePIDOPTERA.
By R. J. TiLLYARD, M.A., D.Sc, F.L.S., F.E.S., Linnkan
Macleay Fellow of the Society in Zoology.
(Plates xxix.-xxx., and sixteen Text-figures.)
There can be little doubt that, in the original Pterygote
Insects, fore- and hindwings were independent in tlight, and the
muscles controlling them were innervated from two separate
sources, viz., the ganglia of the meso- and metathorax respect-
ively. Coincidence of action or beat, then, between fore- and
hindwings, without which flight would scarcely have become
possible, must have been maintained through coordination of the
two sets of nervous impulses sent out from these two ganglia.
Now, in those Orders in w^hich the wings were more or less
hairy, the development of a fringe round the border of the wing
would undoubtedly increase the sense of touch in these organs;
since the macrotrichia, or larger seta? of the wing, are of the
type known as seufiilhe, and were evidently developed at first as
tactile organs. Hence it came about that, in the course of evo-
lution, a further coordination was able to be established between
the posterior portion of the base of the forewing and the anterior
portion of the base of the hind."**" Such coordination did not, at
first, take the form of a definite linking-up of the two wings, but
* It should be borne in mind that the structures here to be discussed
were originally present on both wings; e.(j., frenular bristles occur at base
of foreirinys of certain Mecoptera, as well as of hindwings; and a jugal
lobe is pi^esent at bases of hofh wings in certain Trichoptera and in Micro-
pterj/yldfc. But it is onlj' on the posterior border of the forewing and the
anterior border of the hind that thej^ can come into contact, and so develop
into a coupling-apparatus.
liY R. J. TILLYAKD. 287
consisted merely in the greater development of the sensilla? of
this region of the two wings, their contact acting as a guide in
flight, much as the reins act in the driving of a horse, or the
touch of the hand of one person in guiding another in the dark.
To these structures, whether they act merely as a yiridf, or
whether they are more fully developed so as to fink the two
wings quite closely together, T propose to give the name "Wing-
coupling Apparatus," which 1 have already used in dealing with
the Planipennia(12).
The complete, archaic wing-coupling apparatus consists of four
distinct parts, two belonging to the forewing, and two to the
hind. These are: —
In the forew ing
(1) The area of contact with the hind wing or its bristles.
(2) The bristles developed along the border of that area.
In the hind wing
(3) The area of contact with the forewing or its bristles.
(4) The bristles developed along the border of that area.
In the case under discussion, i.e., when the two areas of con-
tact are located at the bases of the wings, we may name the
parts as follows : —
(1) The area of contact of the forewing with the hind, in so
far as it projects beyond the general contour of the posterior
border, may conveniently be termed the jnyal Johe, a term I have
already employed in the paper mentioned abo\e(l2). A special-
ised form of this area, in which it becomes an elongated, narrow
process passing hplouo the costa of the hind wing, is termed the
juyum, this being the term fii'st applied to such a process by
Com stock in the Lepidoptera(l).
(2) The bristles projecting from this area towards the hind-
wing may be called the jnya/ hristlpg.
(3) The area of contact of the hindwing with the fore, in so
far as it projects beyond the general contour of the anterior or
costal border, may be termed the humeral lobe. I had previously
employed the term jnyal jji-oce.s.<i for this area in the Planipennia
(12). But I now consider that this term is open to objection; as
288
THE PANORPOID COMPLEX, 1.
it would certainly conduce to clearness, if the adjective jncial
were to be employed only in connection with the forewing. As
authors who have written upon the Lepidoptera have generally
alluded to this part of the hindwing as the shoulder or humerus
(a somewhat too general term), there can be little objection in
replacing it by the term here proposed.
(4) The bristles projecting from this area towards the forewing
may be called the fremihwi. This term is already in general use
in the Lepidoptera, and should now be adopted for ihe homo-
logous structures in other Orders,
p The relationships of these
four parts to one another, in
an ideal archaic coupling-ap-
paratus, may be seen from
Text-fig. 1, it being under-
stood that, owing to the
slightly higher level of the
Text-fig. 1 . * forewing, the frenulum passes
beneath the jugal lobe, but the jugal bristles lie above the costa
of the hindwing.
We may now profitabl}^ study the formation of the coupling-
apparatus in the wings of the diiierent Orders comprising the
Panorpoid Complex. From this discussion, we are compelled to
omit the Protomecoptera (in which the bases of the wings have
so far not been discovered in the fossils known), and the Aphani-
ptera, in which the wings have been lost.
Order MECOPTERA. (Text-figs. 2-4).
This is the only Order extant in which all four parts of the
ideal coupling-apparatus can still be recognised. In Text-fig. 2,
I have figured the coupling-apparatus from the wings of two
very archaic families, found only in Austi-alia. In the family
Choristidce (Text-fig. 2a), there is a slightly projecting, but quite
* Ideal archaic wing-conpling apparatus at bases of wings: //•, frenulum;
Fw, forewing; ///, humeral lobe; Hm, hindwing; jh, jugal bristles; ,//, jugal
jobe. (Jugal bristles rest above costa of hindwing, but frenular bristles
pass beneath jugal lobe of forewing),
BY R. J. TILLYARD.
289
definite, jif{/a/ Johp ( //), from the more distal portion of whicli
arises a set of about a dozen small, but stiff and closely set, ji'(j<il
J)ristlp)< (jh), which make contact with the costa of the hindwing
in flight. In the hindwing, there is a small but quite definite
hiviupral lobe (hi), from which spring the two enormous bristles
of the frennl urn (fr). These latter, during fiight, joass under —
and, indeed, quite beyond the interior border of — the jugal lobe
of the fore wing. The whole apparatus is eminently suital)le for
maintaining contact between the two wings in flight, without in
any way linking the hindwing /??•;?? /y to the fore.
In the wings of the family Xannoehorlstidiv (Text-fig. 2/>), which
are specialised by reduction, w^e ty.
meet with much the same struc- il
tures. But in this case there is a
single, strong, jugal bristle on the
forewing, instead of a set of
weaker bristles. The flight of q^
these insects is the strongest of // y\ ■■••... r-'
any in the Order, and the wings J
are closely linked together. The jl
linking is accomplished by the /^
two sets of bristles. The frenu-
lum passes under the jugal lobe, 7 . f k/'"'"^^ -jb
while the jugal bristle passes ^'
above the bases of the frenular ^^^,>'-^^:^:3^3Lijy'h<'^<^><K,^f
bristles, and presses down upon
the costal area of the hindwing,
thus forming a very neat and
perfect coupling-apparatus. ^^ ' ^'"
In the archaic J/^roy^?V/r^(Text-fig.3), there is a peculiar special-
isation not found anywhere else, to my knowledge, within the
Insecta. The jugal lobe beai-s a very distinct, black tubercle,
projecting from its border. This tubercle appears to have been
* Wing-coupling apparatus in Mecoptera. a, In T<vniocliorisfa pallida
E.-P., wings connected, viewed from beneath. I>, In Xannochorida dipter-
o/ri(N Till. , wings disconnected. (Both ;< 88). Lettering as in Text-fig. 1.
290 THE PANORPOID COMPLEX, i.,
formed by fusion of an original set of jugal bristles, as seen in
Chorista. A frenulum is also present, but reduced in size.
In all other representatives of this Order, the jugal bristles
appear to be either absent or very weakly formed; and the whule
coupling-apparatus shows a
gradual reduction from dis-
use, probably owing to a pro-
gressive weakening in the hi
tlight of these somewhat lazy
insects. Text-tig. 3. *
The culmination of this tendency is to be seen in the highly
specialised HUfaridce,, in which fore- and hind wings have again
il ^-ii-^i.*;-^^ returned to complete independence in
"^"■''^'^^^^' llight, the wings becoming greatly nar-
rowed and elongated, with petiolation
'^ of their bases. The coupling-apparatus
D •■' ^^V/^ is here only represented by a vestige of
the humeral lobe, from which projects a
W^ sinule frenular bristle, which ci-osses a
^S,..,^^ similar bristle developed from the ex-
Text-tif'.4.t treme base of the forewing (Text-fig. 4).
Reviewing the above evidence, it is clear that the Archetype
of the Order Mecoptera must have had the complete archaic
coupling apparatus fully developed, very much as it still exists
in the ancient Choriatidm to-day.
Order PLANIPENNIA. (Text-fig.5).
In this Order, the coupling-apparatus is again best developed
in the more archaic families, notably in the Ilpvierubiicke (Text-
fif.5). Jugal bristles are absent; but the frenulum is repre-
sented either by one or two strong bristles {Jr). These bristles
* Wing-coupling apparatus in Merope tuber Newm,, viewed from above,
showing the large, black tubercle at l^ase of forewing: /;/, humeral lobe;
( X 40).
t Reduced wing-coupling apparatus in HarpobiUacus tWyardi E.-F :
}), bristle developed from extreme base of forewing; other lettering as in
Text-fig. l;(x 40).
BV R. J. TILLYARD. 291
project from the apex of a very strongly developed humeral lobe
(h/). During Hight, this latter lobe, together with its frenulum,
passes under the jugal lobe of the forewing, which is hollowed
out beneath for its reception.
The same type of coupling-apparatus is to be found in all the
other families, but it is generally much
reduced in size, and does not appear to be
functional. The tendency in the evolution
of this Order has been to combine a pro-
gressive narrowing of the wings with a
gradual return to complete independence
of fore- and hindwing in flight. In the
highest families, Mynnelaonfidce and Asca- Text-fig.o."
laphid(f., the bases of the wings become shortly petiolate, and the
original coupling-apparatus can only be made out as an extremely
reduced renniant at the very bases of the wings.
For this Order, then, we must assume an Archetype in which
the archaic coupling-apparatus was complete in all except one
particular, viz., that the jugal bristles were absent.
Order MEGALOPTERA. (Text-figs. 6-7).
Throughout this Order, the hairy vestiture of the wings, both
macro- and microtrichia, has been very much reduced. The most
archaic type of coupling-apparatus known to me within the Order
is to be found in a genus not yet described, but allied both to
the Corydalidce and the Sialidcn. This possesses both jugal and
humeral lobes, of small size, the latter with a few small hairs,
the remnants of the frenulum. In Sialis, the two lobes are
present, but apparently hairless.
In the Corydalidce (Text-fig. 6), as exemplified by Archicha all-
odes, the only genus that I have been able to study, the humeral
lobe is much reduced, but the jugal lobe becomes a definite
angular projection (,y7) which presses down upon the costa of the
hindwing. Further, the forewing overlaps the hind for a con-
* Wing-coupling apparatus in Drepanacra humilis McL., viewed from
beneath; ( x 24:). Lettering as in Text-fig. 1.
292
THE PANORPOID COMPLEX, 1.
siderable distance beyond the base; and, by pressing down upon
it, serves to increase the strength of the coupling in this rather
puwerfully-riying insect. This latter type of wing-coupling,
which occurs also in many Trichoptera, and in some of the most
Text-lig.CJ.
Wing-coupling apparatus in Archichau/iode-'i (jitftiftrns Walk., viewed from
above; ( x 12). A, beginning of amplexiform overfold; jJ, jugal lobe.
highly specialised Lepidoptera, I propose to term ainpl>;.rAjurm
(Lat. aiiiplf'.cits^ an embrace). It is always correlated with the
disappearance of the frenulum.
--^hT"^
Text-tig. 7.
Reduced wing-coupling apparatus in Raphidia macidicoUis Steph. ; ( x 40) :
A/, humeral lobe; ,//, jugal lobe; f, roughened tubercular patch.
In the highest family uf the Order, the terrestrial Iiaj>hidiid<p,
both jugal and humeral lobes are present, but without Ijristles.
There is, however, a very peculiar patch uf roughened tubercles,
situated on the posterior margin of the forewing, just beyond the
jugal lobe (Text-fig. 7,^) This may })ossibly help in njaintaining
contact between the wdngs in tlight.
BY R. J. TILLVAHD. 293
From the above evidence, we iinist presume tliat the Archetype
of the Megaloptera possessed a somewhat reduced coupHiig-appa-
ratus, ill which the two lobes were present, the jugal bristles
absent, and the frenular bristles probably (piite vestigial.
Order TRICHOPTERA. (Text-figs. 8-9).
In this Order, the original wing-coupling apparatus undergoes
some remarkable developments, which have attracted very little
notice from entomologists, so far. In all the oldest families, one
can only recognise the jugal lobe, which is the anal lobe, of sys-
tematists in this Order. This appears as a strongly projecting
lobe at the base of the forewing, in such genera as EhyacophUa
(Text-fig. 8, J/). There are neither jugal bristles nor frenulum;
and the humeral lobe is suppressed, or else only slightly developed.
In no case does this jugal lobe pass under the hindwing during
flight; it merely rests upon the costa from above.
In the great majority of genera, the jugal lobe is absent or
vestigial, and new elements of wing-coup- ''**it5j— __^^<<^
ling have made their appearance. This \- ; 7^^.,^.
iieAv type of wing-coupling is awy^/e-t^/orz/i, i[-A;'',' y^
i.e., it is brought about by an overfold of ^..^""^^ ""
the whole length of the anal area of the *" ~~"
fore wing upon the costa of the hind. But
besides this, there are many cases in Text-tig. 8.*
which the amplexiform coupling is strengthened by one of two
new devices, as follows: —
(1) Between the anal vein of the fore wing, and the anal border,
there may be developed a single row of stiii" hairs, all pointing
forwards and outwards, on the underside of the wing. The row^
of hairs upon the costal margin of the hindwing is also stiliened;
so that, during flight, further strengtli is given to the amplexi-
form coupling by the intermingling of these two sets of stiif
hairs. I have seen this type of coupling in an archaic Austra-
lian genus of tlie family Fhryyaiteidce, not yet named.
* Wing-coupling apparatus in Rhyacopltila dorsal U Pict., viewed from
above; ( x 12) : jl, jugal lobe.
294 THE PAXORPOID COMPLEX, 1.,
(2) Much more commonly, and especially noticeable in the
families Lppfocerid(P- and Sericostvmatiche, is a type of coupling
whicli T propose to term multihamidate (Text-fig. 9). In this,
the row of hairs along the costa of the hindwing becomes special-
ised, for a greater or less length, by the de\'elopment of the hairs
into stiif hooks, which grapple the anal margin of the forewing,
and so bring about a very perfect form of wing-coupling. Tliis
type will be recognised as that which also occurs universally
throughout the Order Hymenoptera.
Text tig. y.
Row of booklets developed along the middle portion of the costal margin
of the hindwing in Otcefi-s sp., (Australia); ( x 83).
There are many genera in this Order in which certain of the
macrotrichia become developed into series of large, stiff bristles,
closely resembling those of a typical frenulum. These most
frequently occur along the main veins, or portions of them.
Bearing this in mind, I have searched very carefully for a true
fienulum in this Order, but I have not succeeded in finding one.
In tlie males of the remarkable and highly specialised genus
Plectrotarsits, whose systematic position within the Suborder
Inaequipalpia is still a matter of dispute, I have found a set of
two or three strong bristles in the position of the frenulum. But
they are absent in the female, and they play no part in the
coupling of the wings, which is of an advanced amplexiform type,
with the jugal lobe quite small and non-functional. In two other
very spiny genera, viz., Chimari'ha, and an unnamed Australian
genus having whorls of numerous spines upon the tibite, there
are also stiff" bristles in the position of the frenulum, at least in
the males. Those of the former genus are long and slender, and
are arranged in two or more irregular rows, quite unlike a true
frenulum; while those of the latter are also very long and slender,
irregularly placed, and lying flat along the wing, parallel to the
costa. In neither case do they play any part in wing-coupling.
BY R. J. TILLYARD. 295
These developments must be regarded as isolated specialisations,
which must be expected to occur throughout an Order in which
all parts of the wing remain so well provided with hairs as in
the Trichoptera.
From the above evidence, we must conclude that the Arche-
type of the Trichoptera already possessed a highly reduced
coupling-apparatus, in which only one of the four original paits
w as represented, viz., the jnyal lob'-. This type may be designated
as the archaic jnyati' type of wing-coupling.
Order DIPTERA.
In this Order, owing to the loss of the hind wings, there is no
longer any need for a coupling-apparatus. But we can recognise
the jugal lobe of the forewing, in the form of the basal lobe
known as the aJala in this Order.
It is clear, therefore, that the Archetype of the Diptera re-
sembled that of the Trichoptera in possessing the archaic jugate
tijpe of wing-coupling, in which only the jugal lobe is present.
AVe might note here, parenthetically, that, in the other Holo-
metabolous Order (the Coleoptera), in which flight is carried on
by only one pair of wings, there may still be found evidences of
the original presence of a jugal lobe; e.g., in certain Hydro-
philidfP, where this lobe is quite large and conspicuous, though
it does not seem to perform any definite function.
Order LEPIDOPTERA.
(Text-figs. 10-16, and Plates xxix.-xxx., figs. 18).
We have kept this Order to the last, because, within it, there
are developed the most surprising and interesting of all the
specialisations arising from the old type of basal coupling-appa-
ratus, which we have already studied in the other Orders of the
Complex.
It has been frequently stated that certain families of INIoths,
viz., the JJicropterygidce {sena. /at., including the Eriocraniidcp)
and the HepiaUdcH, are distinguished from all other Lepidoptera
by the nature of their wing-coupling apparatus. To these, Mey-
rick(9) has lately added a third family, the Frutotheoridce, which
296 THE PANORPOID COMPLEX, 1.,
he claims to be intermediate between the other two. These
families together form the 8uborder Jiigatfp* of Comstock, or the
group Micropterygina of Meyrick. The latter author (3, p. 7 97)
thus defines the character of the wing-coupling apparatus in this
group:— "Fore wings with an oblique membranous dorsal process
( jugum) near base, forming with the dorsal margin a notch or
sinus, which receives the costa of the hindwings. Hindwings
without frenulum."
This may be taken as a standard definition of the jug<ife type
of wing-coupling; provided that we add what is perhaps not
quite apparent at first sight, that this true jugum passes under
the costa of the hindwing, and so forms the notch or sinus above-
mentioned, in which the costa is held as in a finger-and -thumb
grip. The nature of this coupling-apparatus is best understood
by ref ei-ring to Text-fig. 1 1 .
As contrasted with this J>'[/"-fe type, the rest of the Order is
usually regarded as forming a single Suborder Frenatae,"^ in which
the majority of the families possess a frenulum, but no jugum. A
good definition of this character has been given by Meyrick (3,
p. 4): — "Generally from or near the base of the costa of the hind-
wing rises a stiff bristle or group of bristly hairs, termed the
freiinlfon, of which the apex passes under a chitinous catch on
the lower surface of the forewing, termed the retinaculum^ thus
serving to lock the wings together; the frenulum is commonly
single and strong in the male, multiple and weak in the female;
the retinaculum in the female is commonly represented by a
group of stiff scales."
However, certain families of the Frenata, including the great
division Rhopalocera or Butterfiies, do not possess this frenulum.
In these forms, the coupling of the wings in fiight is brought
al^out by the downward pressure of the posterior margin or
dorsum of the forewing upon the costa of the hind, the latter, on
its part, pressing strongly upwards from beneath. The humeral
* I fail to understand the use of the feminine plural, since tliese w'ords
are adjectives, and should agree with the neuter phual noun Ltpidoptera.
I propose, therefore, in future, to write them Jngata and Frenata, respect-
ivelj-.
BY R. J. TILLYARD. 297
lobe of the hindwing is generally fairly large in these forms, thus
allowing a greater area of the hindwing to pass under the fore
than would be the case if it were normal in size. Thus it will
be seen that the type of coupling here represented is that which
I have already termed, in the Megaloptera and Trichoptera,
amplexiform. I propose to use this term to distinguish these
groups from those possessing a frenulum, without in any way
thereby indicating a belief that the amplexiform groups in the
Lepidoptera constitute a single line of descent. For it must be
evident that thei'e is no reason why the frenulum may not have
been lost independently along several distinct lines of descent,
leaving us at the present time with several isolated amplexiform
groups, each of which has its nearest relationship, not with the
other Amplexiformia, but with a different frenate group.
As an illustration of this, we find, in the Ca^tiiiidce, a family
in which both the frenulum and the humeral lol)e are well de-
veloped. From such a group, either a typical Frenate group or
a typical Amplexiform group might be descended; the former
b}^ reduction of the humeral lobe, the latter by loss of the
frenulum.
Considering that the condition of the wing-venation is of
greater importance than the structure of the wing coupling appa-
ratus, for the purposes of classification, I have already, in a pre-
liminary leport on this research (13), suggested that the primar}^
division of the I^epidoptei-a into two Suborders should be based
upon the former instead of the latter. I therefore defined two
Suborders as follows: — -
i. Suborder HOAIONEURA.
Venation of fore- and hind wings closel}- similar and of primi-
tive design. (Families Microjderyyidtt {senst. /at.), I'rototheoridcf',
and Hepialidce).
ii. Suborder HETERONKURA,
Venation of hindwing reduced, and difi'ering widely from that
of the forewing. (All the other families).
The reason for discarding the older terms Jugata and Frenata,
298 THE PANORPOID COMPLEX, i.,
in favour of this new division, will be more fully appreciated
when we have completed our study of the Microptprycfidw.
Adopting, then, the above terminology, we may now proceed
to study in more detail the type of wing-coupling apparatus to
be found in the two divisions or Suborders.
i. Suborder HOMONEURA.
Family Micropterygid^ i^»ens. J at.).
(Plate xxix., figs.1-4, and Text-fig. 10).
In this family, I made preparations from the wings of all five
genera available to me, viz., Sahatinca, Micro'ptfivyx, Mnemonica,
Eriocrania, and Mnefiarchcea.
I should like here to thank the many kind correspondents who
have supplied me with specimens of this family, at all times diffi-
cult to obtain; and, in particular, Mr. Meyrick, for a series of
set specimens representing all five genera; Mr. A. Philpott, for
various New Zealand species, both set and in spirit, and for his
valuable observations on the flight of Sabatinca; and Mr. K. J.
Morton, of Edinburgh, for a large number of spirit-specimens,
which yielded excellent mounts and dissections.
In dealing with this family, I was at once struck by the
behaviour of the so-called jugum in cleared and mounted speci-
mens. Instead of arranging itself as a free lobe at the base of
the dorsum of the forewing, I found that it always insisted upon
remaining twisted under the wing itself, and that this position
could only be altered with difficulty, by drawing the jugum out
from under the wing with a needle. (Plate xxix., fig.l, and Text-
fig. 10a, c/).
Referring to figures of this jugum already published, I found
that Comstock(l), Forbes (8), and Meyrick (3, 4) had all figured it
as a projecting lobe, whereas Packard (10, 11) alone had depicted
the true position; though without, apparently, any idea of its
significance.
Turning next to the examination of cleared and descaled
mounts of the hindwing, I found invariably from three to six
lar^e bases of insertion (Plate xxix., fig.3, and Text-fig. 10, h.fr.)
BY R. J. TILLYARD.
299
of a circular shape, situated along the costal border in the region
of the humeral lobe, this latter being only a slight prominence.
Comparing these with the bases of insertion of neighbouring
scales (sc), I found that they were of very much greater size, as
well as of more regular shape. It was evident, therefore, that
they were not the bases of either typical scales or of hairs, but
that they must belong to a series
of strong bristJpx, which had been
removed during the process of
descaling. This led me to infer
the presence of a true Jrenuhi
in this family.
The next step was to examine
set specimens from the underside,
under a fairly high power of the
microscope. To do this, I clipped
off the pin close to the underside
of the thorax, and then inserted
the head of the pin into a piece
of indiarubber, so that the moth
was exposed ventral side upper-
most. By this means the lens
could be brought close down upon
the wings, without touching the
pin. Text-%.10."
The immediate result of this examination was the discovery of
a set of strong bristles, nearly always three ov four in number,
more rarely two, five, or six, constituting a trufi frenuJum, but
directed outwards and upwards at a very slight inclination to
" Wing-coupling apparatus in Sahatinca inronnrueJla Walk., (New Zea-
land, fam. Micropterygicht') : a, base of dorsum of forewing in ? , viewed
from beneath; { x 88): J>, base of costa of hinrlwing in ? (descaled); ( x 83):
c, portion of the same enlarged, to show insertions of frenular bristles
[h.fr.) and insertions of scales (.sf.) from a part of the same wing for com-
parison; ( x 320): d, bases of wings in 6, the forewing viewed from
beneath, the hindwing from above; ( x83): fr, frenulum; Id, humeral lobe;
jl, jugal lobe. {Cf. Plate xxix., figs. 1-4).
b.fr
300 THE PANORPOID COMPLEX, i.,
the costa of the wino- (Text-fig. 10,//-). // is tJiPRc brisfhs, and
not tlip cofifa of thp hind?oing ifi^elf] ichirh hpcomp pnyagpd in fliP
dit or sinus formpd hphvppn the juyum and dorsimi of tJiP forpiciny,
Altlioiigh the difficulty of setting these tiny moths in the natural
position must be veiy great, I actually found three specimens in
which one or both wings had become engaged in the natural
manner; and one of these came from Mr, Meyrick's own col-
lection.
Wishing to have my result tested in the field, I wrote at once
to Mr. A. Philpott, of Invercargill, X.Z., asking him to investi-
gate the coupling-apparatus in the wings of the living insect, and
sending him careful drawings of my results. The onlj^ species
that he was able to stud}^ was a new species of Sabatinca, of
which he reported that the coupling apparatus was exactly" the
same, in structure and position, as that of >S'. inconyrup/Ia, repre-
sented in m}" drawings. He also added some valuable notes on
the habits of flight and rest in this species.*
We come, then, to the following conclusions with regard to the
Jlicroptpvi/g ida- : —
(1) In so far as it is a fact, that a definite and well-developed
frenulum is present in this family, the}" are of a frpnate rather
than lijngafp type.
(2) In so far as the so-called jugum does not pass backwards
under the hind wing, so as to engage the costa in a finger-and-
thumb grip, it is not a true jugum, but must be termed a juyal
lobe; also, for the same reason, these moths are not true jugate
types.
(.H) Since the jugal lobe is turned forwards under the forewing,
and acts as a catch for the frenulum, it is clear that it should be
regarded as an archaic form of rptlnacidum, analogous to, but
not homologous with, the retinaculum found in the true Frenate
forms.
(4) From this, it will be seen that the Microptpvygida- combine
in themselves certain characters belonging to both Jugate and
Frenate types.
* These are given in an Appendix to this Part, on p..'J18.
BY R. .1. TILLYARD. 301
I propose, therefore, to designate the kind of wino-coupHng
apparatus found in tliis famil}^ as Jugo-frenatt\
Apart from the speciahsation shown in the actual nnderfolding
of tlie jugal lobe, and the consequent alteration in the direction
of tlie frenular bristles, so as to lie more nearly parallel with the
costa of tlie hindwing, it will at once be seen that this type of
coupling is the exact homologue of that found in the older Orders
^lecoptei-a and Planipennia. It agrees more closely with the
latter, since both have lost the jugal bristles; but differs from it
in the larger number of frenular bristles, and in the less definite
development of the humeral lobe.
Thus we come to the logical conclusion, that the archaic jugo-
f renate type, found in the older Orders of the Panorpoid Complex,
is represented, at the very base of the Lepidoptera, by a some-
what more specialised jugo-f renate type, in which the retinacular
nature of the jugal lobe, already existing in the Planipennia, is
more accentuated by nnderfolding, so that the fienular bristles
are held in a firmer grasp.
Having now fully reviewed the position in the Micropfn'i/c/irhf,
we may pass on to consider the more specialised families.
Families H E p i A L i d je and P k o t o t h e o r i d ^ .
(Plate XXX., fig.5, and Text-figs. 11-1 2).
In the J/ppia/id(f, I have studied all the Australian genera
available, together with the Pala^arctic genus IL-piahift, of which
I possess a number of specimens. In the FrofofJiPorichf,
through the kindness of Dr. Pc'ringuey, Directoi* of the South
African Museum, Capetown, I have received two specimens for
study, one of which proved to belong to Mr. Meyrick's Proto-
tht'ora pplroHoma (9), while the other appears to represent a new
species, not yet described.
The typical piyum found in the great majority of the Hepi-
alidd-, and also in the genus Prototlipora, is shown in Text-fig. 11,
and also in Plate xxx., fig.o. It is a long and fairly stiff finger-
like process (in most species carrying very long and abundant
hairs), which projects well below the costa of the liindwing
during flight, and engages it in a strong finger-and-thumb grip.
22
302
THE PANOKPOID COMPLEX, 1.,
The resulting flight is, in most cases, exceedingly swift and
strong; indeed, there is one Australian species common enough
about Sydney, Perisspctii< ausfralasice Don., in which the males
fly so swiftly as to be
almost invisible, and
are most difticult to
catch. One has onl}^
to watch, too, the man-
ceuvres which both
sexes carry out during
courting, to be con-
vinced that dexterity
Text-fig. 11.* of flight is carried very
far in this family, and is certainly not surpassed by the greater
number of Heteroneurous forms. We may, therefore, regard
this true jugate type as a high specialisation, very superior to the
jugo-frenate type of the Micro2)terygidce.
In the genus Prototheora, I find no difference from the typical
jugate type of the Hppialidcf. The jugum is long and narrow,
and the costa of the. hind wing is quite devoid of bristles.
In the new (undescribed) species of the Prototheoridce which I
examined, there is a more primitive type of jugum, in the form
of a triangular lobe, only slightly more prominent than that to
be seen in many Trichoptera. As in this latter Order, this lobe
does not act as a true jugum, but appears to rest upon the upper
surface of the hindwing, just overlapping the costa. I do not
know^ whether this character is a constant for this species, having
only examined one specimen. It should be noted that, in study-
ing a long series of HepialidcE, a very similar development mani-
fests itself occasionally in one or both wings, especially in the
genera Charagia and Poriiia. Such occasional developments are
to be regarded as strictly atavistic, and indicate the evolution of
the highly specialised J^i^itwi from the o\dev jug cd lobe.
* Wing- coupling apparatus in Charagia eximia Scott, viewed from
beneath, with hairs removed; ( x 12): j, jugum; j*'^, jugal sinus. [Cf. Plate
XXX., fig.o).
BY R. J. TILLYARD.
303
Some interesting deviations from the normal jugate type ma}'
now be noted in the Hejnalidce : —
(1) In the genus Pielu.% the jugum is a hard chitinous rod,
somewhat truncated at its tip, and placed so as to project de-
finitely beneath the anal area of the forewing (Text-fig. 12). It
is quite impossible to make this rod pass under the costa of the
hindwing, either in the living or freshly-killed insect. But it
will fit quite easily abovfi the costa of the hindwing, near the
base, where there can be found, in specimens that have been on
the wing, a deep groove worn in an oblique direction in the dense
hairs that clothe the area around the humeral cross- vein.
It would appear, then, that this genus differs from the rest of
the family in having the juguin lesting above the costa, not
beneath it, and also in the structure and position of the organ
itself. We should, therefore, regai'd this genus as highly
specialised.
(2) l7i the genus Lpfo, some specimens show a very weakly
formed jugum of very
small size for the bulk
of the insect: while
others have none at
all. The bases of the
wings are densel}'
clothed with long, soft
hairs. The method of
coupling in these mag-
nificent insects, which
are said to have great Text-fig. 12.^
powers of flight, appears to be practically amplpxiform, the anal
field of the forewing being bounded anteriorly by a ver}^ strongly
developed lA, and overlapping the costa of the hindwing to a
considerable extent in flight.
(3) In the small moths of the genus Frau.% the jugum is very
short, yet it undoubtedly secures the costa of the hindwing in
* Jugum of P ie/ as hi/a/inatii)i Don., 9 , viewed from beneath, to show
natural position under anal area of forewing; ( x 6).
304 THE PANORPOID COMPLEX, i.,
the typical manner. Of the New Zealand Porina nmhracnlata,
Mr. Philpott writes, "the jugum is so small as to seem worthless
as a coupling in the manner of finger-and-thumb."
Thus we see that there is a considerable diversit\" in the form
and function of tlie jugimi, as developed in the Hcpialklfc.
The discussion as to how the highly specialised jugum of the
Hefialidcv is related to the jugo-frenate type of coupling-appa-
ratus found in the Microptpryciicht' is best left until we have
dealt with the Heteroneura.
ii. Suborder HETERONEURA.
A. Frenate Types. (Plate xxx., figs.6-8, and Text-figs. 13-1 6).
Mey rick's excellent definition of the frenate tj^pe has already
been quoted on p. 296. We ma)^ now profitably study, in more
detail, the difi'erences between the female and male types.
These differences are clearly correlated with differences in the
power and frequency of flight. In those forms in which the
female flies but little, the frenulum remains small, and composed
of a number of separate bristles, while that of the strongly-flying
male may be very large and strong. But in other forms, in
which both sexes fly almost equally well, the frenulum of the
female may be almost as strongly formed as that of the male,
though never consisting actually of a single bristle.
Text-fig. 13.
Wing-eoupling apparatus of Hippotion i^cro/a {Boisd.), 9 , showing fren-
ulum of eight separate bristles, and retinaculum of hairs arising from
cubitus (C'?0; viewed from beneath; ( x 16): h/, humeral lobe.
Text-fig. 13 shows the frenulum of a female Sphingid {Hippotion
scrofa Boisd.). The bristles composing it are eight or nine in
IBY R. J. TILLYARb.
305
number, all quite separate, but closely set together, as shown in
the figure.
The ret'inaciihim^ or catch, whicli holds these bristles in place,
is a very generalised structure, being nothing more than the
somewhat stittened haii's or scales which arise from the underside
of the cubital vein of the fore wing, and run oblique!}^ forward
and distad. The manner in which the fi'enulum is held in place
by these hairs is closely similar to that by which the jugum of
Pirhis is held in its groove by the hairs on the costa of the
hindwinu'.
Text-fig. 14.
Wing-coupling apparatus of Plusia rerticillata (Guer.), ? , showing fren-
ulum of two strong bristles, and retinaculum of stiff hairs arising from
cubitus [Cii); viewed from beneath; ( x30h hi, humeral lobe.
In the Noctuid genus Flusia, in which males and females fly
almost equally well, we may see a much more liighl}' specialised
coupling-ap[iaratus in the female. Here, the frenulum is com-
posed of only two (sometimes three) bristles, apparently formed
by the fusion of an original greater number, and almost as long
as those af the male. The retinaculum, too, is more strongly de-
veloped, the hairs being stiffer and more closely set, so as to form
a very definite catch for the frenulum (Text-fig. 14).
Turning next to the males, we find, in every case, a highly
specialised condition present (Plate xxx., fig.8, and Text-fig. 15).
Here, the frenulum is represented by a single huge bristle,
which may even reach to more than half the length of the wing,
as in the family Fsychiche. As Packard (10) has already shown,
306
THE PAXORPOID COMPLEX, 1.
if this apparently single bristle be cut acnjss near its base, the
separate interior channels of the bristles of which it is actually
composed can be easily recognised. Thus, we see that the large,
male frenular bristle is not formed by liypertrophy of one of the
original series at the expense of all the rest, but by fusion and
lengthening of the whole set. The longitudinal Huting visible
along the somewhat fiatteiied basal part of this bristle is also
evidence of its composite nature.
Text-tig. J d.
A\'iiii;-coupling apparatus of Hippotio)i scro/a (Buiyd.), ^ , showing single,
strong, frenular bristle, engaged in hasp-like chitinou;; retinaculum
developed from the radius (B); viewed from beneath; ( x 14); hi,
humeral lobe.
The ?rfi'nacii/fwi in the male is a highly specialised structure,
having no homologue in the female, or in any other insect. It
consists of a strong, curved, chitinous catch, directed posteriorly
towards the hind wing, and developed from the underside of the
strong radius of the forewing, not very far from its base. This
structure is hasp-shaped, and forms a very etiicient catch for the
frenulum, which is able to slide to and fro within its grasp,
during ilight, without running any risk of becoming detached
(Text-fig. 15).
It is curious to note the difference in action required for the
coupling (jf the wings in the two sexes, owing to the difference
in position of the retinaculum. In the female, we must move
the hindwiny sharply forward, and then let it fall gently back,
so that the frenulum may become engaged in the catching hairs.
BY R. J. TILLYARD. 307
But it is necessary to move the foreiving sharply forward, if one
desires to secure the coupling of the two wings in the male. In
doing this, the stationary frenulum is caught up by the moving
hasp-like retinaculum; and the grip, once effected, is not easily
lost.
AVe may sunnnarise these results briefly as follows: —
Fetnale. — Unspecialisedyr('>i///?/?/iof from two to nine separate
bnstles set closely together. Unspecialised retinacubim of stiff
hairs or scales directed anteriorly, and developed from tlie under-
side of the cubitus of the forewing.
Male. — Specialised frciiuluiit of a single, large, composite
bristle, formed by fusion of the original series of separate bristles.
Specialised retinacidnrn in the form of a chitinous hasp, directed
posteriorly, and developed from the underside of the radius.
B. Amplexiform Types.
The groups which haxe lost tlie frenulum and adopted the
amplexAfomt type of wing-coupling (in which connection is main-
tained simply by overlapping of the anal area of the forewing
upon the well developed costal area of the hind, including the
enlarged humeral lobe) may be arranged in three super-families;
viz., the Saturniina, the Lasiocampina, and the Papilionina or
Rhopalocera. These three do not appear to be very closely
related.
The Saturniina comprise the large and highly specialised group
of the Atlas Moths or Emperors. These possess neither frenulum
nor proboscis, and have a highly specialised venation. Meyrick
places them as the last of his series Notodontina ; but this
arrangement does not seem to be generally accepted by other
Lepidopterists.
The Lasiocampina form a natural group, if the E upterotidoi
and DrepauidcB be included in them. In the former family, a
frenulum is present; in the latter, most of the genera show a
small or reduced frenulum. In the other two families, the
Bndromidie and Lasiocampidce, the frenulum is absent. These
last show a very high specialisation of the amplexiform arrange-
;enera. For instance, in Lasiocampja, the
308
THE PAXOKPOID COMPLEX, 1.
humeral lobe is greatly enlarged, and is supported by a series of
recently funned ribs or cross-veins.
The Papiliunina furni another natural grouj), wliose origin
appears to be nuich in douljt. Meyrick and Hainpson consider
them to be derived from the Thyrldidiu. The Ilf^jjcriidai^ ho\\ -
ever, grade so closely into the Castniidai in Australia, ^Yhere
forms like Eusckumou are of Hesperiid structure in all but the
possession of the frenulum, that it is ditlicult to cast aside the
belief, held by the opposite school of Lepidopterists, that the
Co^fitluhe closely approximate to the ancestral form from which
the higher ButterHies, through the I/esjM'riid(e, ha\e been derived.
C. The Family Castmid.e.
j\[uch light can be thrown upon the evolution of the two types
of frenulum, in the male and female of Heteroneura respectively,
as well as upon the origin of Amplexitorm types, by the study of
Text -% 1(5.
\\'iH^-ci)upliU!4 appaialus of Sz/uemon hesperoults Feld., ,5 , vieAved from
beneath; ( X 14) : y'/-, frenulum; Fir, forewing; H>i\ liindwing; /•/, re-
tinaculum developed from radius (R); rf\ retinaculum developed from
cubitus iCn)', in 9 , this latter alone occurs.
the family Castaiidtf.^ In this family, both the humeral lobe
and tlie frenular bristles may be seen well developed, in such a
genus as >Si/7iem(m (Text-fig. 16). In the males of this genus, the
* For the supply of material for study in this famih% I am indeljted to
Mr. Geo. Lyell, F.E.8., of Gisborne, Vic.
t3Y R. J. TiLLYAftD. 30<)
frenulum is of the usual type, i.e., elongated, and with all the
bristles fused together into a single, strong bristle. In the
female, there are two long and delicate l^ristles (as in Pln^ia)
accompanied usually by several sliorter bristles at their base.
The retinaculum of the female is (jf the usual type, consisting of
a patch of forwardly projecting scales, developed on the under-
side of the cubitus of the forewing; but this retinaculum is situ-
ated so far from the wing-base, that only the two longer bristles
can become engaged in it, and tlie connection appears to be easily
lost during Hight In the male, this retinaculum of scales upon
the cubitus is refautrd ([ii'ifr cott.spicxoftslt/ {Text-^'^AG, rt'), to-
gether with an additional development of a di(/ht ovrr/o/diur/ of
the edge of the radius in a backward direction {rl). Usually,
during Hight, the frenulum of the male is caught in the curve of
this overfold, as sliown in the figure. But if, at any tijne, the
frenulum slips out of this somewhat precarious hold, it is then
caught up by the still existing retinaculum on the cubitus, and
the insect can continue its Hight in the manner of the female.
Thus we see that, in St/tientoii, the type of coupling-apparatus
for the wings is much closer, in the two sexes, than it is in other
frenate moths; and we can scarcely doubt that the condition in
tlie male of this genus is that which preceded, in the males of
other frenate moths, the more specialised condition that we tind
in them at present.
If now, we turn to the genus Enschemon^ which is claimed by
some authors as a moth, by others as a buttertly, we Hnd that
the male has a well-developed frenulum and retinaculum of the
type found in other frenate moths, i.f., more highly specialised
than in Syufinon. But, on examining the female, I was surprised
to find no trace whatever of a frenulum or retinaculum; and it
it is evident that this sex couples its wings in the amplexiform
maimer of the Butterflies, by means of its large humeral lobe.
Thus, judged only on the form of its wing-coupling apparatus,
the male of Euscheinon is a frenate moth, the female a butterfly.
This genus, then, exactly bridges the gap between the old sub-
divisions Heterocera and Khopalocera. That this is not a mere
310 THE PANORPOID COMPLEX, i.,
chance convergence, I shall hope to prove in detail later on, when
we come to consider the question of kirxal forms; the larva of
Enschr}}io)i being, in almost exery respect, that of a typical
Hesperiid.
From the above evidence, the origin of the Amplexiform type
of wing-coupling in the Butterflies is naturally suggested as fol-
lows. The ancestors of the Butterflies must have had a wing-
coupling apparatus of the type seen in iSt/nrnuni at the present
day. By developing a manner of flight differing from that of
the moths, principally in the greater amplitude of wing-stroke
but lesser number of vibrations per second, the delicate, elong-
ated bristles of the frenulum in the female first began to fail to
hold the wings in position. The retinaculum being already too
far out for the shorter bristles to be of any use, the whole
frenular apparatus would then rapidly become aborted, and the
insect would depend upon the pressure exerted by the large
humeral lobe of the hindwing for keeping the wings in position.
That is to say, the Amplexiform type of wing-coupling first
became adopted in the female only, as we see it in Eui<che))Lon
at the present day. Next, considering the question of the male,
evolution could still proceed along one of two directions: either
the overfolding of the radius might continue to increase, until
an efficacious, hasp-shaped retinaculum (of the type seen in most
male f renate moths) had been developed, thus saxing the stronger
frenulum of this sex from extinction; or the line of evolution
might follow that of the female, presumably by the change in
the method of Might making it impossible for even this strong
frenulum to continue its hold upon the retinaculum, at a time
AS hen the latter had not yet i-eached the hasp-shaped stage. In
that case, frenulum and retinaculum would, in the end, become
aborted, as in the female, and we should arrive at the full Am-
plexiform type exhibited by the Buttertlies in both sexes.
BY R. J. TILLYARD.
311
The following Table will exhibit, in a concise manner, the
differences in the c(jndition of the wing-coupling apparatus studied
above : —
Group
Sex
Frenulum
Retinaculum
on cubitus
Ketinaculum
on radius
Typical
Heteroneura
V
9
6
short unfused
bristles
sin;,de lon^' bris-
tle (original
bristles fused)
present
absent
absent
present
Ca.
duiid't' : —
'Syiitmon
9
two long and
several short
biistles, unfused
present
absent
Syiitmon
6
single long biis-
tle (original
bristles fused)
present
present,
weakly
formed
bJuschemou
9
absent
absent
absent
Euschtmou
6
sin<j;le long bris-
tle (ori<,'inal
bristles fused)
absent
present, as a
strong hasp
Papilionina ^
iSaturniina* ,
Lasiocampiiia" (par- '
tim)
9 6
absent
absent
absent
Form of the Colpling-appakatus in the Archetype of the
Lepidoptera.
Reviewing all the above evidence, it becomes clear that the
oldest existing form of coupling-apparatus, viz., that of the
Micro2)terygidce, is already specialised in a direction from which
* It is not suggested here that the evolution of the amplexiform type in
the iSaturniina and Lasiocanipina has been along the same lines as that of
the Papilionina. In the Lasiocanipina, at any rate, a studj'^ of the con-
ditions to be seen in the Drtpanido' would suggest a direct descent from
Eupterotid-like ancestors with the frenate wing-coupling apparatus fully
developed. Thus the Papilionina are probably of remoter origin than
either the Saturniina or the higher families of the Lasiocanipina.
312 l^Hli PAXOHfOtD COMPLEX, i.,
the true Jugate and Freriate types cannot possibly be deii\-ed,
because of the alteration of the direction of the frenular bristles,
and the turning-under of the jugal lobe. We must conclude,
therefore, that tlie Archetype of the Lepidoptera possessed a
primitive jugofrenate type of coupling-apparatus, of the form
found in the Planipennia; i.'\, with all parts present and normal,
except the jugal bristles, which were absent.
General Conclusions.
Reviewing the evidence afforded by all the Orders of the
Panorpoid Complex, we may legitimately arrive at the following
conclusions : —
(1) The original coupling-apparatus was situated at the bases
of the wings, and was of a primitive jnyo-frenate type, in which
each wing developed a slight lobe towards the other, and these
lobes bore projecting bristles which came into contact, and so set
up a tactile connection between the two wings, to help in the
coordination of the act of Might. By increase in the size of the
lobes, or the length of the bristles, the connection became of a
more intimate kind, a certain amount of pressure being exerted
to keep the wings in position during flight.
In this primitive type, there were originally four elements
represented, viz., the j/iy a/ lobe with its juyal bristleti on the foie-
wing, and the hnuural lobe with ita freyiulum on the hind.
(2) The only Order which has retained this ancient type of
coupling-apparatus in its entirety is the Mecoptera; and, in this
Order, the apparatus is only fully functional in the two archaic
families Churisfiche and San ttochoriaf idee.
(3) By loss of the jugal bristles, and by increase of the size
of the two lobes, there arose the modification of the jugo-frenate
type seen in the Planipennia. This also is only fully functional
in the older families, such as the Ilfnurobiida'.
(4) By loss of the frenular bristles, reduction of the
humeral lobe, and increase in the size of the jugal lobe, there
arose the series of types seen in the Megaloptera, Trichoptera,
and the true J ugatc Lepidoptera {IL'jjLalldce and Frotothcoridoi).
BY R. J. TILLYARD. 313
These parallel reductions must be coiiceiYed of as haYing taken
place quite apart from one another phyletically, and probably at
quite different geological periods. Though passing through the
same series of reductions, these three groups arrive at quite
different final results, as may be seen by comparing llapliidia
(Text-fig.7) with Opcrfh (Text-fig.9) and witli Characiia (Text-
fig. 11).
(5) The highest specialisation of the old jugo-frenate type, still
preserYing the original characters of that type, is the jugo-frenate
type of the Microjyfprygidcp, in which the jugal lobe is folded
under the forewing, so as to become directed forward and out-
ward, and the frenulum becomes engaged in the groove so formed.
Thus the jugal lobe functions as tlip most archaic form of rctina-
cnlnm yet discorcml.
(6) The origin of the specialised jugum of the HejnalidcE and
Protothcoridct is not to be traced dii-ectly from the Micropterygid
type, but fi'om an older, unspecialisefl, jugo-frenate type, such as
we find in the Planipennia. The jugal lobe remains projecting
outwards and downwards, not turned forwards to pass undei' the
forewing. At first, it rested upon the ct)sta of the hindwing, as
in the older forms of Tiichoptera. The frenulum, being useless,
disappeared. A later and higher specialisation led to the length-
ening and narrowing of the jugum, and finally to its adopting
the position seen in most of the Hej^iaUdce.
(7) From this, it follows that no existing type within the
Lepidoptera of to-day represents the true ancestral form or
Archetype of the Ordei\ That Archetype must have possessed
an iiiisppcifdiKcd, jayo-frpuatc coapJuai-apparat^is^ probably with-
out jugal bristles, as in the Planipennia to-day. The de\elop-
ment of the specialised jugo-frenate type of the M icropterycfidix,
on the one hand, and of the specialised jayatc type of the Hcpi-
alidce and Frotothcoridit on the other, must have pi'oceeded upon
diYergent lines.
(8) The frenate forms amongst the Lepidoptera are also de-
rived from the archetypic jugo-frenate foiin, and not directly
from the Microptn-yyidce, as is pro^•ed by the bristles of the
frenulum maintaininti- their original direction. This line was
314 THE PANORPOID COMPLEX, i.,
evolved simply by loss of the archaic ju^al lobe, with correlated
reduction in size and strength of the hindwing, as it became
more and more dependent upon the fore in fliglit.
The most ancient type of frenate coupling is that preserved
for us in the females of the whole series, where the bristles
remain short and unspecialised, and the retinaculum is formed
simply from the brush of stiff hairs or scales, that project for-
wards from the underside of the cubitus on the foi-ewing.
In the males, there is a higher degree of specialisation, the
frenular bristles becoming fused together and greatly lengthened,
while a new and more effective retinaculum is developed from
the underside of the radius in the forewing, in the form of a
posteriorly projecting hasp or catch of chitin.
(9) The amplexiform types amongst the Lepidoptera Hetero-
neura are to be regarded as a series of separate developments
(probably three in number) from originally frenate ancestral
forms. Of these, the most evident connection would appear to
be that uniting the frenate Ca^tiiiid(p, with their well developed
humeral lobe and clubbed antennae, with the very similar but
non-frenate IJpsiwriidfP.
(10) The only portion of the original coupling-apparatus left
in the highly specialised Order Diptera is the jugal lobe, which
becomes enlarged to form the aJula.
We may now exhibit, in the form of a short Table, the state
of the coupling-apparatus in the various existing groups, to which
we must add the Archetypes of the several Orders, as these
results will be required in the final discussion on the Phylogeny
of the Orders. (See pp. 3 16-3 17).
BY R. J. TILLYARD. 315
8uGf4ESTED CLASSIFICATION FOR THK LePIDOPTERA.
The characters here studied, thongli not suitable for the main
subdivision of the Order into two Suborders, may be legitimately
used for the subdivision of the first of the two Suborders which
we have based upon the wing-venation, viz., the Homoneura.
They cannot, however, be used in subdividing the Heter<jneura,
since the three Amplexiform groups do not lie along one single
line of descent.
We may, therefore, adopt the following classfication : —
Order LEPIDOPTERA.
1. Suborder HOMONEURA.
Venation of fore- and hind wings closely similar and of primi-
tive design.
Division A. Jugo-frenata. — With archaic jugo-frenate coup-
ling-apparatus consisting of jugal lobe, humeral lobe, and frenu-
lum; the jugal lobe turned under the forewing, and acting as a
retinaculum for the forewing Family Microjytpvyyidcp (s.lat.).
Division B. Jugata. — With specialised jugate coupling-appa-
ratus; the frenulum absent, the jugal lobe elongated and nar-
rowed, usually passing beneath the hindwing
Families HepiaJidce and Frototheoridce.
•2. Suborder HETERONEURA.
Venation of fore and hindwings dissimilar, that of the hind-
wing being strongly reduced in comparison with that of the
forewing. Coupling-apparatus of frenate type, except in three
of the highest groups, in which the frenulum is absent and the
coupling is of the amplexiform type.
(Division of the immense number of forms into superfamilies
and families must be based upon wing-v^enational and other
characters).
316
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318 the panorpoid complex, i.,
Appendix.
Note on the habits of flight and resting position of Sahatinca.
The following note, communicated to me in January last by
Mr. A. Philpott, of Invercargill, N.Z., throws some new light
upon the habits of this archaic genus: — "Since returning home,
I have been watching for the new species of Sahatinca in a little
bit of bush near my house. The weather is still broken, but on
sunny days I have been able to learn a little about this species.
I have seen several, always on or near certain mossy logs. The
moth sits with its wings held roof-wise over the body. The head
and thorax are held high up, as if ready to jump. When taking
flight, the insect seems to spring into the air with closed wings,
and then to fly forwardi5 in a rather feeble and fluttering manner.
They do not fly far; five or six feet is a rather long distance for
them. The antennae are always held widely separated, pointing
obliquely forward and upward. I am keeping some in a jar with
moss, in an endeavour to get some eggs and larvee, but with no
result so far. When touched, these captives spring an inch or
two sideways, backwards, or forwards with equal ease. The
feeble fluttering flight of this species, and I suppose it to be char-
acteristic of the genus, is in marked contrast to the strong,
dashing, swift motion of the Hepialidce. The jugal lobe is folded
back as in S. inconyruella\ the frenulum is similar to your
drawing.
BIBLIOGRAPHY.
(Of Nos.1-7, already quoted in the Introduction to this work, only Nos.
1, 3, and 4 are referred to in Part i. ).
1. CoMSTOCK, J. H., 1893. — Evolution and Taxonomj^ in " The Wilder
Quarter Centenary Book." Ithaca, N,Y., 1893, pp.37-114.
3. Meyeick, E., 1895. — Handbook of British Lepidoptera. London,
Macmillan and Co., 1895.
i. , 1912. — Micropterygidm in "Genera Insectorum," Fasc.
132. Brussels, M. P. Wytsman, 1912.
8. Forbes, W. T. M., 1914.— "The North American Families of Lepi-
doptera." Psyche, 1914, xxi., No.2, pp.53-65.
BY R. J. TILLYARD. 319
9. Meyrick, E., 1917. — "Descriptions of South African Lepidoptera, "
Ann. S. African Mus., 1917, xvii., Pt. i., No.l, pp. 1-19. (Proto-
theoridce, pp, 17-19).
10. Packard, A. S., 1895. — "Monograph of the Bombycine Moths of
North America." Mem, Nat. Acad. Sci., Washington, 1895, vii.,
No.l, pp. 5-390. (8eep.62).
11. , 1898. Textbook of Entomology. New York. The
Macmillan Co., 1898. (See p. 123).
12. TiLLYARD, R. J. 1916.—" Studies in Australian Neuroptera, No. iv."
Proe. Linn. Soc. N. S. Wales, 1916, xli., Pt. ii., pp.269-332. (See
p. 286, and PI. xiv., fig, 19).
13. , 1917.—" The Wing- Venation of Lepidoptera." (Pre-
liminary Report), I.e., 1917. xlii., Pt i., pp. 167-174.
EXPLANATION OF PLATES XXLX.-XXX.
Plate xxix.
Fig. 1, — Jugum of MicropUryx aruucel/a Scopoli, viewed from beneath ;
( X 150).
Fig. 2. — Frenulum of same, viewed from beneath; ( x 125).
Fig. 3. — Humeral lobe and bases of insertion of frenular bristles in Saba-
tinea inconr/ruel/a (Walker); ( x 250).
Fig. 4. — Frenulum of Micropteryx thunhergeUa Fabr. ; ( x 125).
(Photomicrographs from descaled, cleared and mounted specimens.
The jugal lobe in Fig.l, and the humeral lobes in Figs. 2 and 4, have
become cracked across near their bases, owing to the pressure of the cover-
glass upon old and brittle material. Owing to the same cause, the frenular
bristles have become pressed down upon the hindwing, and their correct
directions somewhat altered).
Plate XXX.
Fig. 5. — Jugum of Charagia eximia Scott, viewed from beneath; ( x 8).
Fig.6. — Frenulum and retinaculum of Cephenodef^ janus janu.^ Miskin, 9 ,
viewed from beneath; ( x 5).
Fig. 7. — Frenulum and retinaculum of Hippotion scrofa Boisd., 9 , viewed
from beneath; ( x 7).
Fig. 8. —Frenulum and retinaculum of Hippotion scrofa Boisd., <J , viewed
from beneath; ( x 6).
(Photomicrographs from set specimens, not descaled. In Fig. 7, the
black bristles of the frenulum appear white, owing to the incidence of
strong artificial light directly upon them).
320
THE GEOLOGY AND PETROLOGY OF THE GREAT
SERPENTINE BELT OF NEW SOUTH WALES.
Part vii. The Geology of the Loomberah District and a
PORTION OF the GoONOO GoONOO EsTATE.
By W. N. Benson, B.A., D.Sc, F.G.S., Professor of Geology
AND Mineralogy in the University of Otago, N.Z., late
LiNNEAN MaCLEAY FeLLOW OF THE SOCIETY IN GeOLOGY.
With two PALiEONTOLOGicAL Appendices by F. Chapman,
F.R.M.S., A.L.S., Palaeontologist to the National
Museum, Melbourne.
(Plates xxxi.-xxxviii., and five Text-figures).
contents. page
Introduction AND Acknowledgments 320
Previous Literature 322
Physiography 323
General Geology: — Comparison of Stratigraphical Succession in
Tam worth, Loomberah, and Nundle Districts 324
Regional Geology —
Devonian Sedimentary Rocks ... ... ... ... 327-341
Devonian Igneous Rocks ... ... ... ... ... 341-345
Igneous Rocks of the Western Series 345-352
General Summary of the Devonian Stratigraphical Succession ... 352
Petrology 363
Bibliography 382
Introduction and Acknowledgments.
In previous papers of this series, the writer has described and
given geological maps of the Districts of Nundle and Tamworth
(1, 3, 5, and 6). The former was the first studied, and broad sub-
divisions only were made in the stratigraphical succession. In
the latter, every zone of rock of any importance was mapped, in
the endeavour to obtain as detailed a knowledge as possible of
feY W. N. BENSON. 32i
the exact sequence of formations and tectonic structures devel-
oped. The present communication serves a twofold purpose
It completes the preliminary mapping of the serpentines of the
main line of intrusions from Warialda to Hanging Rock, a dis-
tance of over 130 miles; and secondly, it carries the detailed
tracing of the horizons determined in the Tamworth District
through into the Nundle District, permitting a more accurate
correlation of the several formations in the two districts than
was formerly possible, and thereby indicating that a much
greater complexity exists in the Nundle region than was assumed.
Several features of interest occur in this district, which are
but barely represented beyond it. Firstly, there is the occur-
rence of the Loomberah Limestone, which runs throughout the
length of the district, and extends into the Nundle region. It
is an horizon absolutely distinct from that of the Nemingha or
Moore Creek Limestones, with a different lithology, and a different
and interesting fauna characterised by the presence of numerous
pentaraeroid shells, and a group of corals which do not occur in
the limestones of the two other horizons. In this, and in the
Nemingha limestones below are several species of Tryplasma^
hitherto very rarely observed in the Devonian rocks of this
State,* while, in the latter, is a very rare feature, namely, the
occurrence of Foraminifera in Devonian rocks.
Apart from the faunal peculiarities of this region, the devel-
opment of igneous rocks is of interest; there are here important
■'■■ Other recorded occurrences of Tryplasma in Devonian rocks in New
South Wales are as folloMs : —
1. An undescribed species found by Mr. Surveyor Campbell in the Parish
of Cuerindi, a few miles south-east of Manilla (Ann. Rep. Dept. of Mines,
1915, p. 189).
2. T. lonsdalei var. 8calariformis, found by Cullen and the writer in
Nemingha Parish (This Series, Part v. Set Bibliography).
3. An indefinite species in the Crawney Limestone, collected by Cullen
(Mem. Geol. Surv. N S. Wales, No. 13, p. 101).
The second of these belongs to the horizon of the Nemingha Limestone.
The first and third are almost certainly on the Moore Creek Limestone
horizon. So that it has now been ascertained that Tryplasma may occur
in all three Middle Devonian Coral-limestones.
322 GREAT SERPENtlNE BELT OP NEW SOUTH WALES, vii.,
masses of keratophyre, which extend into the Nundle District,
but are barely represented in the Tamworth District. Several
varieties of dolerites, not yet seen elsewhere in the Devonian
rocks of New South Wales, appear in this area. Again, we
have the most typical instance of highly altered pillow-lavas yet
discovered in the Serpentine Belt. The serpentine, too, is by
no means as continuous in this region as elsewhere, but is
broken into small, isolated lenticles, and the line of separation
of the Eastern and \^'estern series becomes obscure. Finally,
we may note the occurrence of peculiar tectonic features in this
region, developed to a greater degree than in adjacent areas.
The writer is much indebted to Mr. G. McA. King, the
General Manager of the Peel River Company, for permission to
enter and study the geology of portion of the estate of that Com-
pany, and to numerous farmers in the Parish of Loomberah for
similar kindness. In particular, he is indebted to Mr. Ponto,
of Pendene, for kind hospitality, and to Mr. David Carter for
help in securing the rich collection of fossils obtained from his
property. This was supplemented by specimens from the collec-
tion of Mr. S. M. Tout, who was the first to bring the Loom-
berah limestones under scientific notice.
Mr. Etheridge has very kindly examined the Tryplasmas re-
corded herein, and Mr. F. Chapman, the Chtetetesand Foramin-
ifera. To Mr. Dun, the writer is indebted, not only for the
determination of all the other fossils, aiid the valuable notes
included in this paper, but also for his constant interest in the
work, and helpful discussion of the stratigraphic problems that
arose. To the chemists of the Geological Survey are due the
analyses of the limestones cited on pp.334 and 338, which were
obtained from specimens collected by the writer. To all these
gentlemen, his best thanks are due. Mr. Mingaye's very com-
plete rock-analyses, cited on p. 368, add great value to the petro-
logical studies.
Previous Literature.
The references to this region by previous writers have been
very scanty. It was visited b}^ the Rev. W. B. Clarke in 1853,
and the occurrence of Phillipsastrcea on Cope's Creek was ob-
BY W. N. BENSON. 323
served (8). Professor De Koninck determined this as P. ver-
neuilii{9). The duties of Mr. F. Odernheimer, who made a private
survey of the land belonging to the Peel River Land and Mineral
Company, must have involved the investigation of this region,
but no special attention is directed to it in his paper "On the
Geology of a part of the Peel River District in Australia," pub-
lished in 1855(10). Stonier mentioned the occurrence of serpen-
tine near Black Jack, 1892(11). The present writer described
the southern portion of this district in 1911(1), and 1913(3) in
his papers on the Nundle District, gave petrological details of
the occurrence of keratophyres on Silver Gully, and the pillow-
lavas near the Peel River in 1915(5), and included the northern
edge of the district in the area covered by his paper on the Tarn-
worth District, later in the same year (6). In this paper, the
position of the Loomberah Limestone, as a definite horizon dis-
tinct from the Nemingha and Moore Creek Limestones, was first
remarked.
Preliminary notes on the fossils collected from the limestones
of this district by Mr. S. M. Tout and the writer were made by
Mr. Dun (12); and a number of analyses of specimens of this
limestone were given by the chemists of the Geological Survey
(13). Dr. Jensen cites the results of some investigations not yet
published in detail, which were made by Mr. Guthrie, on the
nature of the soils in the regions of the limestones in this
district(14).
Beyond these, no references to the district in geological liter-
ature are known to the writer.
Physiography.
The region lies for the most part west of the Peel Ptiver,
stretches from the south of the Parishes of Nemingha and Tam-
worth, and includes all the Parish of Loomberah and a large
region comprising the north-eastern corner of the Peel River
Company's estate. Its eastern margin is, in fact, marked by
the deep meandering valley of the Peel River, an underfit stream
with broad alluvial flats between spurs, sometimes sharpened,
but more generally blunted (cf. 15 j. To the east of this, the land
324 GREAT SERPENTINE BELT OF NEW SOUTH WALES, vii.,
rises up into a peneplain-surface of about 3,500 feet in height,
cut in hard rocks, chiefly siliceous phyllites, and very much dis-
sected by deep valleys, some following the structure of the
country, others oblique thereto. West of the river, the topo-
graphy is more complex. The geological formations are arranged
in parallel zones of resistant and less resistant rocks, so that
there is a succession of ridges, the ridge of the Eastern series
directly west of the Peel River, running north from Wallaby
Mountain by Woolomin, composed chiefly of siliceous phyllites
and jasper, the ridge of tuffs associated with the Loomberah
limestone, and the ridge of tuffs and agglomerates which form
the Pyramid Hill Range. To the south, a fourth ridge of con-
glomerate intervenes between the last two. These ridges are
separated by the open valleys of Sandy Creek and Reedy Creek
respectively. To the south of these creeks, there is a series of
more youthful streams, which cut athwart the grain of the country,
and flow generally north easterly or east-north-easterly into the
Peel River. Of these. Cope's Creek is the most important. The
lower portion of the creek is thus roughly parallel to the series
of dip-faults (see p. 360) which cut through the country and may
originally have been determined b}^ one of these. Pipeclay
Creek, Silver Gully, and Hyde's Creek belong to the same trans-
verse system of streams. The same holds even more clearly for
the creek north and west of Wallaby Mountain.
General Geology.
To understand the significance of the features observed in
this district, those to the south and north must be reviewed.
Along a line of section drawn westwards from the serpentine
near Bowling Alley Point, the following sequence was observed
{see 3): —
i. The Lower Banded Radiolarian Claystones.
ii. The Lower Bowling Alley Tuffs and Breccias, in which
there are numerous intrusions of dolerite and spilite.
iii. The Limestone.
iv. The Upper Banded Radiolarian Claystone, interstratified
with a large amount of tuff and breccia, and abundant intrusions
of spiiite and dolerite.
BY W. N. BENSON. 325
V. Upper Bowling Alley Tuffs and Breccias, with some cherts.
With these are two lenses of limestone considerably west of the
main zone.
vi. Mudstonesof the Nundle Series, gradually passing up from
the cherts associated with the Upper Breccias and including a
conglomeratic zone which forms Nundle Sugar Loaf, two miles
west of the township.
In putting forward this sequence, it was noted that there
seemed a possibility of considerable repetition occurring within
it. The distinction between the phyllites and jaspers east of
the serpentine (the Eastern or Woolomin Series), and the rocks
to the west of it, seemed very clear.
In the Tam worth District, however, further facts were dis-
covered. Parts of the Eastern Series were recognised as being
merely infolded and highly crushed equivalents of western rocks,
and the Western Series itself was subdivided as follows (6): —
(a) The Lower Middle Devonian series of radiolarian cherts
and claystones, with intercalated tuffs containing the Nemingha
Limestone, which is often closely associated with a very ferru-
ginous tuff or agglomerate.
(b) The Igneous Zone, consisting chiefly of breccias, agglomer-
ates, and tuffs with dolerites and spilites.
(c) An Upper Middle Devonian Series of radiolarian cherts
and claystones, containing, in its lower part, the Loomberah
limestone, and, in its middle or upper part, the Moore Creek
limestone. In the portion of this Upper Middle Devonian Series
which lies above the Moore Creek limestone (as, e.g., the clay-
shales of Tam worth Common), the cherty character of the sedi-
ments is entirely lost.
(d) The Baldwin Agglomerates, which form the base of the
Upper Devonian Series.
(e) The Barraba Mudstones, forming the Upper Devonian
Series proper. These are often difficult, indeed impossible, to
distinguish lithologically from the rocks of the upper portion of
the Upper Middle Devonian.
It was further shown that there are several repetitions of zones
(a) and (6) in the region lying immediately west of the Serpen-
326 GREAT SERPENTINE BKLT OF NEW SOUTH WALES, vii.,
tine. When, however, we attempt to trace these subdivisions
through the Loomberah District, and to apply them to the suc-
cession in the Nundle District, we are met with difficulties, for
the new factors entering into the geology of the Loomberah
District add to the complexity of the problem. Firstly, there
is a general absence of serpentine throughout the region, so that
the boundary between the Eastern and Western Series is not a
distinct and unmistakable line of fault separating altered and
unaltered rocks, but merely a zone of transition, crushing, and
silicification. Secondly, even where the serpentine does occur,
the rocks immediately west of it have, in many cases, as highly
altered a character as those to the east. Tliirdly, there is de-
veloped a large amount of keratophyre, forming an intrusive
mass or masses, one of which apparently is just below the horizon
of the Nemingha limestones. The southern end of this zone of
keratophyre lies in the Hyde's Creek complex, and a further
occurrence of it extends from Silver Gully to Pipeclay Gully
(5, pp. 133-137). The northern end of the zone is that complex
near Macllveen's homestead in the south-eastern portion of Tam-
worth District(6, p. 572). With this keratophyre is often asso-
ciated a great amount of ferruginous and chalcedonic jasper
derived from magmatic solutions. Finally, there is to be con-
sidered the extremely disturbed and broken character of the
faulting, breaking-up the stratigraphical succession, so that it is
not possible to determine the extent of repetition of series that
is present.
To simplify the reading of the detailed discussion of the several
formations, it may be well to state, at the outset, that the facts
appear to justify the following conclusions concerning the strati-
graphical relationships of the rocks in the Tamworth, Loomberah,
and Nundle Districts.
L That tlie lower part of the Nundle Series does not corres-
pond with the Barraba Series, but to the upper part of the
Tamworth Series, namely, that lying between the Moore Creek
limestones and the Baldwin Agglomerates.
2. That the ridge of conglomerate forming Nundle Sugarloaf
west of that township, corresponds to the conglomerate which
BY W. N. BENSON. 327
runs along tlie eastern foot of the P\n-amid Hill Range (the
►Scrub Mountain conglomerate). This may be approximately
coeval with the Baldwin Agglomerates.
3. The mudstone west of these may be correlated with the
Goonoo Goonoo or Barraba mudstones.
4. That the great masses of agglomerate and tuff, being derived
from eruption at various centres, do not form continuous horizons
except in rare cases, and great care must be used in stratigraphi-
cal correlation based on the lithological features of pyroclast'ic
rocks. Hence arises the difficulty of determining the exact
horizon to which must be referred the Pyramid Hill group of
tuffs, etc., or of recognising definitely the horizon of the Baldwin
Agglomerates.
5. The Moore Creek Limestone is not represented in the
2^undle District, but its appropriate horizon is probably not far
from the township of Nundle itself.
6. TheLoomberah Limestone is represented by the two western
lenses of limestone near the head of Silver Gully, to which special
attention was called during the first survey of the Nundle
District(3, p.574).
7. The main line of limestone which passed through to iiowling
Alley Point and continues to Hanging Rock, belongs to the
Xemingha or Lower Middle Devonian Limestone.
iS. That the pyroclastic rocks which lie directly above the
spilites and dolerite of the Upper Bowling Alley Claystones
represent the " Igneous Zone'' of the Tamworth District.
9. The Lower Bowling Alley Tuff-Breccias, Dolerites, etc., are
a repetition of the same " Igneous Zone "' and its associated
intrusive rocks.
10. Much of the Middle Devonian (Tamworth) Series may bt'
recognised in an altered state in the regions east of the serpentine
in all three districts.
The Eastern Series.
In the Loomberah District, we will describe with the Eastern
Series, not only all those rocks lying east of a line joining all the
outcrops of serpentine, but the lithologically identical masses
which lie to the west of that line, the rocks of the Transitional
328 GRKAT SERPENTINE BELT OF NEW SOUTH WALES, vii.,
Zone (a name indicating a local feature only), distinguished by
their greater alteration, fracturing, and silicification, from the
normal breccias, tuffs, and cherts of the rest of the Tamworth
Series. The actual western margin of the Eastern Series is thus
difficult to define, and that laid down on the map (Plate xxxii.)
is not a sharply marked feature. There is clear evidence that
very much of this Series consists of merely highly altered forms
of similar types of rock such as occur in the Tamworth Series;
while there are few rocks, if any, so utterly distinct as to demand
their inclusion in a definitely older, possibly Lower Devonian
formation, (the suggested Woolomin Series, a term we have now
abandoned).
Most notable are the masses of tuff and breccia, which occur
in the hills east of the Peel River, between Nemingha Creek and
Dungowan, continuing those already noted in the north-eastern
portion of JSemingha Parish (6, p. 547). These cross the Peel
River in the neighbourhood of Dungowan, and strike through
the hills west of the river till they come to the Wallaby Mount-
ain and Cope's Creek. Some of these are very similar indeed
to the normal tuff-breccias of Middle Devonian age, while others,
notably a great mass of almost schistose, highly crushed rock
running across the western end of Wallaby Mountain, consist
of a coarse tuff with abundant inclusions of limestone drawn
out into long lenticles. This may represent the Nemingha
limestone, but there are also instances where such a tuff is asso-
ciated with the Loomberah limestone. Some of the masses of
tuff are very silicified, and one instance of jasperised, ferruginous
agglomerate appears as if it might be the equivalent of the red
tuff of East Gap Hill, described in Part v. (pp. 564-6). This is
to be found on the hills east of the Dungowan Hotel, associated
with a little vesicular keratophyre.
Again, in Portions 52 and 146 in the south-eastern angle of
the Parish of Nemingha, three hundred yards east of the serpen-
tine, is a large belt of agglomerate, together with porphyritic or
granular spilite, inclosing a lenticular mass of limestone a
hundred yards long and twenty yards wide, with somewhat the
reddish crystalline character of the Nemingha marbles, associated
BY W. N. BENSON. 329
with ferruginous tuffs with limestone-fragments such as have
been already mentioned. Traces of a Stromatoporoid, of Favo-
sites multitabulata, Heliolites, and a large cyathopbylloid coral
were observed. These are insufficient to determine whether the
limestone belongs to the Nemingha or Loomberah horizon. It
runs due north, and is cut off to the south by a marked cross-
fault, which also truncates the adjacent serpentine. Limestone
again appears on the ridge in Portion 52, Loomberah, associated
with chert and spilite, all in a very shattered condition. This
lies in the transitional region between the Serpentine Line and
the Tamworth Series, but probably represents an altered form
of the Nemingha Zone.
The great masses of jaspers, which are such a marked feature
of the Eastern Series in the Nundle District, are especially well
developed in the Wallaby Mountain, west of Woolomin, where
a strong band strikes across the mountain, forming the long
ridge which deflects the outlet of Cope's Creek, rising into the
high cliffs that form the eastern side of the mountain, and dying
out in the valley to the north. Other jasper-belts occur east of
this, nearer to Woolomin, and further long bands traverse the
Parish of Woolomin, and others occur east of Dungowan, though
the last are not so extensive as elsewhere. They result from
intense silicification along zones of shattering, and are not
primary deposits.
The igneous rocks, other than tuff, namely the dolerites,
spilites, and keratophyres, occurring east of the serpentine, and
especially in the transitional zone west of the serpentine, are dis-
cussed below (p.342).
The Tamworth Series.
The region occupied by rocks of this division forms the greater
portion of the area considered, and, throughout this area, the
dips are nearly vertical or inclined to the W.S.W. at angles of
about 60°-70°, with only here and there a narrow band dipping
in the opposite direction. The total width of this region is
over four miles, so that, were no repetition of strata present, a
thickness of nearly 20,000 feet must occur, which is several times
greater than was shown to be probable in the Tamworth Dis-
330 GREAT SERPENTINE BELT OF NEW SOUTH WALES, vii ,
trict. We must conclude, therefore, that considerable repetition
by strike-faulting is present, though admitting that the sequence
from east to west is, in the main, an ascending one in the time-
scale.
We shall now endeavour to trace through from the Tamworth
to the Nundle District, several horizons already recognised in
the succession of Middle Devonian strata.
The Nemingha Limestone. — The main limestone of the Loom-
berah District is clearly a continuation of the mass of limestone
which runs through Beedles Freehold and the Gap at Nemingha
(6, pp. 587-9), and then commences by the eastern corner of
Portion 60,* Parish of Loomberah, appearing from beneath the
river-alluvium, and running thence to the south-east through
Portions 59, 58, and 57. It is a brecciated pink limestone asso-
ciated, in Portion 59, with a small amount of very ferruginous,
more or less brecciated keratophyre or porphyritic andesite, lying
on the eastern side of the limestone; and, to the west, is an intru-
sion of porphyritic dolerite with phenocrj^sts of labradorite. The
outcropping mass of limestone is 77 yards wide, but it is impos-
sible to determine the thickness of the limestone owing to the
absence of any evidence of the angle of dip, though adjacent
rocks dip at high angles. It is broken by cross-faults, throwing
it for short distances. In Portion 58 (Mr. David Carter's pro-
perty), the limestone has been duplicated by a fault or faults,
and a short band of the same horizon occurs four hundred yards
north-east of the main zone. The eastern mass is the more
clearly exposed, and covers an area four hundred yards long by
sixty yards in width. Both bands of limestone have a western
wall of keratophyre-breccias, ferruginous rocks apparently greatly
altered by percolating solutions (the Nemingha Red Breccia).
Both masses of limestone, for the most part, lie in ploughed
land, and, in the boulders of limestone brought to the surface, a
large series of fossils was obtained, which confirm the correla-
tion of this limestone with the Nemingha horizon. These were
* Unless otherwise stated, all references to Portion-numbers in the
sequel refer to Portions of the Parisli of Loomberah.
BY W. N, BENSON. 331
determined b}- Mr. W. S. Dun, Mr. Etheridge, and Mr. F.
Chapman. The following forms were recognised : —
Protozoa.
Foraminifera —
I'samiinosphcera neminghensis, sp.nov\, Chapman.
Valvulina plicata Brady.
V. hulloides Brady.
V. oblonga, sp.nov., Chapman,
Piilvinulina bensoni, sp.nov., Chapman.
Radiolaria —
An indeterminate species.
CCELENTERATA.
Zaphrentis sp.
Tryplasma congregationis Eth.fil.
T. lonsdalei Eth.fil.
T. vermifoj'mis Eth.fil.
A cystiphylloid coral.
Syringopora porteri Eth.fil.
S. auloporoides Eth.fil.
Favosites gothlandica Lamarck.
F. multitahulata Eth.fil.
F. pittmani Eth.fil.
F. crumrtieri Eth.fil.
Alveolites sp.
Heliolites porosa Goldf .
//. infersfincta Linn.
An indeterminate Stromatoporoid.
ECHINODERMATA.
Numerous Crinoid ossicles.
Brachiopoda.
' A try pa sp.
Zygospira sp.
Concerning the Tryplasma lonsdalei, Mr. Dun remarks that
it is apparently a solitary form of the T. lo7isdalei-t ype, but the
tabulae are more irregularly and widely spaced than in the
normal form. The wall is thick, and there are short, septal
spines. The diameter of the corallite is 5 mm.
332 GREAT SERPENTINE BELT OP NEW SOUTH WALES, vii,
Several interesting features are seen in this Table. The two
species of Syringopora named have, so far, been collected only from
the Moore Creek limestone, to which also F. crummeri has hereto-
fore been confined. The recognition of three definite species of
Tryplasma is an advance on the single form recorded previously
from this horizon (6, p. 552); while the occurrence of Foraminifera
is a very welcome feature, since, apart from forms occurring in
the Devonian limestones of Germany and South Devonshire, no
other occurrences of Devonian Foraminifera are known. They
were found in a small mass of oolitic limestone about four inches
in diameter. The granules, according to Mr. Chapman, form a
little more than half the bulk of the rock; "they vary in diameter
from 0-46 to 0*7 mm., and only a small proportion are due
entirely to oolitic accretion, whether originally of algal origin or
not it is impossible to say, on account of their present mineral-
ised condition. The nucleus of the oolitic grain, in more than
one case, was seen to consist of an ossicle of a crinoid. There
is a fair amount of iron-staining in the rock-structure which
seems to be entirely secondary, as the stain is de\ eloped more
strongly along incipient fracture-lines than in the grains them-
selves." (See Appendix i.). The ferruginous matter has doubt-
less been introduced from the associated NeminghaRed Breccias.
The analysis of the limestones from this outcrop are given
below (p.334).
Nearly a mile south of the occurrence in Portion 58, the lime-
stone appears again, associated with red, ferruginous keratophyre-
breccia crossing the western end of Portion 55; this outcrop is
nearly half a mile in length.
The next outcrop is considerably to the east as well as south
of the above, and is probably separated from it by considerable
dip-fault, though the intervening alluvium and ploughed land
hide the details of the outcrops. This mass is quite small,
occurring near the corner of Portions 53 and 52.
The limestone on Portion 35 has been moved still further to
the east, and is separated from the last-mentioned occurrence b}-
a large intrusion of dolerite. A small fault traverses this mass,
cutting it into two portions, one nearly 600 yards long and 60
BY VV. N. BENSON. 333
wide, the other 300 yards long and 40 yards wide. The chemical
composition of a specimen of this mass is given below (No. 1787).
A small mass of limestone next appears a mile to the south,
being found in a well sunk by the creek in Portion 32.
No more limestone appears for over a mile, until, in the
southern part of Portion 31, a great band of red breccia is asso-
ciated with a small amount of limestone on its eastern side.
The red band broadens as it continues southwards into the Peel
River Company's Estate, and the limestone now appears to the
west, as well as to the east, of the Red Breccia, which is here
mor.e than a quarter of a mile wide. An important mass of
limestone lies on the western side of Sandy Creek, half a mile
south of the boundary of Loomberah Parish. It consists of
three portions, largely of red encrinital limestone. The aggre-
gate length of the outcrop is 600 yards, and the width 80 yards.
The dip is apparently W.40°S. at 50°. The upper portion is
rather ferruginous; the lower, grey portion is either massive or
brecciated. {See analyses 1785, 1786, infra).
At the foot of Black Jack, the limestone and tuff is suddenly
cut off by a fault, which throws it half a mile to the east. On
the south side of the fault, the two masses of limestone, separated
by the breccia, are half a mile apart, but very soon they are
brought together again by a second great fault, and they continue
over the shoulder of Black Jack, making bold outcrops (analysis
1788), and pass thence down into Cope's Creek. The limestone
appears on the other side of the valley, and continues to Pipe-
clay Creek. This is the portion of the zone which was described
when considering the Geology of the Nundle District (3, p. 574).
In Cope's Creek, the limestone occurs in rather irregularly placed
masses in the red breccia, and one large lenticle has been torn
off and enveloped in keratophyre. It is possible that Clarke's
specimen of PhilUpsastrcea verneuilii was obtained here.
Throughout its length, this limestone keeps the characters
which mark the Nemingha limestone. It is usually thorouo^hlv
crystalline, so much so that, except for the fossiliferous masses
on Mr. Carter's property and south of Cope's Creek, no recc-
nisable fossils have been obtained. The rock is generally white7J • '^^V^
^"■♦■#
334
GREAT SERPENTINE BELT OF NEW SOUTH WALES, Vll.,
pink, reddish or mottled: less commonly, it is grey. It is often
highly brecciated, and in particular in Cope's Creek, the rock is
clearly made up of fragments of corals and of crinoids. The
colouration of the rock appears due to the introduction of ferru-
ginous solutions. The composition of the stone is shown in the
following Table; the analyses are due to the chemists of the
Geological Survey (13).
1781
1782
1783
1785
1786
1787
1788
CaCO,
97-46
98-21
97-68
94 03
97-54
97-55
98-21
MgCO^
0-7
0-69
0-64
0-90
0-71
0-79 1
0-50
MnCO^
0-04
0-06
0-04
0-08
0 05
0 04 1
0 04
Fe^Oa, AUOo ...
0-32
0-24
0-18
0-95
0-17
0-28
0 24
Gangue
1-56
0-98
1-42
4 02
1-52
1-48
1-24
100-09
100-18
99-96
99-98
99-99
100-14
100-23
1781-2 — Red Marble, Portion 58, Loomberah.
1783 — Pink Marble, Portion 58, Loomberah.
1785— Pink Marble, U miles N.N.W. of Black Jack Mountain.
1786— Red Marble, 1^ miles N.N.W. of Black Jack Mountain.
1787 — Pink Marble, Portion 35, Loomberah.
1788 — Pink Marble, east side of Black Jack Mountain.
Concerning the depth of origin of this limestone, we have to
note the absence of evidence of littoral conditions though the
limestone is often brecciated. Radiolarian chert is directly
associated with it. Mr. Chapman's remarks on this point may
be noted. {See Appendix i.).
The Loomberah Limestone.
A second definite zone is that of the Loomberah Limestone.
Just beyond the north-western corner of the Parish of Loom-
berah, namely, in Portion 10, Parish of Calala, is a belt of lime-
stone about a hundred yards long and ten to twelve feet thick,
containing HelioliUs, Tryiilasma., and crinoids. It dips W.5°S.
at 70'. To the east of it are several bands of tuff and a sill of
dolerite, the whole lying within normal radiolarian cherty clay-
stones. Followed to tlie south east, the limestone ceases, and
the tuff-breccia becomes ricli in fragments of limestone. It is
BY W. N. BENSON, 335
displaced by several cross-faults. In Mr. Carter's property
(Portion 58), the limestone comes in a»ain, forming a very im-
portant band. This is about a quarter of a mile in length and
150 feet thick. Its western end is very highly crystalline, so
much so that the rock has passed into a white marble, but,
towards the east, it is less crystalline, and consists of an extra-
ordinary assemblage of broken fossils. Fragments of large
pentameroid shells are the predominant feature, but intermingled
with them are abundant masses of Stromatnporoids, of fragments
of coral, and even angular or rounded pebbles of chert, the whole
giving very clear indication of rather turbulent conditions of
formation, as if they were reef-breccias (see Plate xxxiv., fig.l).
In spite of this, there is no indication of an unconformity, the
radiolarian cherts which lie immediately above the limestone
dipping W.25°S. at 70°, those immediately below W.26''S. at
65°, which, considering the variability of the dip, may be taken
as proof of conformity. Possibl}^ the shallow-water limestone
indicates the occurrence of a regression of the sea. In cases
such as this there is no reason to. expect a mass of coarsely
clastic sediment at the base of the overlying transgressive sedi-
ment {see 16, p. 458).
The second characteristic of this limestone is its remarkable
fauna, which is quite unlike that of any horizon yet discovered
in the Devonian rocks of New South Wales. The following list
indicates the forms recognised by Messrs. Chapman, Dun, and
Etheridge. Of particular interest is the presence of Tryplasma^
as determined by Mr. Etheridge, and Chretetes, described by Mr.
Chapman.*
Zaphrentis^ sp.ind.
Zaphrentis{^ sp. (Sp. et subgen. nov.]).
Tryplasma sp. Intermediate between T. delicatula Eth.fxl.,
and T. vermiformin Eth.fil.
Cyathophyllum sp.
* In a private communication, Mr. Chapman says — "It is interesting
to note your opinion of the hj^drographic conditions at Loomberah.
Ch(xtetes, to my mind, played the same part in the coastal reefs of the
Palteozoie, as HtUoporu does at the present day,"'
336 GREAT SERPENTINE BELT OF NEW SOUTH WALES, vii.,
Diphyphyllum, rohushim Eth.fil.
D. porteri Eth.fi].
Form allied to Omphyma.
Spoiigophyllum, sp.no v., Dun.
Phillipsastrcea grandis, sp.nov., Dun.
Favosites gothhindica Lamarck.
F. gothlandica with multitabulate habit.
F. multitahiitata Eth.fil. (rare).
Favosites, sp.nov. (?), a form with very irregular habit.
Favosites, sp.nov., a dendroid form distinct from those in
other Devonian horizons in this area.
Litophyllum konincki Eth.fil.
Dania sp.
Chcpietes stelliformis, sp.nov., Chapman.
Heliolites porosa Goldf.
Plasmopora sp.
Stromatoporella loomherensis, sp.nov., Dun.
S. be7isoni, sp.nov., Dun.
Actinost7'oma austraie, sp.nov., Dun.
A Treptostomatous Bryozoan.
Pentameriis [Conchidium) cf. kniuhtii.
Pentamerus cf. sieheri.
Strophomena cf. rugata.
A try pa sp.
Murchisotiia sp.
The band of tuff" or breccia, directly below this limestone, may
be traced south almost continuously to Cope's Creek, displaced
here and there by cross-faults. There is frequently a consider-
able amount of limestone-fragments in this tufi-breccia, but only
occasionally are there any large masses of limestone, such as
those in Portions 23 and 33 (the latter of which contains small
forms of Favosites multitahulata. The limestone also occurs in
Portion 32, and in several localities west of Black Jack. One
of these masses has a maximum width of sixty feet, dipping
nearly vertically , and contains pentameroid shells. Cyatho-
phyllum, Heliolites, a Stromatoporoid, globular masses of Favosites
gothlandica and Tryplasiuu(l) which characterise the horizon.
BY W. N. BENSON. 337
and, in addition, there was a section exposed of a form very
similar to Cystiphyllum vesicuiosum which unfortunately could
not be removed for more careful examination.
South of this, where the limestone is split into two bands bv
a zone of tuff, the feature noted in Portion d8, namely, the
abundance of fragments and pebbles of chert in the limestone, is
again present. It must also be mentioned that, throughout the
length of the zone of the Loomberah limestone, the chert over-
lying it contains numerous small inti-usions of tuflP.
The tuff associated with the limestone may be traced down
into the valley of Cope's Creek, and beyond it to the south.
Here, probably, was obtained the specimen of Phillipsastrcea
verneuilii recognised by De Koninck. The small patch of lime-
stone occurring on the top of the hill, a mile and a half W.S.W.
of the Permo-Carboniferous sandstone at ReicheFs homestead,
near Bowling Alley Point, probably belongs to this series. Jt
has been already pointed out that it does not lie on the same
horizon as the Nemingha limestone. . A lenticular mass of lime-
stone occurs near Cann's Plains Creek, still further to the south,
and probably belongs to the same horizon(3, p.574). Thus we
can trace the Loomberah horizon intermittently from the north-
west of the Parish of Loomberah, to within two miles of Bowling
Alley Point, and have thus obtained a second definite horizon
linking the geology of the Tam worth District to that of Nundle.
But there is still another mass of limestone that may possibly
belong to the same horizon, though so altered by recrystallisa-
tion in the region adjacent to the serpentine, that no traces of
fossils have been found in it. This is the lenticular mass of
white marble in the south-eastern corner of the Parish of
Nemingha, Portion 121, mentioned previously {see 6, p.560). It
is about four hundred yards long and sixty yards in width as a
maximum measurement, the main visible portion being only two
hundred by twenty yards in length and breadth.
338
GREAT SKRPENTIN^E BELT OV NEW SOUTH WALES, Vll.,
The chemical composition of the Loomberah limestone is seen
from the analyses below, which were made by the chemists of
the Geological Survey(13).
1395
1220
1784
CaCO,
98-85
94-91
95-43
MgCO,
0-42
0-83
1-27
MnCO,
0-04
Oil
FeoOs+AlaOg
0-22
0-80
0-60
Gangue
0-64
3-55
•
100 09
2-45
100-17
99-86
1395 — Limestone, Portion 121, Nemingha (horizon not certain).
1220 — Limestone, Portion 58, Loomberah.
1784 — Limestone, Portion 23, Loomberah.
Scrub Mountain Conglomerate.
For about a mile west of the zone of the Loomberah limestone,
the sediments are a succession of cherts, radiolarian claystones
and tuff, the latter sometimes broadening out into coarse agglo-
merates, as in the case of the agglomerate to the west of the
upper portion of Cope's Creek. In Reedy Creek, however, the
rocks lose their cherty nature to a great extent, becoming mud-
stones almost indistinguishable from those of the Barraba Series.
There is one very marked horizon in this portion of the section,
a zone of conglomerate which may be termed the Scrub Mountain
Conglomerate. It commences in Portion 17 and continues to
the south-east. It is made up of usually rounded or sub-angular
pebbles chieHy of a richly felspathic keratophyre, a strongly
trachytic type of which was found to contain phenocrysts of
felspar with abundant minute crystals of augite in the base(1474);
there is also some beautiful granophyre(1477, 1479), and, most
interesting of all, an angular fragment of radiolarian chert.
This last is not like the usual cherts of the Tarn worth Series,
being free from any sign of banding, but small amounts of a
rather similar rock do occur here and there in the Eastern Series.
All these lie in a tuffaceous matrix. This has not yet been
studied microscopically, but from a macroscopic examination
BY W. N. BENSON. 339
there did not appear to be the same close relationship between
the tuffaceous matrix and the inclusions (the former derived by
the attrition of the latter) that was so marked a feature of
Macllveen's complex, or in the Baldwin Agglomerates {see 6,
pp.573, 578). This, however, requires thorough investigation.
Should the matrix really prove to be clastic, not merely pyro-
clastic, the occurrence in it of the radiolarian chert and the
keratophyres, etc., might indicate another regression of the sea
at this point, and the exposure and erosion of Lower or Middle
Devonian rocks some distance east(?) of this region. The pos-
sible significance of this is considered later (pp.341, 355).
The zone is very continuous. It forms the long ridge in
Portions 17 and 18, and, southwards from thence, it makes up
the little hills which sharply mark the ends of the ridges that
separate the western tributaries of Reedy Creek. About four
miles south of the boundary of the Parish of Loomberah, it rises
as a long spur into Scrub Mountain, and south-east of here it
forms Rodney Mountain, beyond the limits of the map here-
with, and further to the south-east it rises into Nundle Sugar-
loaf, and crosses the Peel River to form the western end of the
Yellow Rock Hill south of that township. The last three occur-
rences of the conglomerate have been already noted (3, p. 581).
Thus the Conglomerate affords another horizon which can be
traced through from the Loomberah Parish into the region
mapped in the JSundle District. Small fragments of the con-
glomerate occur here and there adjacent to Reedy Creek; they
are perhaps due to repetition by schuppen-i'a,\x\tm^. Round
Mountain, which rises from beside the Main North Road on the
northern edge of the Goonoo Goonoo Estate, ten miles due south
of Tamworth, is apparently composed of this conglomerate.
The Pyramid Hill Tuff.
West of the Scrub Mountain Conglomerate and stratigraphi-
cally above it, the mudstones are interstratified with numerous
bands of tuffs of moderate grain-size, and these, being resistant
to erosion, form the long ridge which runs south-eastwards from
Pyramid Hill. This structure of the range is that of a faulted
340 GREAT S£Rf ENTINE BELT OF KKW SOtJTH WALES, vii.,
syncline. To the west of it there extends the broad, low, undu-
lating region composed chiefly of mudstones, extending to the
foothills of the ridges of Carboniferous rock that pass through
the Currabubula district. These are the mudstones of the
Burindi Series (on the extreme west), and the Barraba Series
which forms the greater portion of the Tamworth and Goonoo
Goonoo Plains. To the south-east, the tulf of the Pyramid Hill
Range apparently extends into the hills west of Nundle.
The Stratiyraphical Position of the NiLudle Mudstones.
The difficult question to be decided is the stratigraphical posi-
tion that must be assigned to the Pyramid Hill tuff. The syn-
clinal structure of the range, the lithological character of the
component rocks and the consequent topographic features, recall
the tuffs which form the hills west of the Tamworth Common.
These tuffs, etc., have been correlated with those occurring south
of the river in Portion 27, Calala(6, p. 580), and lie near, but not
on, the same line of strike as the Pyramid Hill tuff. The beds
which occur between the tuff of Portion 27, Calala, and the
nearest Loomberah limestone, are quite similar to those between
the Loomberah limestone and the Pyramid Hill tuffs, save for
the absence of the Scrub Mountain conglomerate. The first
impulse, therefore, would lead one to correlate the Pyramid Hill
tuff with the tuffs in the Tamworth Common, and, therefore,
with the Baldwin Agglomerate (6, p. 579). Such a correlation
would place all the mudstones of the Reedy Creek Valley,
together with the Nundle conglomerate, into the upper portion
of the Tamworth (Middle Devonian) Series, and would further
demand that none of the region mapped in the Nundle district
should belong to the Barraba Series, to which the western group
of mudstones (the Nundle Series) had previously been referred.
But the study of the tuffs and breccias within the Devonian
Series shows that they do not always form on a single horizon,
but (what might have been anticipated) they formed at different
times, spreading out on either side of their centres of eruption,
and, therefore, lie at varying horizons. Apparently there were
epochs when eruption was more continuous and widespread than
BY W. X. BENSON. 341
at other times, giving more or less definite igneous zones, l)ut
stratigraphica] correlation on tlie petrological features of pyro-
clastic rocks alone is exceedingly unsafe. It is quite possible
that the Pyramid Hill tuff belongs to a period of eruption rather
more recent than that of the Baldwin Agglomerates, and that
the true horizon of the base of the Barraba Series lies at some
indefinite horizon between the Loomberah limestones and the
Pyramid Hill tufi". As the outbreak of the huge eruptions,
which formed the Baldwin Agglomerate of the Manilla-Bingera
region, might reasonably be expected to have been associated
with some crust-movement in adjacent districts, we may put
forward the suggestion that the Scrub Mountain conglomerate
(the lithology of which seems to point to the occurrence of some
crust-movement) might be considered contemporaneous with the
Baldwin Agglomerates. This hypothesis has the merit of in-
volving a less radical change than the former one in the con-
ception of the range of characters of the Upper Middle Devonian
rocks, and is, therefore, adopted in the map. But it cannot be
considered as more than tentative, lacking as yet confirmatorv
evidence. On this hypothesis, the portions of the Nundle Series
lying south-west of Nundle Sugarloaf and Yellow Bock Hill (3,
Plate xxii.) belong to the Harraba Series.
The Devonian Igneous Rocks.
Igneous Bocks in the Eastern Series. — The igneous rocks in
the Eastern Series must be first briefiy discussed, before passing
on to the more important rocks west of the serpentine. They
consist, for the most p^irt, of bands of more or less schistose
breccias and agglomerates, which are fairly abundant, and are
particularly well seen in a zone which runs nearly due north
from the western end of Portion 42, Loomberah, to the eastern
end of Portion 46; while less noteworthy masses occur, east of
the Peel River, in the Parishes of Gill and Nemingha. Massi^e
igneous rocks have a more restricted occurrence. Spilites are
found a mile north of Dungowan Hotel in Portion 52, Gill.
Vesicular keratophyre occurs near by in Portion 51, and in Por-
tion 43 of the Parish of Loomberah is a very small lenticle of a
342 GREAT serI'p:ntike belt of new south Wales, \ ii.,
quartzose dolerite* with zoned andesine-felspar. Larger bands
of fine-grained dolerite and spilite cross the valley north of
Wallaby Mountain and the hills to the east.
Lj neons Bocks in the Transitional Zone — More complex than
these are the features of a series of igneous rocks which lie along
the zone of passage between the Eastern Series, the zone else-
where represented by the Serpentine Line. The most northerly
of these masses are the group of dolerites and spilites by the
Loomberah Bridge in Portion 56, which occupy the greater part
of the small spur east of Sandy Creek. A very good exposure
of the spilite is seen in the road-cutting on the east of the hill,
in which typical pillow-structure is exposed {see Plate xxxiii.).
The pillows are about two feet in diameter, and have generally
a number of vesicles, often filled with calcite, arranged in con-
centric rows within the pillow. The rock itself is greatly altered,
as described below (p. 365). For this reason, the pillow-lavas
are more like the pillow-lavas seen by the writer in Cornwall f
than any others in the Great Serpentine Belt(5, p.l31 ). The
dolerite which occurs on the west side of the spur is also greatly
altered. The original ophitic structure has been preserved in
spite of the mineralogical changes. Magnetite is the only
original mineral; the augite has become platey chlorite, and the
felspar has been changed either to a mass of calcite with a small
amount of albite; or to much-strained quartz or chalcedonj' with
chlorite, spherulitic if in large patches, and sometimes associated
with a little carbonate. The dolerite itself is sometimes slightly
vesicular. These sills of spilite and dolerite are separated by
banded jasperised cherts or highly siliceous cherts which dip
W.4''N. at 60° on the east side of the hill, but dip E. at 60° on
the west side of the hill.
Southwards of these, in the eastern end of Portions bh and
54, and again on the boundary of 53 and 52, are further intru-
sive masses. Spilite occurs on either side of the hill of chert
and jasper in Portion 54, but further to the east keratophyres
are found.
" See footnote to p. 364
t For description of these lavas, see (17).
rJY W. N. BKKSOX. 343
Small intrusions of rather vesicular spilite occur at the south-
western corner of Portion 48, but the next important mass of
spilitic rock is the complex series of intrusions which runs
through the western ends of Portions 39, 38, and 62, and are
associated with serpentine in a complex fashion. The spilite
occurs in a series of overlapping lenticles, decreasing in size
from north to south; the rock is vesicular, full of secondary
chert, in most irregular strings and veinlets {cf. 5, p. 127). These
lenticles are separated from one another by narrow zones of more
or less silicified, crushed chert, into which the serpentine has
been injected. In one instance, on the northern erid of the
complex, the serpentine appears to be injected into the spilite
(though the abundance of drifted blocks makes the observation
uncertain), while at the southern end a small patch of vesicular
keratophyre, 10 yards long and o yards wide, has apparently
been caught up and included in the ultrabasic rock. The whole
structure seems to have resulted from the shattering of a mass
of spilite and keratophyre, by a series of oblique faults, into
which the cherty crush-breccias were dragged and the ultrabasic
rock injected.
South of these are the long, undisturbed, lenticular intrusions
of dense spilite on the western slopes of Wallaby Mountain, and
crossing the creek at its base. These are quite typical of the
non-vesicular spilites in the Eastern Series.
The keratophyres of the Transitional Zone are tirst seen in the
eastern end of Portions 53 and 54, and occur again on the
boundary of Portions 53 and 52. (A small mass of Nemingha
limestone lies just west of this last). In the first of these, they
are vesicular with amygdules filled with quartz, chalcedony,
chlorite, and calcite. They become abundant, however, in the
Water-Reserve between Portions 51 and 52, are vesicular with
abundant quartz in the amygdules, and microscopic examination
shows the abundance of quartz in the groundmass. To the
south, the mass of keratophyre broadens out into the largest
area of igneous rock within the limits of the map. Though only
about two hundred yards wide at the northern end of Portion
35, it becomes half a mile in width to the south, and continues
344 TtHEat serpentink belt of new south walks, vii.,
thus through Portious 36 and 37, at the southern end of which
it is cut oif by a fault, which throws it eight hundred yards to
the east. Thence it continues for a mile further, exhibiting
some very interesting features described below (p. 345). This
belt of keratophyre forms the western margin of the Transitional
Zone; except where jasperised, the rocks to the west of it ob-
viously belong to the Tam worth Series. In Portion 52, a small
amount of such jasperised rock occurs, the relations of which
have been obscured through the ploughing of the land. More
distinct is the small mass of jasper in Portion 35, in contact
with a keratophyre rather rich in magnetite.
The main mass of the keratophyre is a rock of fine to medium
grain-size, consisting chiefly of albite, a few phenocrysts lying in
a trachytic or spongy base, dotted with finely crystalline mag-
netite. A little quartz is present in the base together with some
chlorite. Occasionally, the rock is quite coarseh^ crystalline,
and contains a little augite, approaching the character of albitic
quartz-dolerite (1414).
Along the eastern margin of this intrusion in Portion 36, the
rock is distinctly brecciated, this facies passing imperceptibly
into the massive rock. In microscopic examination, it has the
character of a flow-breccia, and sometimes appears to have been
originally more or less glassy. Fragments of albite occur with
very irregular outline and often slight strain-structure, also
irregular fragments of quartz and small grains of uralitised
augite, lying in a cryptocrystalline base (1411). In some speci-
mens, the matrix is the predominant portion; in others, the
larger fragments of crystals, evidently derived from a shattered
dolerite and quartz-keratophyre, form the dominant feature of
the rock. With such large fragments of single crystals may be
associated chips broken from a minutely trachytic keratophyre
(1406), such as are so abundant in the keratophyre-breccias of
Pipeclay Creek (5, pp. 15 1-156).
In places, the keratophyre becomes enriched in magnetite, and
a dark magnetite-keratophyre occurs by the jasper in Portion
35, as noted above, and in the south of Portion 37. The best
examples of magnetite-keratophyre, however, are to be found in
BY W. N. BKNSON. 345
a small complex, half a mile south of the south-eastern corner of
Portion 37, on the head of Bog Hole Gully, in which are ex-
hibited many of the features noted in the Hyde's Creek complex
(5, pp. 133-4). The keratophyre varies from a purely felspathic
type to a type irregularly blotched with areas enriched in mag-
netite, in a matrix of evenly coloured rock. There is also a
mass of more or less homogeneous magnetite-keratophyre, which
is, however, rather slaggy or vesicular, the openings being filled
with calcite (see p. 373). The margin of this irregular group of
intrusions is strongly jasperised, and the jaspers contain irregu-
larly distributed masses of haematite. There can be little doubt
that here also the ferruginous jasper has been developed by
solutions emanating from the keratophyre.
Towards the southern end of the zone of keratophyre, where
it is sharply cut oflf by a fault, the rock becomes much brecciated,
and peculiar spherulitic masses are developed, which, when
microscopically examined, prove to result from intense secondary
silicification {see pp. 374-375). The rock is traversed by numer-
ous veins of quartz, and, in optical continuity with the grains in
these veins, there are roughly circular areas in which quartz has
entirely replaced the minutely granular groundmass of the rock,
and forms a matrix in which lie embedded the more or less
idiomorphic felspar-laths. Each of these areas weathers out like
a spherule.
►Some long strips of highly silicified claystones are enveloped
in this mass.
71ie ly neons Hocks of the Western Series.
1 . The Pyroclastic Ptocks.
The Igneous Zone. — One of the most continuous and strati-
graphically useful lithological horizons in the Tamworth Dis-
trict is the Igneous Zone, which there lies about 800-1,600
feet above the horizon of the Nemingha limestone. This
is not a definite horizon, but merely a lithological zone in
which pyroclastic rocks were constantly present, and were often
associated with intrusive rocks. This zone is not so clearly
marked in tlie Loomberah District, but seems to be here repre-
34G ORKAT SKKPKN'I'INE BELT OF NKW SOUTH WALKS, vii..
sented b}^ a fairly continuous band of pyroclastic rocks, which
lies a short distance below the Loomberah limestone. Sills of
dolerite do not occur in this, but lie a short distance below it
as. for instance, in Portion 61, Loomberah (here the felspar is a
labradorite with a marginal zone of oligoclase), and in Portions
59, 58, 24, and 35, in which also the felspars are free from albi-
tisation. The tuffs, however, are more continuous than the in-
trusive rock, and may be traced right through into the Nundle
region .
The Nemingha lied Breccia. — A second igneous zone occurs in
the Loomberah District, which is of more distinctive nature and
stratigraphical value than the one last noted. It is in close
association with the Nemingha limestone, and may be termed
the Nemingha Red Breccia, It is especially marked by the
presence of a deep red colour. The breccia of which it is com-
posed has evidently been subjected to the action of solutions
rich in iron, soda, and carbonates, which have introduced much
magnetite and htematite into the rock-fragments, and converted
their felspars into albite and carbonates. The zone appears first
well developed in Portion 58 (Mr. Carter's property), tlie breccia
lying along the western side of each band of limestone {see p. 330).
It occurs again in the western corner of Portion 55 near the
Jasper Knob, and traces of it are seen in the fields south of
here. It does not form a definite band, however, until Portion
32 is reached, and there it is marked by a mass of porphyritic
dolerite. From this point, it extends to the south, closely asso-
ciated with the limestone, which at first lies on the east side
only, but, nearer Black Jack, appears also west of a band of red
breccias five hundred yards in width. Half a mile north of
Black Jack, the limestone and breccia-zone is thrown nearly half
a mile to the east and its width is nearly doubled, but a second
fault crossing the northern face of the mountain while moving the
centre of the band of breccia about a hundred yards further to
the east, causes it to return to a width of approximately three
hundred yards. It is here greatly broken, and numerous masses
of Nemingha limestone occur in it. The same horizon has been
traced across Cope's Creek, and as far as Silver Gully. Here,
BY W. N. BENSON. 347
the quartz-keratophyre breccia, associated with the limestone,
described and figured in a previous paper (5, p. 137), belongs to
this horizon. In the southernmost extension of the zone, the
strong impregnation of the rock by ferruginous solutions, and
its consequent red colour, are not always present.
We find in this another zone, which can be traced all through
the Loomberah District, and into the Nundle District. It is
also seen, in all probability, in the Tam^\'orth District. The
masses of red, ferruginous breccia associated with the southern-
most masses of the Nemingha limestone in the south-east of the
Tamworth Region, doubtless belong to this zone, and it is a
matter of great difficulty to determine how much of the pyro-
clastic rocks on East and West Gap Hills in that parish rightly
belong to the Nemingha Red Breccias or to the Igneous Zone.
The Silver Gully Agglomerate. — A short distance westward of
this zone, commencing at Black Jack, is a second mass of coarse
breccia associated with limestone, which, however, is nearly
always in the form of small fragments except on Black Jack
itself. This may be traced southwards across Cope's Creek, and
obtains a great width in Silver Gully. A small lenticle of lime-
stone appears on the slope to the north of this creek. It is
apparently heavily faulted near this creek. Beyond Silver
Gully, it continues as a narrowing band containing small lenti-
cular patches of limestone, but is cut off by a fault before reach-
ing Hyde's Creek.
Though the limestones on this zone are of the Nemingha-type,
though in small lenses, the lithology of the breccias, the occur-
rence of fragments of coarse dolerite, etc., and the absence of
red colouring, are features so distinct fiom those of the Nemingha
Red Breccias, that the two masses cannot be considered to be
on the one horizon. We may conclude, therefore, that the Silver
Gully Agglomerate is a third horizon of pyroclastic rocks, inter-
mediate in position between the Igneous Zone and the Nemingha
Red Breccias.
This intermediate horizon appears to be repeated on the
hills west of Wallaby Mountain. It commences immediately
south of the fault which truncates the keratophyre, and here is
348 GREAT SERPENTINE BELT OF NEW SOUTH WALES, vii.,
nearly half a mile wide, and includes a band of a peculiar por-
phyrite (see p. 369, specimen 1435). In lithological character,
the agglomerate is very like that in the band described last. It
may be traced continuously to the south, becoming narrower all
the way, until it dies out near the Pipeclay Gully.
Massive Igneous JRocks.
The Keratophyres. —The most striking feature of the massive
igneous rocks is the occurrence of a second long zone of kerato-
phyres extending from near the northern end of the map to the
southern. The northernmost occurrence in this zone appears to
be the few small sills at the head of Oakey Creek, in the Parish
of Nemingha; but, further south, there is the keratophyre-
complex by Macllveen's, which clearly belongs to the zone here
discussed. It has been already described (6, pp.57 1-3). South
of the Peel River, keratophyre appears again in Portion 55,
Loomberah, where there is a very inconspicuous outcrop of vesi-
cular, magnetite-bearing keratophyre to the M'est of Sandy Creek.
Beside it, there rises Jasper Knob, a huge mass of jasper, some-
times almost saccharoidal or miarolitic, with chalcedonic quartz
and finely divided platey haematite, not at all smooth and uniform
like the jaspers of the Eastern Series. The hill is about 200
yards long and over 100 yards in width, and is crowned by great
crags of jasper rising about 150 feet above the creek. To the
north and south, the country-rock is more or less jasperised
claystone. There can be no doubt that the presence of the
jasper is due to the impregnation or replacement of the claystone
by ferruginous solutions emanating from the keratophyre, just as
was shown in earlier papers {e.g., 5, pp. 133-4, 137, 164; 6, p. 57 2).
There is, however, a remarkable disproportion between the size
of the mass of jasper and the small amount of visible keratophyre.
Passing to the south, no more keratophyre belonging to this
zone appears for a distance of about four miles. Then it is seen
forming the low ridge in the east of Portion 31, the red breccias
and Nemingha limestone occurring along its western margin.
It is here a porphyritic rock, with large phenocrysts of acid
oligoclase in a finely trachytic base, in which a little quartz is
BY W. N. BENSON. 349
present. From this point, it may be traced across Reedy Creek
to the point where it is cut off by the important cross-fault men-
tioned below (.sec p. 351). This portion of the zone includes a
very unusual fades of keratophyre, namely, an aphanitic rock
with a semi-perlitic fracture. Microscopical examination proves
that this is a minutely crystalline variolite(si?e 1405, p. 369).
The zone is then thrown to the east, about 600 yards, by the
above-mentioned fault, but, a short distance further to the south,
it has been thrown back to the west, and continues along the
eastern side of the Nemingha limestone across Cope's Creek.
In this portion east of Black Jack, it is not entirely uniform,
some portions being rather less acid than others (see, e.g., the
description of 1383, p.370).
The keratophyre continues south of Cope's Creek, its intrusive
nature being rendered clear by the large block of limestone 600
yards south of the creek, which has been torn off from the
Nemingha horizon and enveloped by it. It may be traced from
this point down to Pipeclay Creek, where it is extremely sili-
ceous, and thence to Silver Gully. The features of this region
have already been briefly described (5, pp.137 and 154-5). Of
particular interest in this southern extremity of the keratophyre
zone is the occurrence of the Silver Gully keratophyre-complex
(jaspers, magnetite-keratophyre, etc.) which is intrusive into
the limestone. The brecciation of the keratophyre, and its pas-
sage into what seem to be breccias connected with the Nemingha
Red Breccias, are subjects well worthy of detailed study, which
the writer, by his removal from Australia, is unfortunately pre-
vented from making.
The Dolerites.
Three separate types of dolerite occur within this area,
grouped into three intermittent lines of intrusion. They may
be termed — reading from north to south — the Hypersthene-, the
Porphyritic, and the Albitic Dolerite.
The Hypersthe7ie-Dolerite. — Thin belt of dolerites has been
already mentioned as lying a short distance below the Igneous
Zone. Its northernmost occurrence is by the river-bank in Por-
25
350 GREAT SERPKNTINE BELT OF NEW SOUTH WALES, vii.,
tion 61, whence it may be traced througli Portions 59, 58, and
57. The largest mass lies nearly three miles further to the
south, namely, in Portions 24 and 35. This is an almost granit-
oid rock characterised by the presence of strongly zoned plagio-
clase, chiefly labradorite, and hypersthene in association with
augite and ilmenite. An analysis of this rock, 1387, is given
below (p.368).
It is not clear whether any of the dolerites further to the
south, such as that in Portion 37, should belong to this group,
though they were originally possessed of basic felspar, now partly
changed to albite and prehnite.
The Porphy7'itic Dolerite is intimately associated with the
Nemingha Red Breccias, and, therefore, has invaded a somewhat
lower horizon than the hypersthene-dolerite. It appears in
typical form, east of the limestone in Portion 59, as a handsome,
dark green rock (1407). It has pale green phenocrysts of acid
labradorite and dark chloritised augite, while the peculiar spongy
fabric of the base suggests a relationship to the keratophyres.
A much larger mass occurs in Portion 32. This occurs intrusive
into the Nemingha Red Breccia. In between these two masses,
the large and small fragments of porphyritic rocks in the
Nemingha Red Breccia have features strongly suggestive of close
association with, if not original membership of this group of
intrusions. The ferruginous, albitising solutions have, however,
strongly attacked the rocks, changing pyroxenes into iron-ores
and carbonates, while plagioclases, originally basic, have passed
into albite and carbonates. These have thus become kerato-
phyres through the alteration of dolerite-porphyrites, a process
analogous to that described by Neithammer(18) and E. B. Bailey
(19), and they are associated with fragments of normal trachytic
keratophyres, more or less impregnated with ferruginous solutions.
It is possible that these porphyrites may be connected with
the porphyritic spilites on East Gap Hill, in the Parish of
Nemingha(6, pp. 596-8). The latter pass without break into a
red breccia identical in all respects with the Nemingha Red
Breccia. The porphyritic rocks are not, however, identical. The
East Gap Hill rock has suffered to a greater or less degree from
BT W. N. BENSON. 351
metamorphosing solutions, which have replaced the basic felspar
by albite and minutely granular quartz-mozaic.
This correlation would involve some alteration in the details
of the tectonics and stratigraphy assumed previously for the
neighbourhood of East and West Gap Hills (see below).
7%e Albite- Dolerite. — This zone commences to the west of the
keratophyre in Portion 37, and occurs first in a series of small
isolated lenses running south-east until the first great cross-fault
is reached. No more appears for half a mile to the south of
this, but, beyond the second cross-fault, a large sill appears, and
continues thence without a break across Cope's Creek to Pipeclay
Creek, a distance of over two miles. The sill is nearly 200 yards
in width, and its dip is approximately vertical. At its southern
end, the dolerite is faulted against the eastern mass of the Silver
Gully agglomerate, and a great ridge of coarsely crystalline,
ferruginous jasper has developed in the plane of movement. (The
appearance of this ridge would lead one to expect that a little
ferruginous keratophyre would be found along its western face.
Time has not permitted the writer to search for this).
Petrologically, this zone is rather varied; a specimen taken
from the northern portion shows a mixture of sub-ophitic and
sub-variolitic textures, with clear albite, and augite (1378); but
west of Black Jack, in the main zone of rock, a dolerite occurs
(1376), the albite of which is very dusty and associated with
prehnite, and has evidently been derived from a more basic
felspar, with associated alteration of the augites to chlorite.
This zone of dolerite, lying as it does between the Serpentine
Line and the Nemingha Limestone, is probably to be correlated
with the dolerites that extend from the mouth of Sheep Station
Creek (three miles south-east of Pipeclay Creek) to beyond
Hanging Rock {see 3, Plate xxi.). We may also consider that it
finds a repetition in the zone of dolerites and spilites that lie
west of the Nemingha Limestone, and extend from near Bowlino-
Alley Point across Moonlight Hill and Tom Tiger into Swamp
Creek. In these, many examples of the sub-ophitic to sub-
variolitic albite-dolerites have been observed (5, PI. xxv., fig. 2).
352 GREAT SERPKNTINE BELT OF NEW SOUTH WALES, vii.,
Summary op Devonian Sthatigraphical Succession.
Before attempting to draw up an amended columnar section
(Text-fig. 1 ; see also 6, p. 549), we must point out the uncertainty
of some of the data upon which it is based. Owing to the rarity
of exposures showing the angle of dip, reliance must be based
TufF Qnd
MudsJ'one
wil'h Radiolar'ia
OcruD Mounlri
Conclome''ahe t
"inl'o C\uv)b^o•^e
Loornberoli
Uini«rsl"one
Cfierr^ Radio- i
larian flay^rooe )
Dolenfe ■
I anon C/ayslron
KeratoV>h:<re — >
Cher/y Radio
I a nan Clays Tone
Text-fig. 1.
Generalised cokunnar section of the Devonian rocks of the Loomberah
District.
upon the correlation of certain leading zones with other zones,
the horizons of which have been determined in the Tamworth
District. This correlation is often lithologieal only, and the
zones compared are sometimes pyroclastic accumulations, which
may vary greatly when traced in a lateral direction, or may
BV \v. N. 6E*rsoN. 553
completely disappear, or they may even be sills, which, in addi-
tion, may transgress from one horizon to another, while the
presence of an indeterminable amount of strike-faulting permits
frequent repetition or complete disappearance of some beds.
Hence detailed accuracy is impossible. One must either leave
the succession a chaos, or attempt to piece together the isolated
fragments of evidence, making as few and as reasonable assump-
tions as possible. The latter course appears preferable.
If reference be made to the earlier columnar section (loc. cit.,
supra), the lowest portion of phyllites, jaspers, etc., may fairly
be correlated with such rocks in the Eastern Series as are not
comparable with Western types. They are separated from the
Neraingha Limestone by the intrusive mass of the eastern kerato-
phyre, a great irregular sill, and an unknown width of transi-
tional rocks. There is possibly a fault west of this, causing the
disappearance of all but the small mass of the Nemingha Lime-
stone in Portion 52, which itself would then be a repetition of a
portion of the main zone of the Nemingha Limestone. Beneath
this main zone, there is also keratophyre closely similar to that
to the east of it, and tentatively correlated therewith. The
jSTemingha Red Breccia reaches its maximum thickness near
Black Jack, and is possibly there near the point of its eruption.
The Nemingha Limestone is not confined to one horizon only,
such as the base of the Red Breccia, but may occur in large
lenses at various levels within this breccia, though since the
breccia must have been rapidly formed, all such lenses of lime-
stone arc practically coeval ((/. 6, p. 575). The depth of the sea
during the deposition of these limestones must have been con-
siderable to permit of the rapid accumulation of such a thickness
of breccia, though Mr. Chapman's remarks suggest (see p. 390)
that this locally may have risen above sea-level and permitted the
foriiiation of oolitic limestone. Such local production of islands
of pyroclastic material was shown to be probable in the Tam-
worth District. The intrusion of the porphyritic dolerite seems
to have been closely connected with the formation of the
Nemingha Breccias.
'J'he deposition of radiolarian cherts and claystones followed
354 GREAT SERPENTINE BELT OF NEW SOUTH WALES, Vli.,
but was shortly interrupted by the outbreak of the volcanic
action which formed the Silver Gully Agglomerate extending
from Black Jack to Hyde's Creek, the centre of eruption
being probably near the southern margin of the map herewith
(Plate xxxii.).
Further radiolarian sedimentation followed, with minor
volcanic outbreaks, until the more extensive eruptions occurred
which liave produced the pyroclastic rocks of the Igneous Zone,
which are not as thick in the region under discussion as they
are in the districts to the north and to the south. Associated,
apparently, with this eruption was the intrusion of several
dolerite-sills lying below the horizon of the pyroclastic rocks of
the Igneous Zone. The sills exposed within the Loomberah
Parish are of normal composition, that to the south of Loomberah
being albitic. Whether the keratophyres below the Nemingha
Limestone were injected at this period or not, cannot be deter-
mined at present; they were evidently formed under a consider-
able cover of sediment. All the tuffs contain fragments of coarse-
and fine-grained keratophyric material (but not fragments of
ferruginous jasper).
Radiolarian sedimentation ensued once more, and the deposi-
tion of the Loomberah Limestone followed, the limestone rising
in places above the surface of the sea, so that it has distinctly
the features of a littoral formation. Nevertheless, the radio-
larian mudstones above and below it are quite of the normal
tvpe, the former containing abundant small intrusions of acid
tuff
Following the deposition of the Loomberah Limestone is a
series of radiolarian claystones and tuff. The most important
centre of eruption was near the head of Cope's Creek, the bands
dving away rapidly to the north, but continuing some distance
to the south. A less important centre of eruption lay near the
north-western end of the area mapped, in Calala Parish. Above
these agglomerates, the radiolarian clayslone gives place to a
radiolarian mudstone, softer, and not quite so finely granular as the
former rock. No sign of the Moore Creek Limestone is present,
though it might be expected near this region of change in char-
feV W. N. BEMSON. 355
acter of sediment. In this series occurs the zone of the Scrub
Mountain Conglomerate, which, commencing in the south-west
of Loomberah Parish, may be traced to beyond Nundle. There
is at present no evidence to show the change in geographic con-
dition which led to the development of this formation, but the
suggestion has been made above(p.341) that it represents a con-
glomerate laid down by a sea transgressing over a region which
has been upwarped during the eruption of the immense masses
of pyroclastic material forming the Baldwin Agglomerates, which
extend intermittently from Tamwoj-th to Bingara. If this be so,
the Scrub Mountain Conglomerate is approximately coeval with
that outbreak.
No angular or lithological unconformity, however, has been
recognised between the mudstones above and below that con-
glomerate. The mudstones above this region (which, if the pre-
sumed stratigraphic position of the conglomerate is correct,
belong to the Barraba Series) are exactly like those below. They
contain the Pyramid Hill tuffs, the centre of eruption of which
was near the Pyramid Hill Range, for they die out to the north-
west and also to the south-east, as is indicated by the topograjDhy
of the regions beyond the area mapped. Higher beds than this
have not been tiaced in this region.
'I'lic fSerpenliiie^^ etc.
The band of serpentine in the south-eastern c(n-ner of the map
of the Tam worth district, continues for nearly a mile till, near the
iiuiin road in Portion 83, Nemingha (behind Mr. White's resi-
dence), it is sharply cut off by a small fault, which lias displaced
the eastern series a distance of about 100 yards to the north-west.
The serpentines of the above band ai-e of varying width, and
largely schistose, and do not show an^^ relief. A notew^orthy
feature is the occurrence in the serpentine, in Portion 105, of a
very coarse pegmatitic mass of albite and quartz, which has
been shattered and seamed with further veins of quartz.
Similar albitic veins occur elsewhere in the serpentine (4, pp.69 1-
2). South of the above-mentioned fault, no continuous zone
of serpentine is met with. A small lenticle, about 60 yards in
356 GREAT SERPENTINE BELT Ot NEW SOUTH WALES, vii.,
length appears in Portion 54, Loomberali, and then no sign of
serpentine until Portion 39. Here, nearly six miles from the
last occurrence, the serpentine is found in intimate relation
with a mass of vesicular spilite. A single band of spilite has
apparently been broken by oblique faults, and, into these,
peridotite has been injected. The northernmost patch appears
actually to invade the spilite, but the rubbly nature of the ex-
posure renders this uncertain. The masses of serpentine to the
south have been thrust between the fragments of the sheared mass
of spilite, and, beyond these, .serpentine occurs here and there in
the line of shearing passing southwards up a small gully. Again
there is a gap of two miles free from serpentine. It then appears
south of Cope's Creek, as a narrow zone, widening to the south,
extending through a small saddle into the watershed of Pipeclay
Creek. This mass of serpentine contains abundant intrusions of
dolerite, of the type frequently present in such association, already
described from the neighbourhood of Bowling Alle3^ Point, and
Moonbi(5, pp.156-7; 6, pp. 6 15-6),
No further serpentine is seen till on tlic soutbern side of the Hat
t>l)ening at the mouth of Hyde's Creek, when a small patch appears
on the north side of the Peel River in Portion 9, Parish of Dun-
gowan, and is continued south of the river for a couple of hundred
yards, until it is cut off by the southern boundary -fault of this
ancient senkungsfeld-area (see below, p. 360). Its further
development commences in 8heep ^Station Creek a mile east of
here, and continues to Hanging Rock, as already described (3,
pp. 582-5).
Gratioithjire.
A single narrow \ein of creamy-white granophyre occurs in
Portion 35, and is, perhaps, to be correlated with the veins of
granophyre and porphyry associated with the gi-anite in the
Nundle District, though the neare^^t known occurrence of this is ten
miles away. It consists of fairly idiomorphic but very kaolinised
plagioclase, zoned but appai*ently acid in the main. These form
prisms 0-8 x O'l mm. in cross-section, with interstitial quartz
and sometimes granophyric intergrowth. Small, brown-green
BV \\. N. BENSON. 357
prisms of hornblende alsu occur, with fibrous extensions, and
rarely large grains of magnetite.
The Penno-Curbonij'crous Sandstones.
Nothing need be added to what has already been published con-
cerning these rocks (3, pp. 586-7) save in regard to their relation
to the faulting, which is discussed in a later section (p. 360).
Tertiary Basalt.
The occurrence of the Tertiary basalt is confined to a few locali-
ties. It is found only in necks and dykes. The largest neck is» the
mass of Black Jack itself, a roughly circular area of basalt in
which a thickness of about 300 feet of this rock are exposed.
There is absolutely no sign of underlying gravels, and the varying
height of the lower limit of the exposed basalt seems to depend on
the relation of the surface-topography to the margin of a vertical
column of rock. Half a mile to the north is another small plug of
basalt cutting through tlie limestones. This is only 100 yards in
diameter. A third occurs near the mouth of Hyde's Creek (5,
p. 172).
Finally, a small dyke of basalt has been noted near the head of
Reedy Creek, running in a direction parallel 1o tlie trend of the
dip-faults. These are (piite normal olivine-basalts, ccjiicerning
which no special petrological features are Avorthy of record.
Becettt Drift and Allaviam.
These terms are employed in the same sense as before (6,
l)p. 590-51)2). The greatest accunudation of drift is in the large
Hat near the head of Reetly Creek, a topograpliic feature Avhose
significance is not quite clear. The alluvium of the flood-plain of
the main river includes about eleven feet of loam lying above
gravel, which, in the single test known to the writer, reached a
thickness of thirty feet. The thickness of tlie upper stratum of
loam is subject to rapid variation. ;Mr. R. H. Dowe informed the
Avriter that in Portion 59, a fence four feet in height was com-
pletely covered in twenty-five years, while nearby, in the hole left
by a tree which had burnt down to the roots, it was obvious that
358 GREAT SERPE?JTINE BELT OF NEW SOUTH WALES, vii.,
the lloo'l-plain level had risen at least eight i'eet durnig the life ot
that tree.
Foldinu and FaidtiiKj.
A\ e are iiu\n in a position to smnniarise the teetonic struetuie
of the district, as deduced from the assumptions made in our
stratigraphical correlations. The most ob\ious fact is a general
close folding- along an axis ruiniing in a direction averaging N. 22*^
W.-8. 22^ E. (magnetic). The folding force came from the east,
and the eastern side is most affected. There is a median zone of
great crushing into which the serpentine.s have been injected, and
a western zone with a dip at first nearly vertical though predomi-
nantly Avesterly, decreasing in amount as we pass in that direction.
East of this Serpentine Line, however, the strike N.22"W. is by
no means universal, but a more nearly meridional strike is quite
common, as shown by the calcareous breccia on the i^outh-west of
Wallab}^ Mt., which runs obliquely to the Serpentine Line, or by
the tuffs, etc., outcropping by the roadside in Portion 43, or l)y
the northern end of the serpentine mass in Cope's Creek. This
recalls the virgatioii of strike in the Parish of Nemingha (6, map),
or that in the region of Mundowey on the Namoi(2 or 7, map).
But superposed upon this oblique strike is the regional strike
shown in the great lines of shear in wliich are developed the red
jaspers, such as those that form tlie cliffs of AVallaby ]Mt., and
extend to the mouth of Cope's Creek.
AVe concluded that the eastern mass of keratophyre, in the zone
west of the Serpentine Line, may have a fault along its western
margin concealhig the limestones, for the -most part. Another
])]-()l)ably occurs along the east side of the western keratophyre.
Between these, the chert dips E. SS'^X. at 85° in places, and is,
thei-efore, slightly overfolded (see Text-fig. 2). West of the
limestone, the beds dip to the west, but there is doubtless a con-
siderable amount of repetition by strike-faulting, with, occasion-
ally, a reversal in dip.
The Pyramid Hill Range structurally is a great syncline, cut by
one or more strike-faults. Tlie analysis of the probable folding
and strike-faulting thus indicates a structure comparable witli that
of the Tamwortli District and regions further north.
BY W. N. ftENSON.
359
^
o o
§ §
§ s
•? 'V ^ y)^
■■ vL^T'-'^^j'
i^ o
V-~i~-^^I —
\ — ■
'^ \^^
^ r:--t
R
g^-
^^
"^ -•
V)
_..
1^
-^— _'.
Text-fijr.-i.
Geological section across the Loombeiali District along tlic line AI5C
on Plate xxxii.
360 GREAT SERPENTINE BELT Oi N'EAV SOUIH WALES, Vli.
We lia\e now to consider a second group of faults, conceriiiiig'
wliicli little lias yet been mentioned. These run in a direction
appro.ximately at right angles to the main line of strike. One is
thus reminded of 8uess' discussion of Haw -faults or "blatter "(20),
and might suggest that they were here caused by the differential
folding thrust exerted between the Tamworth region, buttressed by
the Moonbi Granite, and the Xundle region buttressed by the
Duncan's Creek Granite; for, in those regions, strike-faults, such
as occur in the Loomberah region, are not nearly so .strongly
marked a feature. Against this, however, is the absence of any
dehuite evidence as to the relative age of these faults and the
granite-masses, and the certainty that considerable vertical, as well
as horizontal, movement must have occurred along the fault-planes.
The dip-faults are clearly 3'ounger than the folding and strike-
faulting, and, though the evidence is not yet conclusive, the dis-
placement of the serpentine from its position on the Peel River,
near AVarden's (Portion !), Dungowan), to the mouth of Sheep
Station Creek, half a mile to the cast, may well be due to the
marked dip-fault which runs from Hyde's Creek to the Peel River,
'file dip-faulting then Avould be later than the intrusion of the
ultrabasic rock. Furthennore, it was concluded that the large
open area at the mouth of Hyde's Creek owes its origin to the
removal, by the ri\er, of an ui-faulted block of soft Permo-Car-
boniferous rocks among the harder Devonian beds (3). The
southern boundary of this area is marked by the above-mentioned
dip-fault; the northern boundary is parallel thereto, and, in all
probability, has been determined by a similar fault. The eastern
and western limits of this area are suthciently nearly parallel to
the main strike to suggest that they were determined, in some
degree, by movement along lines of strike-faulting. Hence part at
least of the movement along these fault-planes must have occuiTed
after the deposition of the Permo-Carboniferous Sandstones. This,
however, was perhaps posthumous movement on fault-planes first
formed during the late Carboniferous folding. The author's
removal from the State has prevented his completing the detailed
study of the faults around this down -thrown area.
(Coiitiimtd oil p.-li)-'>.)
361
NOTES AND EXHIBITS.
Mr T. Steel exhibited the gizzard of a Muscovy duck with a
lateral, vermiform appendage, three-quarters of an inch in length.
Mr. Fred Turner exhibited a specimen of Geranium molle
Linn., a European annual weed, found growing in Moore Park,
in 1891, a species not included in the late Dr. Woolls' "Plants
Indigenous and Naturalised in the Neighbourhood of Sydney "
(1891).
Mr. J. L Froggatt showed specimens of the common House-fly
(Musca domestica Linn.) bred from maggots collected from stale
and rotten liver in which blowfly-maggots had previously fed and
matured, at Moree, N.S. W.; the first time, as far as the exhibitor
could ascertain, that this species had been reported as breeding
in purely animal matter.
Mr. W, W. Froggatt exhibited specimens of three introduced
beetles of economic importance — A^iobium domesticiim Fourer,
a wood-borer, introduced from Europe in timber, recently found
damaging floors in several localities; Rhizopertha dominica Fabr.,
an Indian beetle, becoming a serious wheat-pest in South Aus-
tralia, and at Sydney; Spheno])ho7'us striatus Fahrs., a Brazilian
beetle, whose larvse attack the basal portion of the stems of
Banana-plants in Southern Queensland and in the Tweed River
district. Mr. Froggatt also communicated particulars respecting
recent migrations of mice in inland districts, and of small, car-
nivorous Marsupials {FhascogaJe Jiavipes Waterh.) which prey on
them.
Dr. Tillyard showed a specimen of shale with a fossil insect-
wing, from the roof of the coal-seam of the Sydney Harbour
Colliery, described in a recent paper.
Mr. Fletcher showed a series of Brachyscelid galls from one
branch of a tree of Eucalyptus resinifera^ some of which had
incorporated leafy branchlets.
362
ORDINARY MONTHLY MEETING.
July 31st, 1918.
Professor H. G. Chapman, M.D., B.S., President, in the Chair.
Miss Hkather fSiLERRiE, B.Sc, Science Research Scholar,
University of Sydney, was elected an Ordinary Member of the
Society.
The Donations and Exchanges received since the previous
Monthly Meeting (26th June, 1918), amounting to 2 Vols., 28
Parts or Nos., 1 Bulletin, 1 Report, and 5 Pamphlets, received
from 27 Societies, etc., were laid upon the table.
363
THE GEOLOGY AND PETROLOGY OF THE GREAT
SERPENTINE BELT OF NEW 80UTH WALES.
Part vii. The Geolohv of the Loomberah District and a
PORTION OF THE GOONOO GOONOO ESTATE.
By W. N. Benson, B A., D.Sc, F.G.8., Professor of Geology
AND Mineralogy in the University of Otago, N.Z , late
LiNNEAN MaCLEAY FeLLOW OF THE SOCIETY IN GeOLOGY.
With two Pal^.ontological Appendices by F, Chapman,
F.R.M.S., A.L.S., Palaeontologist to the National
Museum, Melbourne.
(Plates xxxi.-xxxviii,, and five Text-figures).
[Continued from p. 360].
Petrolooy ... 363
Notes ON THE Loo.MKERAH Fossils ... 375
Bibliography 382
PAL.i<:oxTOLoaicAL Appendices 385
Petrology.
The only rocks of ypec-ial interest, to which attention will be
devoted in the sequel, are the massive Devonian Igneous rocks. It
was hoped to make a complete study of the rocks which occur as
traginents included in the coarse Silver Gully Agglomerate and
the Scrub Mountain Conglomerate, but the securing of sufficient
material was prevented by want of time. The attention of future
investigators is directed to this point. Of the massive rocks, we
find, in this region, several types not elsewhere noted in the
Middle Devonian Series, namely, hypersthene-dolerite ; porphyritic
dolerite with basic felspar; various types intermediate between
doleriteor spilite, and keratophyre; a semiperlitic variolite; and a
peculiar pseudospherulitic or blotched, silicified, pyroxenic kerato-
364 GREAT SERPENTINE BELT OF NEW SOUTH WALES, vii.,
phyre. The conclusions previously reached, as to the mode of
origin of mag'netite-keratophyres, and their relation to normal
keratophyres, and jaspers are quite confirmed.
Dolerites^ and Sjnlifes of the Eastern Series and the Transitional
Zone,
In Portion 43, tliere occurs a very quartzose dolerite, No. 1398.
The felspar occurs in perfect prisms and prismoids. It is an ande-
sine only slightly zoned and very turbid. The augite is in idiomor-
phic grains, with a normal optic axial angle. It is more or less
replaced by chlorite. Ilmenite forms large angular grains; and
quartz is very abundant in large interstitial grains, and is never
granophyric. The spilite (1390) which occurs in Portion 36 hy
tlie western boundary of Portion 47, is a very fine, even-grained
* In a review of the earlier parts of tliis series of papers, Professor
Johannsen (Journ, of Geol., 1917, p,494) writes: — "The term dolerite
is apparently used in a diflerent sense from that common in the United
States, where it signifies a coarse-grained basalt containing a basic plagio-
clase. The writer speaks of albitization proceeding inwards in the felspar,
by which he apparently means that the sodic rims are secondary. It would
seem more probable that the zonal rims are primary. The rock thus ap-
pears to be an augite-andesite." It is, therefore, desirable to state that
the albitic rocks indicated are structurally, and in their mode of occur-
rence, similar to those termed albite-diabases by Flett and Dewey (Geol.
Mag., 1911), and differ from them chiefly in the better preservation of the
pjTOxene. Subsequent research has shown that they are associated with
rocks with basic plagioclase, which are quite similar to, sa3% the Devonian
diabases of Germany. The name dolerite was adopted in conformity with
Dr. Marker's practice (see Petrology for Students, 4th Edition) to indicate
the original nature of the rock. The term augite-andesite might have been
misleading. The secondary nature of the albite was inferred partly from
comparison with the analogous rocks studied by Flett and Dewey. Sub-
sequent work led the writer to doubt the universality of the albitisation,
and the present investigation leaves him with the feeling that the albite
may sometimes be a primary magmatic crystallisation, sometimes the
result of secondary post-volcanic replacement bj" magmatic solutions,
Analj'ses show that such albitisation is accompanied by an addition of
soda. In the case of analcite, it has been shown that these two stages
merge into one another, and it is suggested that the same may hold in
regard to albite,
BY W. N. REXSON. 365
roc-k with small phenocrysts of augite and albite. lii ; trueture, it
is closely comparable with the usual types of spilite in the Eastern
Series, but it ha.s been ratlier crushed. The highly altered pillowy
spilites and associated dolerites in Portion 56 remain to be men-
tioned. The dolerite (1377) was originally ophitic, and though the
rock has been completely decomposed, the structure has been
retained. The felspar has been replaced by strained (juartz and
chlorite, the latter when forming a large area being generally
spiiernlitic. Tiie silica is sometimes ehaleedonic, and may have
narrow margins of carbonate. A second form of alteration is to
calcite, with a small amount of I'esidual albite. The augite is
entirely changed to platey chlorite. A little magnetite remains.
The spilites forming the pillows are represented by Nos. 1112 and
1 1 24. In these, the original structure is more or less lost, owing to
the decomposition of the constituent minerals. Traces, however,
indicate that the felspar formed prisms, which are now replaced
eitlier by chlorite (1124) or a clear, colourless, intermediate
material (1112), which is perhaps a zeolite. The pyroxene w^as
idiomorphic (1112) or ophitic (1124), and is now^ changed to
chlorite in green plates with splierulites, or associated with dolerites
or calcite. The formation of chlorite is accompanied by the
sejiaration of magnetite.
TliP Igncoiin Rochs in tlip Tmnwoiih Serifs.
The Dolerites.
The Dfjleritrs icit]i hasic felspar occur in tlie nortliern of the tw^o
series of intrusions already described, those with albite are in the
southern intrusions. The freshest example of the first group is a
rock (1387) from the large intrusions in Portions 24, 35. It has
a structure intermediate between the granitoid and subophitic.
The felspar is strongly zoned, tlie main portion being labradorite.
the margins of the grains oligoclase. The ferromagnesian minerals
are abundant, augite in sub-idiomorphic grains being predominant.
Hypersthene, however, occurs in considerable amount in large,
roughly idiomor]ihic grains (upon which tlie augite is sometimes
moulded), occasionally containing idiomorphic plagioclase. Tlie
26
366 GREAT SERPENTINE BELT OF NEW SOUTH WALES, vii.,
hyperstbene is decomposing peripherally in the usual manner.
Biotite occurs in very small .amount in minute plates. Titaniferous
magnetite is abimdant in irregular masses moulded to some extent
upon the felspar and pyroxenes. Apatite forms abundant large
prisms, and quartz occurs in a considerable amount in granular
mass in interstices. A very complete analysis of this rock by Mr.
Mingaye is given below. It is of particular interest as being the
first instance of a hypersthene-})earing rock observed in the
Devonian series.
Closely related to this, but less ricli in coloured constituents is
the coarsely granular dolerite in Portion 61 (138-1). In this, the
rhombic pyroxene is entirely decomposed, and is represented by
pseudomorphs of talc and chlorite. Prehnite also occurs as an
alteration-product. The small ])atch of dolerite east of tlie kerato-
phyre in Portion 37, is apparently of compound nature, containing
both normal and albitic dolerites. The first of these is exemplified
by specimen No. 1 409, a coarsely granular dolerite, with a felspar
chiefly labradorite, though clear patclies of oligoclase-andesine
occur. The felspar is largely altered to prehnite. The augite has
been partly alteied to form chlorite, and crystals of magnetite are
present. Quartz and liypersthene are a])sent.
The Alhitie Dolerites. — Tlie albite-dolerite in the occurrence
mentioned above is a very dear, fresh rock (No. 1378), fine-
grained, and with a glomero-porphyi'itic to ophitic fabric, with
small, fui^grained, sub-variolitic patches. The felspare give no
sign of zoning; they are quite fresh albite-oligoclase, and, in the
variolitic areas, there are skeletal extensions from the ends of the
crystals consisting apparently of albite. The pjn^oxene is often
quite undecomposed; at other times, it is replaced by chlorite.
There are peculiar spherulitic aggi-egates of haematite in the vario-
litic portions. Ilmenite occurs in small crystals. Closely related
to the above is the dolerite (No. 1394) occurring by the boundary
of the Peel River Company's Estate, a mile due south of the last
occurrence. Tlie felspars, though albite, are a little dusty. The
augite-crystals are partly chloritised, forming irregular crystals.
BV W. X. BKXSON. 367
These two rocks clearly belong to the same group of dolerites as
figured in Part iv., Plate xxv., fig. 2.
The next mass of dolerite (No. 1393), half a mile to the south-
east of the point of occurrence of No. 1394, is a rock of medium
grain-size, with all)ite-felspar, intei*stitial quartz occasionally
granophyric, and augite almost completely changed to chlorite; a
little magnetite also is present.
The dolerite in the belt of rocks which crosses Cope's Creek, is
exemplified by No. 1376. Though the felspar in this rock is now
a very dusty albite, it has certainly been derived from a more })as:c
mineral. It is sometimes a little spongy, containing prehnite, wliieli
also occurs interstitially in large clear patches. The augite is
entirely replaced by cldorite; ilmenite and apatite occur in ratlier
large grains.
The Por])]i(/n'tic Dolerites, generally closely associated with the
Nemingha limestone, may be illustrated by the description of three
specimens. It occurs in Portion 59, immediately east of tlie lime-
stones, and is a fresh-looking, dark green rock with phenocrysts of
felspar and augite(No.l407). The felspai* is an acid hibradorite
and is only slightly zoned. The augite-phenocrysts are, in parts,
quite fresh, while some are completely changed to pseudomorphs of
chlorite, and carbonates in one instance, with a further replacement
of the centre of the pseudomorph by quartz and chalcedony.
There are also small phenocrysts of magnetite. The base has a
spongy fabric similar to that of quartz-keratophyre. It consists of
short laths of dusty labradorite, an abundance of minute crystals
of magnetite, and a good deal of inter.^titial quartz.
368
GREAT SERPENTINE BELT OF NEW SOUTH WALES, Vll.
The following- analj^ses, generously contributed by Mr. J. C. H.
Mingaye, F.T.C., F.C.>S., show the composition of this rock, and
also that of the dolerite with basic felspars.
No. 1887.
No. 1407.
SiO,
51-14
58-17
AL03
14-47
16-07
Fe„0,
:vm
2-60
FeO
8-28
3-96
MnO
0-22
0-18
NiO, CoO
abs.
abs.
MgO
5-80
3-22
CaO
9-64
8-18
BaO
■■trace
0 07
SrO
tpresent
•i-trace
Na.O
2-43
2-22
K,0
057
1-90
Li„0
abs.
abs.
h;o-
0-34
0-22
H0O+
2-52
1-78
COo
F '
TiO.
abs.
0-93
0-75
0-55
ZrO:
abs.
abs.
p-o;
0-08
0-16
Fe8.
abs.
007
8O3'
abs.
abs.
CI
"trace
* trace
Cr,03
"trace
abs.
V^03
0 03
0 02
1
99-87
100-30
i
Specific gravity
2-929
i 2-763
No. 1387 — Dolerite with hypersthene and basic felspar. Portions 24, 35,
Parish of Loomberah.
No. 1407 — Porphyritio Dolerite, Portion 59, Parish of Loomberali.
These analyses should be compared with tliose given on page 602
of the fifth paper of the present series (6).
No. 1416, from Portion 32, is very similar in general eliaraeter
to No. 1407, though rather more decomposed. The presence of
chlorite replacing the inner portion of a zoned plagiochise-pheno-
cryst is a noteworthy feature.
What appear to liave been originally similar rocks, thougli now
BY W. N. BENSON. 369
altered by the ferruginous and albitic solutions which traversed
the Neming-ha Red Breccia, occur in fragmenta by the limestone in
Portion 58. Specimen No. 1116 is a good example. of these. It has
a ijub-variolitic to pilotaxitic base, consisting of small laths of acid
plagioclase, from which grow-out skeletal felspars or sub-radiating
microlites, with finely divided interstitial magnetite, together with
larger well-formed magnetite crystals, interstitial quartz, and a
very little chlorite. Here and there are irregular areas of calcite.
The large phenocrysts of felspar are entirely replaced by albite,
dotted with calcite. The trace of the original zoning is very
obvious.
A peculiar type of porphyrite occurs interstratified in the
agglomerates between Black Jack and Wallaby Mountain. It is a
dense fine-grained, black rock (No. 1435) with phenocrj-sts of
fresh augite, usually singly twinned, and large crystals of plagio-
clase which are extremely full of inclusions of chlorite, apparently
replacing augite-grains, or portions of glassy groundmass, con-
taining skeleton-crystals of ilmenite. The groundmass of the rock
luis a basaltic texture more like that of some lamprophyres than of
spilites. It consists of long prisms of augite and of plagioclase
with .skeleton-ilmenite, together wdth a considerable amount of
platey chlorite.
Variolite is represented by a single example (No. 1405). It
occurs in the Peel River Company's Estate half a mile to the south-
west of the soutliern corner of Portions 31 and 37 (Loomberah).
It is a dark green, apbanitic rock with a very poorly developed
perlitic fracture. IMicroscopically, it proves to be an extremely
finely crystalline rock, composed of sub-radiating sheafs of minute
felsiDar-microlites, with fine grains of magnetite arranged among
them. There are a few small phenocrysts of albite and augite.
Th e Ke ra toph i/res.
The keratophyres may be grouped into five main divisions. i.The
pyroxenic keratophyre. ii.The keratophyre proper. iii.The mag-
netite-keratophyre. iv.The nodular siliceous keratoph\Te. v.The
keratophyre-breccia. These pa.ss into one another.
370 UKEAT SERPENTINE BKLT OF NEW SOUTH WALES, \ li.,
The jjijrojienic keratoplii/res are those line- and coarse-y rained
types of rocks, wliicli are intermediate in composition between tlie
dolerite-spilite group, on the one hand, and the t'elspathic kera-
tophyre on the other. They may or may not contain quartz. As
instances of these may be deiicribed specimen No. 1414. This is a
pinkish rock of medium grainsize and syenitic appearance, occur-
ring a quarter of a mile soutii of the point where Sandy Creek
enters the Parish of Loomberah. It is granitic in texture, con-
sisting of grains of albite up to 1 mm. in diameter, irregular
prisms of augite partly uralitised or altered to chlorite, prisms of
apatite, and a minor amount of spliene in irregular grains.
The western keratophyre-zone is exemplilied bj' two rocks, which
may be classed with the pjroxenic keratophyre-group. At the
northern end of this zone, in Portion 31, the keratophyre (No.
141.3) is porphyritic with phenocrysts of albite, augite, now com-
pletely chloritised, and ilmenite associated with small crystals of
apatite in a glomero-porphyritic .aggregate lying in a pilotaxitic
base of laths and .small phenocrysts of albite with interstitial
quartz. Further south, on the same belt, just east of Black Jack,
occurs a rock (No. 1383) rather more pyroxenic than this. It is
dark green in colour, consisting of well crystallised albite-laths,
generally about 0*2 mm. in length, together with small phenocrysts
of the same mineral 1 mm. in diameter, partly replaced by chlorite
zonally arranged. Interstitially between the felspar-laths is a
small amount of devitrified glass. The augite occurs in small
grains often more or less chloritised, and, in addition, there is a
fair amount of magnetite. The keratophyre of this zone is, how-
ever, usually more acid than this, as instanced by specimens
described below, and the highly siliceous rocks near Pipeclay
Creek.
Bearing in mind Neithammer's suggestion (18), supported by
E. B. Bailey (19), that keratopliyres are often only albitised
porphyrites, we may include in this group specimen No. 1123 from
the Nemingha Red Breccias (Portion 58). It is closely alHed to
No. 1116 described above (p. 369), and contains idiomorphic
l)henocrysts of plagioclase, in wliicli a little original andesine
BY W. N. BENSON. 371
remains, though most is replaced by albite or calcite. There are
also small pheiiocrysts of fresh augite and magnetite. The ground-
mass is extremely tine-grained, witii a traehytic structure dotted
with tlnely crystalline magnetite.
The karaiophyrrs proper^ which consist essentially of acid plagio-
clase, are rather less abundant than other rocks. They usually
have present in them some other mineral, and as this becomes suf-
ticiently important to distinguish the rocks, the necessity' arises of
adding a qualifying mineral name to the term keratophyre. The
keratophyre on the border of Portions 35 and 36, Loomberah, is
one of these containing the least amount of minerals other than
albite. It consists of minute laths of albite in pilotaxitic to traehy-
tic groundmass, in which a few small phenocrysts of albite-mag-
netite and chlorite apparently replace augite. The felspars of the
groundmass are also associated with a very small amount of inter-
stitial quartz, and there appears to be a small intergrowth of the
felspar-laths with quartz, forming little irregularly-shaped patches
with the same optical orientation throughout. The groundmass is
cut by a few fine-gTained veins of clear albite. The most felspathic
of the keratophyres associated with the Nemingha Red Breccia
(No. 1123) is a highly porphyritic rock with phenocrysts of plagio-
clase, now albite, with a large amount of calcite, a little perfectly
fresh augite, and *i few large grains of magnetite lying in a
groundmass of very finely traehytic felspar, dotted with magnetite
dust and containing a very little interstitial quartz. Finely divided
calcite is also distributed in cloudy masses throughout the rock,
which has evidently been gi-eatly affected by carbonating solutions.
No. 1413. from near the head of Bog Hole Gully, indicates the most
felspathic member of the group of ferruginous keratophyres,
recalling the features of the brecciated keratophja-es from Hyde's
Creek (No. 1296, described in an earlier paper; 5, p. 151), though
the nodular structure is not so well developed. It conies from
near the southern end of the eastern mass of keratophyre. The
more ferruginous types of this series are considered below.
Qu((rtz-l'eratoi}]ii/res are more numerous. Some, such as No.
1134 from the mass of keratophyre east of Black Jack, contain
372 GREAT SEKPENTIXE BELT OF NEW SOUTH WALES, vii.,
small plieiiociysts of albite in a base of not very tine grainsize.
This is spongy in texture, consisting- of laths of albite, with a small
amount of magnetite-dust; rather larger, well formed laths lie in
the mass of microlites, together with large grains of quartz often
with chalcedonic margins. Exactl}' similar rocks have been
described from Pipeclay Creek, a mile and a half further to tlie
south-east along the same band of rock(5, pp. 155-(3). Another
type of quartz-keratophyre occurs in the creek in the Water
Iveserxe between Portions ol and o3. This is exemplitied by
specimens Nos. 1388 and 1421. The former contains quartz, form-
ing a general groundmass, in which are laths of albite with a
general trachytic aTrangement, .sometimes less abundant than the
quartz, but generally more so. Scattered about is chlorite (rei)lae-
ing augite), and crystals of magnetite. J^inesof shearing traverse
the rock, and the magnetite is often congregated in these.
As a final example of siliceous keratojjliyres, we may instance
Xo. 1401, from the extreme southern end of the eastern mass of
keratophyre. This is a very fine-grained rock, crushed slightly
and intersected by veins of quartz, often very narrow. The ground-
mass consists of irregular patches of minutely granular, untwinned
grains of felspar, in wliicli are embedded innumerable minute laths
of acid plagioclase with a more or less trachytic arrangement;
between these is a small amount of Hnely divided magnetite-dust,
and small irregular grains of quartz occur among this. There are
phenocrysts of albite, and a few irregular patches of chlorite. The
magnetite also occurs in the lines of crushing, being evidently
introduced secondarily.
The mafjnetife-l-eralophi/res may be subdivided into those whicli
contain quartz, and those which do not. We will consider the latter
first. Near the head of Bog Hole Creek, in the south-eastern end
of Portion 37, is a small mass of htematitic iron-ore, and by it is a
very liaematitic keratophyre (No. 1427), which consists of fine-
grahied, tracliytic albite-latlis, witli small phenocryst-laths 0-5 mm.
in length, densely impregnated with more or less oxidised mag-
netite. The rocks contain angular patches of trachytic kerato-
phyre, almost free from iron-ore. The latter is aggregated in the
BY W. X. BENSON. 373
cracks. Analogous specimens occur elsewhere, generally not so
oxidised.
The complex of magnet it e-keratopliyres at tiie liead ol Bog Hole
Gully, contains the same \ ariety of rocks as does that near Hyde's
Creek, previously described (5, pp. 150-151). 2\odular kerato-
pliyres occur (No. 1410), in which the rock is made up oi' frag-
ments oi' trachytic keralophyrc, \ery euriched in magnetite in the
centre oi' each iragment, but Tree Irom it at the margin. Between
the fragments are strings of more coarsely crystalline keratophyrc
and calcite, apparently introduced at tiie same time as the second-
ary magnetite, i.e., during or inmiediately after the brecciation.
Theealeite is thus to be considered as derived from the magmatie
solutions. In another example (No. 1120), the magnetite-kerato-
phyi'e is amygdaloidal, the vesicles containing calcite or quartz, and
deep green epidote, and occasionally large grains of magnetite.
The magnetite is i-ather une\enly distributed, but, in the irregu-
larly bounded regions in which there is very little finely divided
magnetite, there Is a roughly proportional amount of large ragged
aggregates of magnetite, sometimes surrounded by calcite, and in
particular, the magnetite is aggregated into crevices and around
the vesicles as described from Macllveen's complex (6, p. GOO).
Associated with this same complex, is the quartz-bearnig kerato-
phyrc very poor in magnetite (No. 1113) tlescribed above.
The Qaarlz-maynetite-kendophijres are of several types. The
most common is exemplified by specimens from the \\'ater
Reserve, betAveen Portions 51 and 52, by one (No. 1117) Irom
near the Jasper Knob in Portion 55, and by No. 1101 in Portions
35, 3G. All these rocks are vesicular, the vesicles being tilled
usually with (piartz and chalcedony, together with a little epidote
and occasionally felspar. Karely, they are filled with chlorite and
quartz. The groundmass of the rock consists of a spongy mass of
laths of acid i)lagioclase with interstitial quailz, dotted with
minute crystals of magnetite. The pheiuxnysts are albite some-
times undergoing" replacement by f|uartz, magnetite, rarely augite
(No. 1101). A fi-e(iuent feature occui-s around the margin of the
vesicles or in large irregular ci-acks. The spongy, texture of the
374 GREAT SERPENTINE BELT OF NEW SOUTH WALES, vil.,
grouiidmass here becomes exceedingly minute in grainsize, and
^e^y dark with finely divided magnetite. The groundmass is a
mosaic of minute interlocking grains of quartz. If felspar be
present, it is entirely untwinned, and the abundance of the mag-
netite prevents its recognition by the bright-line method. The
distribution of lath-shaped areas in the rock free from magnetite,
which w^ere doubtless originally felspar, indicates a general poorly
developed trachytic structure. Lai-ge areas rich in magnetite
appear to replace pyroxene-phenocrysts, and, in addition, there
are abundant phenocrysts of albite. This rock is possibly an
altered form of an augite-porphyrite.
The group of the Silicified-Nodular or Blotched Keratoplinres
comprises a series of very remarkable rocks occurring in the
southern end of the eastern zone of keratophyre. In hand-speci-
men, they are very fine-grained or aphanitic green rocks with
masses of silica, weathering out into beads set irregularly or in
long rows as in spherulitic rliyolite, or covering the whole surface
of the rock with a network-pattern of cracks weathered out between
more resistant silicitled patches. Microscopically, the rock, in its
unaltered state, Avould be classed among the more basic pyroxenic
keratophyres. It 1ms a trachytic base of small laths of acid felspar,
with an abundance of chlorite pseudomorphs after granular augite,
and scattered small grahis of magnetite. In this base are pheno-
crysts of albite, sometimes arranged in groups (the glomero-
porphyritio structure), a few irregular masses of chlorite, which
possibly represent original pyroxene-phenocrysts, and some small
phenocrysts or aggregates of magnetite. Well defined prisms of
apatite often appear in the vicinity of the phenocrysts. The
resistant portions of the rock prove to be rounded or irregular
regions in which the pyroxene has been completely removed, and
the felspar-laths remain apparently unaltered, lying in a matrix of
quartz which is in optical continuity over an irregularly bounded
area. Each resistant region or "pseudospherulite" may consist of
several such areas of quartz. Where the rock is traversed by a
small vein of quartz cutting through one of these replacement-
areas, the quartz in the matrix of the rock, on either side of the
BY W. X. BKNSOX. 375
vein, will often be in optical continuity with that in the vein.
►Sometimes, the rock contains only a few of such replacement-
areas ('^.^., No. 1408), which are generally rather sharply bounded,
but, in other specimens, the whole of the groundmass is replaced
(No. lttL'5), and very little indeed of the coloured constituents
remain.
In addition to these ^■eins oi' quartz, there are occasionally [e.g.,
in No. 138G) to be found veins less than 0-3 nmi. in diameter, which
consist of a line-grained mosaic of water-clear albite, which may be
associated with a little calcite.
The brecciated keralophijres Avhicli occur along the eastern mar-
gin of the eastern keratophyre zone, have already been sufficiently
described {see p. 344).
No further remarks need be added to the description of the
Devonian pyroclastic rocks given in previous papers. A number
of peculiar forms of alteration of the rock-fragments in the
Nemingha Red Breccia have been noted, but the general conditions
attending the development of that formation are now sufficiently
well estaldished, and these details are merely confirmatory.
Notes on the Loombekah Fossils.
As pointed out above, the fauna of the Loomberah limestone has
been found by JNlr. W. S. Dun to contain a number of new and
interesting forms, concernmg which he has kindly contributed the
following valuable notes, and to these are added further remarks
by tlie [)resent author, which are placed between brackets: —
"This limestone is, in most cases, of detrital origin, and, especi-
ally in the ease of the Rugose Corals, the preservation is so unsatis-
factory that, although manv^ of the forms are new, the specimens
are so imperfect, that it is impossible to obtain sufficient evidence
to make the diagnosis proper to a new species."
"Zaphrentis, sp. ind. — One specimen. The transverse section
shows 61 septa, with a dense psendocolumella, and limited stereo-
plasma. (See Plate xxxv., fig. 6.) The longitudinal section shows
very irregularly placed tabulae strongly concave."
^^Zaphrentis, (?)sp. — A single specimen of a very interesting
376
GREAT SERPENTINE BELT OF NEW SOUTH WALES, Vll.
Zaplireutuid coral. It is .simple, with a diameter of 13 mm. There
arc 08 septa, with tliick septal walls. The eosta? are very pro-
nounced, projecting for about 2 mm. The bases of the septa are
thick for a limited distance, the greater extent being free from
stereoplasma, Ihietuate, and not (juite reaching to the centre. This
is a most distinct form, and may constitute a new subgenus of
Zaphrentis, but the material available is too limited to enable a
satisfactory diagnosis to he drawn up.'' {See Plate xxxiv., lig. 1,
and Text-liu'. \\.)
Milhmel"re
Tfxt-lig..S.
Tianfjver.sc and uhliijuc sectiuiis of Z«;>/<re/<//s- s}).ii(j\'.(?suhgcii.iiov.), ( x 4),
Loomhcrah Liniet>toiie.
^'Cyathophijlh(m, sp. ind. — A large, simple type, the transverse
section only being available. Diameter, 16 '5 mm. The septa have
limited stereoplasma, and there are 58, comprised in two cycles.
The shorter septa are irregular in length, and frequently anasto-
BY W. N. BENSON,
^77
inose witli the primaries. The dissepimeiitul stnietiires are limited
in number and irregular (^'"c Te<t-fis'.4). There is no Australian
Cyathophylloid of this type hitherto described. The specific
characters are the sparse de- x'^^^Si^^'^
velopment of dissepimental
tissue, the steroplasmic layer ,^^\^
on all septa being developed
to a rather abnormal extent."
Text-fig. 4.
rraiisverse section of Cyal]iOphullnm sp.
iiov.. ( X 2), Looniherah Limestone.
'■''S'pon<.io2)hjjlhim, (?) sp.nov.
— An aberrant type of S'pou-
yopJiyllnm, in which the
structure of the corallite-wall
is masked, its place being-
taken by dense vesicular tis-
sue. (8ee PI. XXXV., figs. 2 and
?u and Text-fig.5). The septa
are more persistc^nt than in
other Australian Spongophylloids, and, in most cases, spring from
the replaced wall, there being a space of ai^out omm. of vesicular
tissue separating the se[)t;il ends of adjoining corjUHtes, Dissepi-
mental tissues are dense, and the \esicles are small. The septa in
tlie central area are 24-:i0 in number. To the naked eye, this
remarkable coral has the general appearance of a l'hi/li]jsastrcca
hut, under the lens, the septa are seen to be non-contluent. The
absence of a true wall ma}^ not necessarily separate it from the
Spongophyllums, as specimens of the typical S. biparlUa from the
Silurian of Yass, have, in some cases, a portion of the corallite-
wall undeveloi^ed. It is even possible that this may represent a
stage l)etween Spornjojili/fUum (sensu stricto), and Endophylhim.''
[A recent statement by Dr. S. Smitli appears to illustrate this
remark (21) — "Among the Rugose colonial corals there are cer-
tain genera, or certain species within a genus, in which the eoral-
lites have lost their epitheca, and are united by their dissepimental
tissue. The septa of adjacent corallites in the astraeiform colony
tend to become confluent, but all stages of this development, from
378 GREAT SERPENTINE BELT OF NEW SOUTH WALES, vii.,
that in which it is incipient to that of perfect confluence, are to be
found. In some eases, on the other hand, the septa of one eoralUte
do not extend to those of another, and leave an intervening space
Text-fig. 5.
Transverse section of Sponyophylhim sp.nov., ( x 4), Loomberah Limestone.
to tlie sole occupation of the dissepiments. AstrcTiform colonies
make their appearance in widely divergent stocks, and at different
periods of time; being, it w^ould seem, the ultimate terms in a pro-
gressive development along a w^ell-defined line."]
BY W. X. BENSON. 379
^^Phillipsastnea (jrandis, sp.nov. — A very large type of Phillip
sastrma, the centre of the coralHtes being 2 cm. apart" [2'5, 1*7,
and 1*6 cm. in three adjacent eorallites in the specimen illus-
trated in Plate xxx\-., figs. 4 and o.] ''The septa are variable in
length and number; there are 23-28 of the longer, and the shorter
reach to the limit of the central area. This species is quite distinct
from any of the Australian forms 1 have seen, and no closely allied
species from extra-Australian i-egions are known to me." [There
is a .species, P. gifias, occurring in the Lower and Upper ]\Iiddle
Devonian beds of North America, Onondaga and Hamilton Series,
in whicli the centres of adjacent eorallites are more tlian 2 cm.
apart (22); but further details concerning this form are not avail-
able liere. P. vernp/nilii, which was recognised by De Koninck(9)
in a specimen obtained l)y the Rev. AV. B. Clarke, from Cope's
Gully, near Hanging Rock, had from 28-32 septa, and the centres
of the adjacent eorallites are 10-12 mm. apart. It is extremely
pro])able tliat this form came from a flevelopmeut of the Loom-
herah limestone near the head of Cope's Creek, just outside the
limits of the area mai)ped in Plate xxxii.]
''Favofiltes gotJilandica/'
"Fuvosites^ sp. ind. (sj). nov. f). — Transverse sections of one
specimen oidy are available. Tliis form is distinct from any of the
types of the Au.stralian Devonian rocks, and is remarkal^le for the
very irregular form of the eorallites. (See Plate xxxv., fig. 7).
Additional material will probably prove it to be a new species."
[In addition to the forms recognised by IVIr. Dun, mention may
be made of a very interesting slide, which, unfortunately, has been
mislaid, and cannot here be figured. It was cut from a form,
apparently F. gotldandica, but in whicli the wide spacing of the
tabulas was replaced for a distance of about fi mm., by closely
])acked tabulae, as near together as in F. multitahulata. The
change occurred quite .sharply on one side, and more gradually on
the other, and at the same level in adjacent eorallites, as if brought
out by a rapid change and slower reversion in the local conditions.]
^^Plasmopora sp. — A single specimen of a small Plasmopora is
contained in the collection. It is quite indeterminable specif!-
380 GREAT SERPI^XTINE BELT OF NEW SOUTH WALES, vii.,
cally, though distinct from forms in the Nemingha limestont% and
the Mur rum bid gee beds."
[Ileliolitpfi 2^orosa. — This occurs in numerous large masses, the
exact shape of which is not apparent. Measurements of the
dimensions of the parts of the form agree exactly with those
given by Mr. Etheridge for corals of this species in the limestone
of Moore Creek (23).
CJutiHps .^tdliforriiU, sp.nov., Chapman. — An account of this
form is given in Mr. Chapman's interesting note in the second
appendix to this paper. While he refers to the rarity with which
it occurs out of the Carbonifei'ous system, it is interesting to
note that De Koninck remarks that the specimen of Campo-
phyUiim^tJpxiiOfiimi received by him from Quadong was enveloped
in a mass of some size of Cluefetes (/o/d/)issi(9, p. 54)].
"The Stromatoporoids, w^hich form the best index of the age
of the Loomberah limestone, are well preserved. They are re-
presented by the forms StromafojwrpJIa loomhprfii^i^, S. hensoni,
and Acfinoi^froma aiisfralp, all new forms, and an indefinite species.
The appearance of these is distinctly Devonian, the forms closely
resembling typically Middle Devonian species. As has been
pointed out by Heinrich (24), the discrimination of the different
species of Stromatopora is extremely diHicult, and the final test
is the conventional use of measurements of the interlaminar
spaces, and the spacing of pillars. Numerous averages of mea-
surements have been taken in the case of Professor Benson's
specimens, and it is thought that the erection of new species is
justified." [As will be seen from the photographs in Plates
xxxvi., xxxvii., and xxxviii., all enlarged two and a half diame-
ters, and from macroscopic examination, the f<il lowing appear to
be the characteristics of these species:
SfromafopovpUa loomhi'vensis. — The coenosteum is roughl}^ hemi-
spherical, sometimes reaching as much as 20 cm. in diameter. It
is divided into concentric latilaminie, about ."1 nnii. wide. The
vertical section (Plate xxxvi.) shows that the lamin.'e aie very
definite and continuous, except for the occasional presence of
crossinu' zooidal tubes. There are about sixteen lamime in the
HV \\. \. BKNSON. 381
space of a centimetre. They are connected by radial pillars,
whicli extend pei'pendicularly across all the lamina? making up a
latilamina, and may continue even beyond these. These pillars
also lie about sixteen in the space of a centimetre. Occasionally,
there are Caunopora-tubes, one of which (appearing in figure 8)
is about 3 mm. in greatest diameter. A tangential section shows
the presence of astrorhizte.
StromatirporpJIa benaoitl (Plate xxxvii.). — The shape of the
ccenosteum cannot at present be stated. The latilaminif are less
definitely marked than in the preceding species. The lamina.'
are so spaced that about fifty occur in a centimetre, while the
radial pillars are also about the same distance apart. They con-
tinue across several laminae, at times as many as thirty or forty.
The nariow, tabulate, zooidal tubes also continue across as many
as ten laminje, but are difficult to distinguish from the ordinary
intei'pillar spaces. Small astrorhizse are recognisable in the
vertical section. The Caunopora-tubes are abundant, and have
a diameter of about 0'5 mm., and show infundibuliform tabulse.
This form appears to be similar in some respects to FaraUel(ypora
dartinijtonensis (Carter), from the Middle Devonian of Devon-
shire.
Both these forms of HtvomatoporeJla show a more definite net-
work, and more continuous radial pillars than the majority of
the forms discussed by Nicholson (25).
The ccenosteum of *S'. loomherensis is unusually large, and its
lamina? and pillars are unusually far apart; in S. bensoni, on the
other hand, they appear to be closer together than is normal for
the genus. In addition to the two forms mentioned, there is a
large form apparently belonging to this genus, in which the pre-
servation is less perfect. In this, the laminai' and pillar-spacing
is farther apart than in the case of S. loombereiisis, there being
about ten laminte and nine pillars in the space of a centimetre.
The pillars are apparently short.
Actinostroma aiisfrale (Plate xxxviii.) is characterised by well
developed horizontal or slightly wavy laminse, with scarcely
noticeable latilamination. The type-specimen is a Hattish frag-
382 GREAT SERPENTINE BELT OF NEW SOUTH WALES, vii.,
ment, so that the form of the Cffinosteuni is not ascertainable.
There are thirty-eight laminiie in the space of a centimetre,
joined by pillars extending across as many as fifty lamina?, in
some instances. The lamina? and pillars together form a recti-
linear meshwork. There are about thirty-five pillars in a centi-
metre. No zooidal tubes are recognisable, but there are numer-
ous Caunopora-tubes, about 0*25 mm. in diameter, with infuii-
dibuliform tabulae. This form is extremely like A. clathratmn
Nich., and especially that variety of the species which occurs in
the Middle Devonian rocks of Germany. This latter species has
also been recognised by Professor Nicholson in the Devonian
rocks of Western Australia. Until the publication of his Mono-
graph (25), it was usually considered to be Stromatopora concen-
trica Goldfuss. The specimen here described as A. austraJe may
•come from the Loomberah limestone, but it is more probable that
it was found in Portion 163 of the Parish of Nemingha, "Beedle's
Freehold," in the Nemingha limestone.]
BIBLIOGRAPHY.
1. Benson, W. N. — "A Preliminary Note on the Geolog^^ of the Nundle
District." Report Aust. Assoc. Adv. Sci., 1911, pp. 100-106.
2. "The Geology and Petrology of the Great 8eipentine
Belt of New South Wales." Part i. General. Tliese Proceedings,
1913, pp. 490-517.
3. Part ii. Nundle District, ihkl., pp.569-o96.
4. Part iii. Petrology, IhkL, pp. 662-724.
5. Part iv. Spilites, keiatophj^res, etc., ibid., 1915,
pp. 121-173.
6. -— Part v. Tamworth District, ibid., pp.540-624.
7. Part vi. Western New England, 'dtid., 1917, pp. 223-
282.
8. Clarke, The Rev. W. B. — Remarks on the Sedimentary Formations
of New South Wales. Fourth Edition, 1878, p. 132.
9. De Koninck, Prof. L. G. — Descriptions of the Pala?ozoic Fossils of
New South Wales. Memoirs (4eol Surv. N. tS. Wales, No. 6.
10. Odernheimer, F. — "On the (Geology of part of the Peel River District
in Australia." Quart. Journ. Geol. Soc, 1855, pp. 399-400.
11. Stonier, G. A.^ — "The Geology of Swamp Oak and Niangla." Records
Geol. Surv. N. S. Wales, 1892, p.64.
BY W. X. BEXSON, 383
12. Dun, W. S. — PaLvontologist's Report. Aim. Kept. Dept. Mines N.
S. Wales, 1915, p. 189; 1916, p.218.
13. MiNGAYE, J. C. H. — Chemist's Report. Ann. Rept. Dept. Mines N.
S. Wales, 1915, pp. 193-4.
14. Jensen, H. I.— The Soils of New South Wales. Sydney, 1914, p. 16.
15. Dayis, W. M. — " Meandering Valleys and Underfit Rivers." Annals
of the Assoc, of American (iieographers. Vol. iii., 1914, pp. 3-28.
16. Ulrich, E. 0. — "Revision of the Palwozoic Systems." Bull. Geol.
Soc. America, 1911, pp.281-680.
17. Dewey and Reid — "The Origin of the Pillow-lavas in Cornwall."
Quart. Journ. Geol. Soc, Vol. Ixiv., 1908, pp.264-272.
18. Neithammer, G. — "Die Eruptivgesteine von Loh Oelo auf Java."
Tschermaks Min. Petr. Mitth., 1908, p. 21 8.
19. Bailey E. B. — (In) " The Geology of the Glasgow District." Memoir
Geol. Survey of Scotland, 1911, p. 127.
20. SuESS, E. — The Face of the Elarth. Sollas' Translation, Vol. i., p. 120.
21. Smith, S. — "On AuUna rotiformis, PhiJIijjsastrwa hennahi, and
Orionastnva.''' Quart. Journ. Geol. Soc, Vol. Ixxxii., 1917, p.281.
22. Grabau and Shimer — North American Index of Fossils, Vol. i., p. 69,
23. Etheridge, R. — "On the Corals from the Tam worth District, chiefly
from the Moore Creek and Woolomol Limestone.'" Records Geol.
Surv. N. S. Wales, Vol. vi., 1899, pp. 151-182.
24. Heinrigh, M. — "On the Structure and Classification of the Stroma-
toporoidea." Journ. of Geol., Vol. xxiv., 1916, pp.57-60.
25. Nicholson, Prof. H. A. — "A Monograph of the British Stromato-
poroidea." Monographs of the Palipontographical Society.
EXPLANATION OF PLATES XXXI. -XXXVIII.
Plate xxxi.
Topographical Map of the Loomberah District.
Plate xxxii.
Geological Map of the Loomberah District.
Plate xxxiii.
Pillow-lavas exposed in the road-cutting by the Loombeiah Bridge, eleven
miles south-east of Tamworth.
Plate xxxiv.
Fig. 1. — Section of the Loomberah Limestone, showing its fragmental
chai'acter; and transverse sections of Zaphrentis, sp.nov., (?subgen.
nov.), Z, Treptostomatous Bryozoans, B, valves of Fenfamenis, P,
and indeterminate fragments; ( '< 3/2).
384 (iREAT SERPKXTIXE BELT OF NEW SOLTTH WALFIS, vii.
Fig.'i. — Transverse section of SpoiujopliylhmK {?)sp.nov. , frcmi the Looiii-
berah Limestone; ( x 3/2).
Fig.3. — Longitudinal section of S.{'t), sp.nov,; ( x 3/2).
Plate XXXV.
Fig. 4. — Approximately longitudinal section of PhiUipsaMrfva ;/ rand is, sp,
no v., Dun; ( x 3/2); Loomberah Limestone.
Fig. 5. — Transverse section of P. qrandis, sp.nov,; ( x 3/2).
Fig.6. — Transverse section of Zaphrenfis, sp.ind.; (x3/2); Loomberah
Limestone.
Fig. 7. — Transverse section of FarosUes, sp.nov.; { x 3/2); Loomberah Lime-
stone.
Plate xxxvi.
Fig. 8. — StromatoporeUa /oomherensis, sp.nov.. Dun; vertical section; ( x 5/2).
Fig. 9. — S. loomberensis, transverse and partially tangential section showing
astro rhizfe; ( x 5/2),
Both from the Loomberah Limestone.
Plate xxxvii.
Figs. 10-11. — StromatoporeUa hensoni, sp.nov,, Dun; transverse sections
showing Caunopora-tubes; { x 5/2).
Plate xxxviii.
Fig. 12. — Actinoxfroma aiistrale, sp.nov., Dun ; transverse and partially
tangential section showing Caunopora-tubes; ( x 5/2),
Fig. 13. — A. austrah, vertical section; ( x 5/2). Probably from the Nemin-
gha Limestone.
385
Appendix i.
Devonian Foraminifera ; Tamworth District, Xew South
Wales.
By Fredk. Chapman, A.L.S., F.R.M.S,, Palaeontologist to the
National Museum, Melbourne.
(Plates xxxix.-xli.)
Introduction.
At the request of Dr. AV. N. Benson, B.A., F.G.S., I am giving
the details of foraminiferal evidence which I noticed whilst
examining some micro-sections of limestone from the Nemingha
horizon of the Tamworth Series The method of studying fora-
minifera from rock-slices, without accompanying specimens show-
ing the exterior of the test, is not entirely satisfactory, but the
fact that these organisms are of Devonian age is, in itself, of
sufficient importance to merit a record of the occurrence, fora-
minifera being extremely rare in Devonian faunas.
The Rock-structure and its Stratigraphical Association.
The foraminifera under notice occur in a well-developed oolitic
limestone in which granules form a little more than half the
bulk of the rock. The granules vary in diameter from 046 to
0*7 mm., and only a small proportion are entirely due to oolitic
accretion; whether originally of algal origin or not it is impos-
sible to say, on account of their present mineralised condition.
The nucleus of the oolite-grain in more than one case was seen
to consist of an ossicle of a crinoid. The majority of the grains,
however, are microgranulitic in structure, either in the nucleus,
or more rarely throughout the entire granule. Certain of these,
from their more irregular outline, and often without an external
coat of concentric oolitic deposit, led me to suspect their fora-
miniferal relationship, which conclusion is borne out by further
study of the specimens. Some of the perfectly spherical grains
28
386 DEVONIAN FORAMINIFKRA. TAMWORTH DISTRICT,
are, I feel convinced, of a radiolarian nature, and here and there
one can detect a central sphere. There is a fair amount of iron-
staining in the rock-structure, which seems to be entirely
secondary, as the stain is developed more strongly along incipient
fracture-lines than in the grains themselves.
From Dr. Benson's work on the rocks of the Tamworth Dis-
trict,* it is seen that the Nemingha horizon can be correlated
with the lower part of the Middle Devonian.
Previous Records of Devonian Foraminifera.
Tn turning to consider the occurrence of foraminifera in other
parts of the world, we find only one authentic record of these
minute fossils, viz., that of Terquem's, who figured and described
a few forms from the Middle Devonian of Paff'rath in the Eifel.f
On p. 41 7 of Terquem's note, he states that the foraminifera were
found in some sand contained in a Megalodon-^heW. They were
in the condition of casts. The material was probably referable
to foraminifera and ostracoda, but generally indeterminable.
Some spheres covered with thick and sharply pointed spines he
referred to Orhnlina. These were very common, and measured
0-48 mm. in diameter. A pyriform cast was referred to Lagemi-
lina (a subgenus of Lagena). A cristellarian w^as also noticed
by Terquem, presenting the arrangement of chambers seen in
CristeUaria vetusta from the Lias. A fusiform cast was com-
pared to Fusulina; and numerous globular casts, consisting of
two or three chambers, were identified as Glohigerina. Judging
from my own observations of radiolarian structures in the
Devonian of Silesia and Bavaria, I am inclined to think that
Terquem's prickly Orbulinse may have more than a fancied re-
lationship to the orbicular radiolarians. In connection with the
present work, I have recently examined some disintegrated lime-
stone from Paffrath, from shells in the National Museum, and
* " The Geology and Petrology of the Great Serpentine Belt of New
South Wales. Part v. The Geology of the Tamworth District."' Proe.
Linn. Soc. N. S. Wales, 1915, Vol. xl., Pt.3, pp. 540-624, Pis. xhx.-lviii.
t Terquem, O. — "Observation sur quelques fossiles des epoques pri-
maires." Bull. Soc. Geol. France, .ser.3, Vol. viii., 1880, pp. 414-418,
PI. xi.
BY F. CHAPMAN. 387
have succeeded in finding a small but representative series of
foraminifera, radiolaria, and ostracoda which I hope to describe
shortly.
The only other allusion to Devonian foraininifera that I have
so far come across, is that by E. Wethered, in his paper on the
limestones of South Devon.* In speaking of the organic con-
stituents of the Gouiatite limestone (Upper Devonian) from a
quarry at Whiteway Farm, near Chudleigh, Mr. Wethered says
— " The specimens collected as typical of this limestone show it to
be quite different in structure from the beds below. There is a
fine crystalline groundmass in which are several fragmentary
remains of organisms. One of these appears to be a foraminifer,
and it is especially interesting as being the only one found in
my slides of the South Devon limestones. The Goniatite lime-
stone seems to have been formed by an accumulation of small
shells, foraminifera, etc., which have been filled in with a fine
calcareous mud." Wethered does not, however, refer the fora-
minifer he saw to any particular genus.
Description, of the Foro,rtiini/era.
Fam. ASTRORHIZID^.
Subfam. SACCAMMININ.E.
Genus PsammospH/Era Schulze.
PsAMMOSPH.ERA XEMINGHENSI8, n.sp. (Plates xxxix , figs. 1-2;
xl, fig.lO; xli., figs.11-12).
Description. — Tent rudely spherical to elongate Hask-shape.
Wall composed of granular particles neatly fitted together, form-
ing one or two layers, and distinct from the large crystalline
grains of the matrix more or less completely infilling the cavity
of the test. Apertures apparently between the components of
the test- wall or, more rarely, as a slight protuberance on the
surface.
Diameter, circ. 0*4 to 0'75 mm.
Observations. — The above form has a certain resemblance to
* "On the Microscopic Structure and Re.sidue.s Insoluble in Hydro-
chloric Acid in the Devonian Limestones of South Devon." (^uart. Journ.
Geol. Soc, Vol.xlviii., 1892, pp.377-387.
388 DEVONIAN FORAMINIFERA, TAMWORTH DISTRICT,
Saccamndna, but it never has the orifice so strongly protuberant.
Psammosphcei'a, as a genus, has not been found in fossil deposits
older than the Jurassic, from which system Dr. Haeusler* has
recorded the living species P. fusca Schulze. Tlie present
species is less than half the size of the Jurassic and recent forms.
P. nemiiighensis is by far the commonest foraminifer in the
Nemingha limestone, representing about 90 per cent, of the
entire foraminiferal remains
Fam. TEXTULARTID^.
Genus V a l v u l i n a d'Orbigny.
Valvulina PLicATA, Brady. (Plate xxxix., figs. 3, 4).
Vaivnlina plicata Brady, 1873, Mem. Geol. 8urv. Scotland,
Expl. Sheet 23, pp. 06, 95, etc.: Idem, Pal. 8oc. Mon., Vol. xxx.,
1876, p.88, PL iv., figs.lO, 11. Chapman, 1907, "Notes on
Fossils from the Collie Coal-field," Bull. Geol. Surv. W. Austr.,
No.27, p.l6, PI. ii., figs.lOa-c.
Observations. — Several specimens occur in the microscope-
slides, which compare closely with Brady's species from the
Lower and Upper Carboniferous Limestone of England and
Scotland, as well as in the Fusulitta-heds (Carboniferous) of Iowa,
U.S.A. The writer has described a diminutive specimen of the
above species from the Carbopermian Sandstone associated with
Glossopteris leaves from Collie, W. Australia.
An example occurs in one of the slides, which indicates a
plastogamic union of two individuals, as frequently seen in Bis-
corbina, Vaginulina, and other genera.
Valvulina bulloides Brady. (Plate xxxix., figs.5, 6, 7).
Vaivnlina bulloides Brady, 1876, Pal. Soc. Mon., Vol. xxx ,
p.89, PL iv., figs.12-15. Chapman and Howchin, 1905, "Mon.
Foram. Peimo-Carboniferous Limestones of N. S. Wales," Mem.
Geol. Surv. N. S. Wales, Pal., No.14, p.l3, PL i., tigs.9a-c.
Observations. —This species closely resembles a high and tumid
Globigerina, but is easily distinguished by the open umbilical
cavity on the lower surface, and the valvuline aperture.
* Quart. Journ. Geol Soc, Vol. xxxix., 1883, p.26, PI. ii., fig.l.
BY F. rHAPMAN. 389
V. bulloides was previously confined tu the Carboniferous and
Carboperinian. Dr. H. B. Brady records from the Futtulitia-hed^
of the Upper Coal- Measures of North America, the Calcaire de
Naraur of J^elgium, and the Fusulina-Limestone of Miatschkovo,
near Moscow. The same species was described by Mr. Howchin
and the writer from the Carbopermian limestone (Branxton Stage
of the Upper Marine Series), of Wollongong, N. 8. Wales.
Not uncommon in the Nemingha Limestone.
Valvulina oblong a, sp.nov. (Plate xl., fig. 8).
Description. — Several examples of an elongated valvuline type
occur in the slides. They remind one of a short, stout Haplo-
phraginium, or the Lituola nautiloidea Lam., figured by Dr. Brady
from the Carboniferous of Northumberland,* but show the de-
pressed base and characteristic aperture of these Devonian and
Carboniferous modifications of Valvulina.
The longer diameter of a fairly complete specimen in section
is 1 mm.; the shorter diameter, 0*65 mm.
PuLViNULiNA BENSOM, sp.nov. (Plate xl., fig. 9).
Description. — This species differs from the majority of the test-
sections in the micro-slides of the Nemingha limestone in having
a finely granulate shell-wall, which may, at one time, have been
of hyaline structure. The section figured shows a certain amount
of depth, although sliced, and indicates a pulvinuline type of
shell with few, lobulate chambers, and a wavy, peripheral margin.
Jt was probably depressed on the superior face, and slightly
conoidal on the inferior.
The nearest Palaeozoic form to this seems to be P. hrorchijana
Brady,t from the Carboniferous Limestone of Namur, Belgium,
but diifers essentially in its rounder outline and less inflated
chambers. It belongs to the Fii/viiiulina eleyaiis group, and,
from its size and contour, indicates a shallow-water variant of
that type of shell.
* Pal. Soc. Mon., Vol. xxx., 1876, p.63, PI. viii., figs. 7a, b.
t Pal. 8oc. Moil., Vol. xxx., 1876, p.l40, PI. vi., tigs.r2a-t'.
390 DEVONIAN FORAMINIFERA, TAMAVORTH DISTRICT,
Summary.
The components of the Nemmgha limestone appear to be
largely foraminiferal, comprising, so far as the evidence from
microscope rock-sections shows, an abundant species of Psammo-
sphcera —P. neminghensis; and the following more or less sparsely
represented — Valvulina plicata Brady. V. hnlloides Brady, and
V. oblouya, sp.nov.; whilst Pulvinulina appears to be present in
P. bensoni, sp.nov.
The genus Psainmosphcp.ra is more at home in deep rather than
in shallow water, though it has been found in depths as little as
45 fathoms. The other genera indicate a fairly shallow-water
habitat.
One special point of interest stands out conspicuously in the
f <n-aminifera of this limestone, viz,, the rather strong resemblance
of the rare Devonian foraminiferal fauna to the Carboniferous
and higher beds of the Palaeozoic.
Radiolaria seem to form a considerable proportion of the
granules in the Nemingha limestone, but their mineralised con-
dition precludes any reference to definite genera.
As regards the probable depth at which the rock was formed,
one might reasonably assume it to be of moderately deep water
origin from the occurrence of Psavimosp/ict^ra and the radiola-
rians. On the other hand, oolitic grains are indicative of quite
shallow-water areas, though this latter constituent is by no means
predominant, and may not materially affect the above conclusion.
The general condition of deposition must have been such as
would obtain in a moderately deep inland gulf with quiet sedi-
mentation and accumulation of ooze-forming organisms: and it
is just possible that the oolite-grains may have been blown into
the area of deposition from dunes of shore-sand origin in the
vicinity.
In conclusion, I would tender my sincere thanks to Dr. Benson
for giving me the opportunity of examining this interesting lime-
stone, and facilitating this attempt to describe a unique faunula
by furnishing the microscope preparations.
BY F. CHAPMAN. 391
EXT>LAXATIOX OF PLATES XXXIX.-XLI.
Plate xxxix.
Fig. 1. — P.^ammospJiii ra iH//iiit;/htn.-^l.^, sp.nov. Section through test show-
ing shell-wall and introverted aperture.
Fig,2. — P. nemitif/henms, sp.nov. Section showing interior thickened with
secondarj'^ ealcitic deposit.
Fig. 3. — Valni/ina ;>/?Va/o Brady. Section through and a little above the
hase.
Fig.4. — r. plicata Brad3'. Two tests, probahlj- in plastogamie union.
Fig.o.— r. biUloide-^ Brady. Flattened and almost complete test.
Fig.O. — r. /)«//o/c/es- Brady. ^Median section.
Fig. 7. — r. IndlokleM Brady. Nearly complete test.
All figures magnified Trl diams.
Plate xl.
Fig. 8, — Vulrnltna oh/oiu/a, sp.nov. Basal section; ( x .Vi).
Fig. 9. — Pith'inu/lna h('n-<ou't, sp.nov. Almost complete test; ( x .12).
Fig. 10. — Section through P>iainmof<p]ufra nem i nf/hensls, ^howmyr relative
thickness of test; ( x 144).
Plate xli.
Fig. 1 1. — Section of Nemingha limestone with oolite-grains, Psiuninosplntra,
and (V)radiolarian; ( x 2S).
Fig. 12. — Ditto, with oolite-grains and Pscuninosplittra; ( x 28).
392
Appendix ii.
Note on a new Species of CHiETETES.
By Fredk. Chapman, A.L.8., F.R.M.S., Palaeontologist to the
National Museum, Melbourne.
(Plate xlii.)
Introduction.
The genus CJuptetes, founded by Fischer in 1837,* is almost
entirely confined to the Carboniferous. Nicholson, in liis
'•Tabulate Corals of the Palaeozoic Period,*'! says "The species
of ChfHetes as here defined, are not known to occur out of the
Carboniferous (and possibly the Devonian) rocks,'' but he does
not specify any occurrence from the latter system. Zittel states
that the genus is also found in the Lias and Upper Jurassic;
It is, therefore, deeply interesting to find an Australian species
of the C. radians type in the f^oomberah Limestone,§ which, if
Dr. Benson's final conclusions bear out his preliminary investi-
gations, belongs to the upper part of the Middle Devonian. The
peculiar habit of the corallites in this genus, in showing partial
fission of some of the calices by imperfect partitions which are
tooth-like projections in horizontal section, is well shown in the
present species. Further, it is important to note that our species
shows the presence of a dark line in the septal wall, notwith-
standing its supposed absence from that of C. radians.\\
* Oryet. de Gouv. de ^Sloscou, p. 159.
t London, 1879, p. 266.
::: See Zittel. Traite de Paleontologie, 1888, Vol. i., Pt. i., p. 628. Also
Eastman-Zittel, Text-book of Paleontologj-, Vol. i., 1918, p. 118.
§ See Proc. Linn. Soe. N. 8. Wales, 1915, xl., Pt.8, pp.546, 549, and 559.
I Eastman-Zittel says {op. cit., pp. 117) — " Walls thoroughly amalga-
mated, common to adjacent corallites.*' Nicholson [op. cit., p.263) says
'• Rough fractures (generally, but not always) expose the interior of the
tubes; and thin sections, whether transverse or longitudinal, show that
the walls of contiguous corallites are entirelj' and undistinguishabl^- amal-
gamated or fused with one another, the originallj' duplex character of the
partition between neighbouring corallites being in no case recognisable."
BY F. CHAPMAN. 393
Ch^etetes stelliformis, sp.nov. (Plate xlii., figs. 1-3).
Description. — Corallum mas.sive, moderately large. Corallites
crowded, .slender, and gently curved. In cro.ss-.section roundly
polygonal and occasionally elongated, averaging about 0-5 mm.
in diameter; with one to four, but generally three, blunt peg-
like teeth projecting into the cavity, representing imperfect
fission of the calices by longitudinal partitions on the calicular
wall. Tabulai well developed, from 05 mm. to 1-5 mm. apart,
sometimes in continuous planes across the corallum, but often
more irregular and strongly curved. Unlike other species of
ClKtfples, a fine but distinct dark line is seen dividing the walls
of the cells (see remarks antea).
Observations. — This coral is of the type of the abundantly dis-
tributed Cluvietes radians Fischer, from the Lower Carboniferous
of Hussia, England, and elsewhere,* but is distinct in many
points, as in the more roundly polygonal calices, the more
numerous longitudinal partitions appearing as projecting tooth-
like processes in cross-section, and in the strong fusion-points
where calice-wall and tabulae meet. In longitudinal section, this
latter feature is verj^ marked, and, at the point of junction, forms
a stout cross with pointed arms.
ChceteteK depressus, Fleming sp.,t is characterised by much
smaller calices, measuring 0*2 mm. to 0'26 mm., or about
one-half the diameter of C. stelliformis. The diameter of the
calices in C. radians, on the other hand, is practically the same
as that in C. stelliformis, viz., about 0 5 mm. In the excellent
figures of C. radians given by Dr. Nicholson in his " Tabulate
Corals,"; those of the Russian Carboniferous specimens show a
much thicker calicular wall than in C. stelliformis, whilst the
» Oiyct. de Gouv. de Moscou, 1837, p. 160, PL xxxvi., fig. 6. See also
Lonsdale, in Murchison, Verneuil and Keyserling, Geol. Russia in
Europe, Vol. i., 1845, p.595, PI. A, fig.9.
t Favosites depressns Fleming, Brit. Anim., 1828, p. 529. F. capillar is
John Phillips, Geol. Yorkshire, 2nd Pt., 1836, p. 200, PI. iii., figs. 3-5.
Alveolites clepressa Fleming sp., Edwards and Haime, Mon. Brit. Foss.
Corals, Pt. iii.; Mon. Pal. Soc, 1852, p. 158, PI. xlv., figs. 4, 4a.
:*; Supra cit., PI. xii., figs. 4, ^a-d.
29
394 NOTK ON A NEW SPECIES OF CHiETETES.
tabula? are remarkably regular and horizontal, as distinguished
from the irregular and strongly curved tabulae in the Australian
Devonian species. On the same Plate, Nicholson has figured
(fig. 4c,d) sections of a coral from the Carboniferous of Shap,
Westmoreland, which he refers to C. radians, and which ap-
proaches the Australian species in its thin walls and more con-
spicuous projections.
EXPLANATION OF PLATE XLII.
Chcetetefi stelUformif^, sp.nov.
Fig. 1. — Exterior of fractured surface of specimen, showing small size of
corallites, their gentlj^ curved habit and interrupted platforms of
tabulse; 1| nat. size.
Fig. 2. — Transverse section of corallum, showing strong development of
partitions and occasional fission of corallites; ( x 16).
Fig. 3. — Longitudinal section, showing complex nature of wall, curved
tabulse and longitudinal partitions; ( x 16).
395
STUDIES IN AUSTRALIAN MECOPTERA.
No. ii. The Wing-venation of Chorista australis Klug.
By R. J. TiLLYARD, M.A., D.Sc, F.L.S., F.E.S., Linnean
Macleay Fellow of the Society in Zoology.
(Plate xliii.; and four Text-figures).
In collecting the evidence required for my work upon the
Panorpoid Complex, it was found necessary to examine the pupal
tracheation of examples of all the Orders there studied. The
only Order in which this had not yet been done (for at least one
example of the Order) was the Mecoptera; in which so little is
known of the life-histories, that it appears that only two observers,
Brauer in Austria and Miyake in Japan, have ever succeeded
in following up the complete life-history of any single species.
Neither of these authors was seized with the importance of
making a study of the pupal wing-tracheation, although it is
evident that the opportunity was offered to both of them."*^ The
work of both was carried out upon the genus Panorpa, which is
abundant enough in the Northern Hemisphere, but does not
occur in Australia.
It was therefore necessary for me to set about the discovery
of the pupa of some Australian representative of the Order. The
only species that is at all common round Sydney is Harpohittacus
tiUyardi Esb -Pet. But this belongs to a family that is, in man}''
ways, the most highly specialised of all the Mecoptera; whereas,
for my purposes, it was clear that an archaic representative of
the Order was to be preferred. I therefore decided to follow up
the life-history of the rare Chorista australis Klug, of which I
* Brauer's work was done long before theComstock-Needham Theorj' of
Venation saw the light. Mij^ake's work is of recent date, but its objective
was not venational, and the pupal wings were not examined.
396 STUDIES IN AUSTRALIAN MECOPTERA, ii.,
had obtained only about seven specimens in the course of as
many years.
Last year, I had the good fortune to find this rare species
fairly commonly, for the first time since I had begun collecting
insects around Sydney (about fourteen years). More fortunately
still, the area in which they occurred was a very restricted one,
only about a quarter of a mile from my house at Hornsby. In
this locality, I took, in the course of three weeks' collecting in
April, 1917, nearly one hundred specimens of this species.
Several of the females laid eggs, but unfortunately none of these
were fertile. I therefore watched carefully to see where the
females appeared to be ovipositing, and determined to select a
small area where there would seem to be a probability of finding
the larv?e in February and March of this year.
As nothing is know^n of the life-history of this species, I worked
upon the supposition that its larva would behave much as that
of Panorjm; i.e., that it would become rapidly full-fed, and then
burrow into the ground and remain there for the greater part of
the year, only pupating a week or two before the emergence of
the imago. This supposition proved to be fairly correct.
Digging and sifting of the soil in which the larva was expected
to occur was begun in February and continued into March. The
details of this arduous but interesting work are best left to be
given in a full account of the life-history, which I hope to pub-
lish later on. Meanwhile, it will be sufiicient to say that, one
day in March, freshly turned pup?e were obtained, and that these
were at once made use of for the purposes of this paper.
Owing to the shape of the pupa, and the direction of the wing-
sheaths, which are laid more parallel to the abdomen than is usual
in pupse, it was not found practicable to use the method adopted
previously in the case of Chryso'pa. It \vas necessary to separate
the wings from one another carefully, and to cut them off at their
extreme bases with sharp scissors. As the wings are excessively
delicate, this was not easy to do successfully. The photomicro-
graphs obtained show how the delicate wings became creased
near their middles, in the endeavour tu separate the glued ex-
fiY R. J. TILLYARb. 397
tremities; and also how, in the forewing, a portion of the fat-
body, lodged in the extreme base of the wing, got carried over
on to the wing wjien the cover-slip was let down upon it. Apart
from this, the residts obtained were quite satisfactory.
The Mecoptera are generally regarded as being an extremely
archaic Order of Insects; and I'ightly so, for both in their general
structure and in their Palpeontological record, the evidence tends
to emphasise the fact that they are quite as old as any existing
Holometabola, if not older. Within the Order, the genus
Chorista is one of the most archaic types still extan^, and shows
close relationship to forms found in the Permian. I was there-
fore quite prepared, and indeed expectant, to find a very primi-
tive and complete tracheational development in the pupal wings
of this insect; and I was relying upon this to solve certain
problems concerning the fusion of some of the main veins near
the base of the wing, which cannot be determined with certainty
from the imaginal venation. But the result obtained was quite
different from what was expected; and, if there is any truth at
all in the Comstock-Needham Theory of Wing- Venation, we must
regard Chorista as a very highly specialised type in this one par-
ticular, if not in any other.
The facts of the case can be gathered very readily from a
reference to the photomicrographs in Plate xliii., as well as the
drawings in Text-fig. 1. There are only tioo inain trachea' enter-
ing the winy-radiment. One of these belongs to the costo-radial
group, and passes into the main stem of the radius. In the fore-
wing, after giving off a very short and slender branch to R^, it
passes on along the radial sector. At the forking of Rs into
R2+3 and R4-f-5, it follows the upper branch, giving oft' a fairly
strong branch-trachea along the lower. Arriving at the forking
of R.^ from R3, it bends weakly down into the latter, and does not
reach the tip of the wing. The lower branch of this trachea passes
along Ri^o into R5, giving oft" a short branch to R4, and finally
ends up by bending down into Rgt- In the hindwing, there is
only a minute vestige of the branch into Rj. The main trachea
passes on into Rs; arriving at the fork of the sector, it gives oft'
398
STtJDlES In AUSTRALIAN *IECOPTERA, ii.
only a weak branch into R2+35 and passes on into R44-5. Branches
of about equal strength are given off to both R4 and Rg. The
former passes up along a cross- vein into R^, while the latter per-
forms a similar evolution nearer to the apex of the wing, ending
up in R4.
lA Cuj Ca, M4b l^+a
w
• Text-fii
ings of a pupa of Ghorista auKtralis
King, 9, freshly-tunied; ( x21).
Senii-diagranimatic, the tracheation represented by continuous black
lines, the imaginal venation b^^ dotted lines; k, the point at which
Cuj meets a short cross-vein from M, eventually fusing with M in the
imaginal venation : ^[t, median tiacliea; Rf^ radial trachea. For rest
of lettering, see p. 408.
The second main trachea of the wing is of smaller calibre, and
belongs to the cubito-anal group. It enters the wing-rudiment
along the media. In the forewing, it is quite unbranched, and
passes finally into M3 in an almost straight line. In the hind-
wing, it gives off short branches to the cubital fork and also to
M3_^4, and its distal end passes into a cross-vein below M.^.
A second pupa was examined and its wings photographed, but
the negatives obtained were not very satisfactor3\ In this case,
BY R. J. TILLYARD. 399
the distal endings of some of the tracheje were different from those
recorded above; e.g.^ tlie trachea in R.j of the forewing passed
down into R4 via a cross-vein. The median trachea of the fore-
wing also gave out a short branch to the cubital fork.
Only two pupje were available for study. There can be little
doubt that further minor variations would have been found had
a longer series of puppe been examined.
There is only one conclusion to be come to from this result.
It is, that the Mecoptera, as exemplified by the archaic genus
Cho7'ista — and, therefore, presumably, by other existing genera —
are highly specialised as an Order along the same lines that we
find in the Hymenoptera, Trichoptera, and Diptera, viz., by the
reduction of their wing-tracheation. The cause of this reduction
is almost certainly the same in all four Orders, viz., that the
trachese do not grow into the wing-rudiment until the latter has
been fully formed, with the venational scheme completely laid
down. In such a case, there is no longer any need for a trachea-
tional scheme, to guide in the laying down of the venational
scheme originally based upon it; the only necessity is to supply
the wing with oxygen. Consequently, there will be a tendency
to reduce the tracheal supply to the minimum necessary for this
purpose; and, also, for the tracheae to cease to follow their
original paths, and to take instead the path of least resistance.
Both these tendencies are well illustrated by the case here
studied.
The same tendency, with many stages still preserved to us, has
been noted already in the Order Plectoptera; an Order which,
curiously enough, approaches most closely to the Holometabola,
in that it possesses, in its sub-imaginal stage, what appears to
have been Nature's first attempt to evolve a true resting-stage or
pupa. The pupa of the archaic Mecoptera closely resembles the
imago in everything except tlie form of its mouth-parts and the
non- expansion of its wings. Thus it only differs from the sub-
imago of the May-flies in being unable to fly, and in slightly less
mobility of its free appendages. In the Order Plectoptera, there
are a number of genera known in which the remaining wing-
400 STUDIES iX AUStRALlAS' MECOPTERA, ii.,
ti'aclieEe all come off from a single stem. This must be regarded
as a higher specialisation than that found in the Mecoptera, or
in any Holometabolous Ordei'. For, in all of these, it would
appear that tu-o is the minimum number of wing-tracheae yet
reached in the course of evolution, i.e., one from each of the main
tracheal groups.
It is interesting to compare the courses of these two trachea^
in the four Holometabolous Orders which show reduction. Of
the four, the only one which retains the media intact and separate
from the base onwards is the Mecoptera. In this Order, the
trachea belonging to the cubito-anal group passes into the media,
which is as far forward as it could possibly get. This shows
clearly that, in the ancestral form of the Mecoptera, the median
trachea had already become attached to the cubito-anal group, as
it has in most Orders of Insects. In the Hymenoptera, the media
is suppressed in the imaginal venation, and only a bare vestige
of its trachea is to be seen in a very ancient family, the Siricidce.
The principal trachea of the cubito-anal group passes along the
cubitus, and may or may not be accompanied by a separate anal
trachea behnv it. In the highest forms, this latter becomes fused
with the cubital trachea; so that the condition of two trachea?
only, in the wing-rudiment, is there fully reached. In the
Trichoptera and Diptera, in which the media is fused basally
with either the radius or the cubitus for a greater or less dis-
tance, the trachea of the cubito-anal group passes likewise along
the cubitus.
From a study of this character only, then, it is clear that the
Mecoptera present a more archaic stage than that seen in the
Trichoptera and Diptera; and hence there is no evidence here
against the presumption that both these Orders are derived from
the older Mecoptera, as Handlirsch supposed. The Hymenoptera,
on the other hand, cannot be derived from the Mecoptei-a, since
they show a more archaic stage in the Siricida' and some other
families (where more than two trachea3 are still present), even
though they stand far in advance of the Mecoptera in having the
media eliminated from their venational scheme.
BY R. J. TlLLYARlJ. 401
Turning now to the Orders Lepidoptera and Planipennia, we
see that both of tliese preserve the complete set of tracheie in the
pupal wing. In this character, then, they are both of them
much more archaic than the Mecoptera and the other three
Orders discussed above. It must be clear, therefore, on this
character alone, that the Lepidoptera cannot be descended either
from the Mecoptera, as Handlirsch supposed, or from the Tri-
choptera, as Meyrick would have it.
Let us now return to the question of the interpretation of the
imaginal wing-venation in Chorista. The tracheation having
failed us, what other evidence have we to go upon in working
this out?
First of all, it may be stated that the main scheme of venation
in Chorista^ like that of the Trichoptera, is so primitive that
there can be no hesitation in naming the veins, even though we
cannot call upon the trachea? as evidence. The subcosta, the
radius, the radial sector with its dichotomous branches, and the
anal veins can all be named without any misgivings. It is only
the media and cubitus that offer a difficult problem. By re-
ferring to Text-fig. 1, it will be seen that the main stem of the
media, in both fore- and hind wings, is continued straight on by
a strongly convex vein right to the wing-border. All the rest
of the media lies anteriorly to this. The media being properly
a concave vein, it follows that there is at once a strong supposi-
tion that this straight prolongation of the media is really Cuj,
and that the result has been brought about by a fusion of this
latter vein, not far from its original at the cubital fork, witli the
main stem of the media.
If we examine the imaginal venation carefully, we shall see at
once strong evidence in favour of this view. Picking up the
base of the cubitus, which is a very weakly chitinised vein, and
following it to the cubital fork (en/), we see that the apparent
cross-vein from this point to the media may very well be the
basal portion of Cuj, arching up to unite with the media at the
point k; and then leaving it again as the straight, strong, convex
vein in question. If that be so, then Cuo hi the forewing is the
402
STUDIES IX AUSTRALIAN MECOPTERA, II.
vein that continues the hue ut" the stem of Cu; while, in the
liimlwing, it must fuse with lA for a short distance, and then
separate from it again towards the wing-margin.
We must now ask for definite proof that this supposition is
the correct one. That proof is forthcoming from two separate
sources, viz., the pupal wing-venation, and the imaginal wing-
venation. We may take these two separately.
(1) Evidence from the papal wing-venatwii: — In the freshly
turned pupa, less than one day old, the fusions of veins that take
place in the imaginal venation are not completed, but only just
beginning. (The photomicrographs in Plate xliii., are from a
pupa at least three days old; those taken from a freshly turned
pupa showed the separate veins much more distinctly, but unfor-
tunately the negatives were partially spoilt through the use of
stale developer, and are not good enough to reproduce).
Text-fig. 2.
Basal part of hindMing of same pupa as in Text-fig. 1, { x 43), to show the
radial {Rt), and median {Mt) tracheie, the approach of Cui to M, and
the partial fusion of Cuo with lA.
Text-fig. 2 shows a camera-lucida drawing of the basal part of
the hind wing of a freshly-turned pupa. In this, the separate
veins Cu., and lA can be clearly seen running alongside one
another for some distance, and then diverging again towards the
wing-margin. This fixes the position of cuf dX the point already
BY R. J. llLLYARD.
403
selected. The upwardly arching branch above ctif is seen to be
connected witli the media by a i<hort crobS-rcAn, descending from
the latter vein. In this pupal tracheation, then, the line of the
median stem is not continued by Cui, but by the true media; it
is only a rearrangement of the positions of these parts in the
imaginal venation that brings Cuj into the position of continuing
the line of M.^
The case for the forewing is still simpler. Here there is no
fusion of Cuo with lA, and the parts of the cubitus can be seen
to be quite separate from both M and lA.
{'2) Evidence from the imaginal venation: — The evidence that I
propose to bring forward
here is of quite a novel
kind, and does not ap-
pear to have been used
hitherto in any research
upon wing- venation; yet
it is of the utmost vahie.
In the wings of all
Holometabolous Insects,
two kinds of hairs or j /-x-
setyy are developed. \ L-^L_^___ 2A.
These I have called ma-
crotrichia and mic7'otri- Text-tig.o.
chia respectively, t The Part of the imaginal venation of the forewing
former are in most ^^ Chorista australis Kkig, in the region of
..,-,, ,1 the cubital fork (c?//), to show distribution
cases, restricted to the „ ^, - • i- / .,^\ i 4-\ ■ *- ^
' of the niacrotnchia; ( x 27) : k, the ponit at
veins; the latter are which Cui has fused with M; .r, cross- veins.
spread indiscriminately,
and much more abundantly, over the whole of the w ing. In the
Mecoptera, the macrotrichia are found upon all the main veins
and their branches, but never wpott the true cross-veins. Hence
'"■ Compare the somewhat analogous condition in the forewing of Myr-
meleontidte, where Cuia captures Mo close to its origin.
t "Mesozoic Insects of Queensland, No.l.'' These Proceedings, 1917,
xHi., Part 1, p. 195.
404 STUDIES IN AUSTRALIAN MECOPTERA, ii.,
we have, here, a fine criterion for determining whether any sup-
posed cross-vein is really such, or not.
Let us apply this to the basal portion of Cuj, which, arching
up in the imaginal wing-\ enation to meet M, might well be mis-
taken for a cross- vein. From Text-fig. -3, we see that all the
cross-veins around it are destitute of macrotrichia. Not so this
vein, however; for it is seen to carry a series of closely-set macro-
trichia, which are continued along the rest of Cuj.
Further, it will be seen that the macrotrichia de\eloped upon
the media are less in number than those upon the cubitus, for
any given length; those upon the cubitus being about twice as
luunerous. In this character also, the continuity of Cuj from
cuf'u]) to its fusion with M, and thence along the strong, con%ex
vein already mentioned, is clearly indicated.
Putting the two sets of evidence together, there can be no
doubt that the correct interpretation of the condition of the
media and cubitus is that which is given in the Text-figures.
Descrijytion of the Wing-Venation. (Text-fig. 4).
Having solved the above problem, we can now name all parts
of the wing-venation without any doubt whatever. The result
is shown in Text-fig. 4. It will be seen that the wing-\'enational
type is essentially similar to that of the Lepidoptera, Trichoptera,
and Diptera, except for the fact that a considerable number of
unspecialised cross-veins are developed as supports between the
main veins. These cross-veins cannot be regarded as part of
the original venation, since they do not carry macrotrichia, and
are absent or very weakly developed in the oldest genera, both
fossil and recent.
The Sabcosta {Sc) is a straight \ ein terminating about halfway
along the costal margin in the hindwing, somewhat beyond half-
way in the forewing. In Chorista, it gives off only a single
cross-vein in the costal space, near the base (occasionally, as in
Text-fig. 4, a second near its distal end). This is the homologue
of the humeral cross-vein in Lepidoptera and Trichoptera, and is
a true cross-vein, since it does not carry macrotrichia.
BY R
TILLYARD.
^A
in
M
\
405
406 STUDIES IN AUSTRALIAN MECOPTERA, ii.,
Tlie Radius {R) is a strong convex vein branching at the
radial fork {rf) into Rj and Rs. The main stem, Rj, reaches
the wing-margin well before the apex. Around its distal end is
formed the pierostigma {'pt), a hardened membranous area
bounded by the costal margin above and the pApvostuimatic furrow
{ptf) beneath. Within this area are sometimes developed one
or more ptpvostiy mafic vpinhfs, which are also parts of the
original venation, not cross-veins, but true branches of Rj, since
they carry macrotrichia. The Radial St^cfor (Rs) has the typical
dichotomous branching seen in the Trichoptera, giving rise, in
each wing, to four branches, R.,, R3, R4, and R., respectively.
The separation of R., from R;. takes place far distad, under the
pterostigma; that of R^ from R5 much further basad, not far
from the first forking of Rs into R2-1-3 and R4_|^5. All these
branches are connected and strengthened b}' a number of cross-
veins, whose position varies greatly in different individuals.
The .]ffidia (M) resembles the radial sector in the manner of
its branching, as in Trichoptera; except that, in the forewing of
Chorista, the most posterior of its four branches, M^, divides
into two parts, M.^^ and Mj,^. The cross-vein connecting M^ near
its base with Cuj carries no macrotrichia. and must, therefore,
be regarded as a true cross-vein, and not a branchlet from Cu^ .
Hence there is no evidence for the existence of the two branches
of Cuj, which I have called Cuj^ and Cun,, and which are to be
found in the most archaic Lepidoptera and in the Planipennia,
but not in the Trichoptera.
The Cubitus (Cu) is two-branched, as in Trichoptera. The
anterior branch, Cuj, arches upwards to connect with M, and
then continues the line of M to the wing-margin as a strong,
convex vein. The posterior branch, Cua, in the forewing, simply
continues the line of the main stem of Cu. In the hindwing,
however, it fuses for some distance with lA, and then separates
from it, and runs beneath Cu, to the wing-margin, at about half-
way along the wing.
Three Anal Veins are developed in each wing. These remain
BY R. J. TILLYARD. 407
quite distinct throughout theii- courses, except for the partial
fusion of 1 A with Cuo ah-eady mentioned in the liindwing.
A number of cross-veins, irregularh' placed and excessively
variable in different individuals, are developed as supports
between the branches of the media, the cubitus, and the anal
veins.
It should be noted that, in the imaginal venation, the main
stem of the media is fused with the radius for a short distance.
In the pupal venation, these can be seen to be quite distinct.
►Summary.
(1) The tracheation of the pupal wing of Chov'if^ia aia^t rails is
highly specialised by reduction, there being onl}- two main
trachete entering the wing. Of these, one belongs to the costo-
radial group and passes into the radius; the other belongs to the
cubito-anal group and passes into the media.
(2) In the freshly-turned pupa, the fusions of veins that are
found in the imaginal venation are not accomplished, and hence
it is possible to interpret the whole of the venation with certainty.
(3) Main veins and their branches, in the imaginal \ enation,
carry macrotrichia; cross-veins do not. From this, it is de-
monstrable that an apparent cross-vein below M is in reality the
basal portion of Cuj.
(4) The imaginal venation is specialised in having M fused
basally with R for a short distance; in having Cu, continuing
the line of the main stem of M, the short basal portion of Cuj
resembling a cross-vein below M; and also in the partial fusion
of Cug with lA in the hind wing.
(5) The rest of the venation is of a generalised type, closelj"
resembling the typical Trichopterous plan, but with an extra
branch to M4 in the forewing. Cross-veins are fairly numerous,
but variable in position and number in difterent individuals.
408 STUDIES IN AUSTRALIAN MECOPTERA, ii.
EXPLANATION OF PLATE XLIII.
Fitf. I. — Wings of a pupa of Chorista aujif m/ is Klu^r, 9, three cla3's old;
(xl8).
Fig.2. — Basal part of forewing of same; ( x 48). Note the Alar Trunk-
Trachea at extreme base of wing, giving off the two wing-traehea?.
Fig.3. — Basal part of hindwing of same; ( x 32).
(Photomicrographs of tlie freshly dissected wings in water).
Explanation of CoMSTOfK-NEEDHv^M Venational Notation, as t^sed
IN the Text-fk hires.
lA, 2A, 3 A, the three anal veins — C, eosta — Cu, cubitus; Cuj its upper,
and Cu2 its lower branch — ruf, cubital fork— ^/V, frenulum — M, media;
Mj, M„, M3, M4, its four branches, the last of which divides into ^1^ and
M4b in the forewing —;??/, median fork — pf, pterostigma — ptf, pterostig-
matic furrow — R, radius, dividing into Rj, its main stem, and Rs, the
radial sector; this latter has four branches. R.,, R„. R^. and Rg — /•/", radial
fork — rf, fork of radial sector — He, subcosta.
409
THE "SPRINGING" OF TINS OF PRESERVED FRUIT.
By W. W. L'Estrange and R, Greig-Simith, D.Sc, Macleay
Bacteriologist to the Society.
It occasionally happens in the canning of fruit that some of
the tins, after a time, begin to swell. -Normally, the top and
bottom of the container are slightlj^ concave, indicating a partial
vacuum within If, however, one or both is convex, the tin is
abnormal and is alluded to as a "springer." Such cans are
rejected, and, as there may be many sprung tins, the loss may be
considerable to the canner. The cause is generally ascribed to
leaking containers, but as some fruits are more prone to produce
" springers " than others, it appeared that much might be gained
from a study of the phenomenon. It is certainly curious that,
while pears and some varieties of plums with tight stones are
frequently aftected in this way, peaches and other stone-fruits
are but rarely so, even when they are processed or manufactured
in precisely the same way. Our attention was called to the
matter, and an examination was made of some of the affected
tins.
But before dealing with the results of this examination, it may
be well to briefly describe the method of processing the fruit.
The pears, generally the Bartlett variety, are picked, pared, cut
in two, cored, in some cases sliced, and washed in running water.
They are then put into empty containers, and placed on a
travelling belt which carries them past a tap from which boiling
syrup is run in up to within a quarter or half-an-inch of the top.
They pass on to a machine which automatically fits on the lids
and turns the edges. The cold pears reduce the temperature of
the syrup, which is generally at lbO°F., when the lid is fixed
and the container made tight. The tins are put into baskets
and passed through a boiler. The passage occupies 24 minutes,
and the tins are in the boiling water zone for 16 minutes. On
30
410
THE SPRINGING OF TINS OF PRESERVED FRUIT,
emerging from the boiler, the ends are bulged, but they become
concave as the tins cool, while stacked on the floor near the
boiler. When cold, they are taken to the store, and built up in
tiers so that any springers may be detected. After a time, they
are labelled and despatched.
The springers appear to be of two kinds. One contains syrup
in an actively fermenting condition: the other is quite still, but
the syrup has a sharp, slightly acid taste noticeable only to the
expert taster, and the fruit is apparently quite fit for human
consumption. The fact that one or both ends of the container
are bulged is, however, sufficient to cause the condemnation of
the tin.
The time during which the fruit is cooked seems to he suffi-
cient to thoroughly pasteurise the (contents of the tins, but to
gain some definite information upon the matter, two tests were
made in the laboratory with fresh pears. These were peeled,
halved, and cored. 'J'he bulb of a thermometer was inserted in
the body of one of the halves and the tin was filled up with boil-
ing syrup, and put into boiling water. The temperatures of the
pear and of the syrup were read off' at intervals.
At start
2 minutes
5 minutes
10 minutes
15 minutes
20 minutes
24 minutes
Temperature in °C. of the
pear
syrup
18°
27°
42-5'
65-5'
78°
85°
89°
81°
84°
87°
90-
92°
94-
95°
Container taken out and placed on bench
28 minutes
90°
87-5°
30 minutes
90°
85-5°
35 minutes
88°
78-5°
40 minutes
85°
74°
The conditions were not quite the same as in processing under
factory-conditions, inasmuch as there was no lid on the con-
tainer, and it was not completely submerged in the boiling water.
BY W. VV. L ESTRANGE AND R. GREIG-SMITH.
411
For this reason, these temperatures are k)wer than would occur
in practice, but even as, in the experiment, the pears and the
syrup were over 70°C. for at least 30 minutes, and over 85* for
20 minutes, the exposure was quite sufficient to pasteurise the
contents
One point about the experiment was, that the tin was put
into the boiling water immediately the syrup was added; whereas
in practice, a certain time must elapse before the tin is capped,
put into a basket and pushed into the boiler, where it is some
four minutes before the zone of boiling water is reached. It was
considered that an interval of ten minutes would completely
cover this interval, and, accordingly, in another laboratory-test,
the container was allowed to stand for ten minutes after receiving
the boiling syrup, and before it was put into the boiling water.
Temperature in ""C. of the
pear
syrup
At start
5 minutes
10 minutes
15 minutes
20 minutes
24 minutes
45°
50°
67°
76°
80-5°
83°
47°
67-5°
80°
85°
88°
89-5°
Container taken out and placed on bench
28 minutes
84°
87-5°
30 minutes
83-5°
86°
40 minutes
79°
78°
Even with the ten-minute interval before placing the tin in
the boiling water, the fruit and the syrup were well over 70° for
at least 25 minutes, and this should be enough to destroy all
vegetating yeasts and bacteria.
A number of faulty tins, ten in all, were received in October,
1916. They consisted of pears, greengages, and plums. They
were examined bacteriologically, and no growths were obtained
from the contents of seven. A tin of pears contained a small
active yeast, Saccharoinyces Zopfii, which has been shown to be
responsible for the "puffing" of tins of golden syrup in America.
412 THE SPRINGING OP TINS OF PRESERVED FRUIT,
It actively fermented fruit-syrup, and was undoubtedly the cause
of the springing of this particular tin. Another tin of pears
contained a mixture of inactive yeasts and moulds. As the}'
did not ferment fruit-syrup, it was clear that the cause of the
trouble had disappeared. A tin of greengages contained inactive
bacteria and moulds, and, as in the previous case, the agent
which had caused the fermentation had died or had been killed.
Several of the containers showed clear evidence of having leaked
at some time.
A second lot of six faulty tins of pears was obtained in June,
1917, and the notes upon these are as follows.
No.l. The container had a faulty lid, the tinplate was spong\',
and foaming syrup was oozing out of a central pin-hole. 'J'he
syrup was actively fermenting and contained yeasts.
No.2, The container had a small quantity of syrup, the bulk
having apparently leaked out. The syrup contained yeasts and,
when set aside under aseptic conditions, it fermented vigorously.
No. 3. The contents appeared to be normal, but the syrup con-
tained bacteria and yeasts.
No. 4. The fruit appeared normal, but the syrup was thin, and
contained bacteria and yeasts.
Nos.5 and 6. The contents were normal, and the syrup con-
tained bacteria only.
All the containers, with the exception of No. 6, showed signs
of having leaked at some time. No. 6 was the only one that ap
peared to be intact.
The bacteria in the tins were the same. On dextrose ajjar,
they grew as short, irregular rods, and hs chains, especially in
the condensed water. They were Gram-positive, and, as they
had no action upon any sugar, they could not have been respon-
sible for the springing of the retainers.
The yeasts w^ere all of the culture-type, that is, they were
varieties of Saccharomyces cereinsice. This type is associated
with the manufacture of beer, and one would not expect to find
it in a factory where there is so much fruit utilised. In a factory
such as a fruit-cannery, wild yeasts and torulse should be present
in abundance, and it certainly was extraordinary to find only
BY W. W. i/eSTRANGE AND R. GREfG-SMlTH. 413
culture- or beer-yeasts in the preserved pears. It was subse-
quently discovered that, adjoining the cannery, there is a factory,
where a large amount of ginger-beer is produced. It is safe to
conclude that the yeasts came from this source.
In October, 1917, a third lot of six tins of pears was examined,
and, with one exception, all of them showed signs of leakage.
The contents were sterile, and, from the presence of soldered
vent-holes on two of the containers, it was concluded that the
lot had been reprocessed, that is to say, they had been put
through the boiler after evidence of springing had been detected.
One of the tins contained a strongly foaming, sterile syrup, and
as this was oozing from one of the ends, it was evident that the
re-processing had been of recent date.
We had an opportunity of inspecting the processing of pears
in February, 1918, and, from our observations, we concluded
that the root of the trouble lay with the closing of the containeis.
l he margin of the lid is painted with a mixture containing Hour
or starch, and when the closing machine overlaps the edges of
lid and container-top, the mixture fills up the spaces between
the interlocked edges. The joint appears to be tight, and doubt-
less the starchy mixture makes a good lute. But the pressure
exerted by the air contained in the tin (it constitutes from one-
twelfth to one twenty-fourth of the contents) must be consider-
able during the passage of the container through the boiler. The
luting mixture is supposed to form a jelly in contact with the
hot water, on the one side, and the hot syrup on the other, and
it is quite reasonable to suppose that the jelly may be forced out
of the joint by the pressure of the contents. The tins appear to
be sound immediately after processing: it is when they are
stacked in the store-room, that they show signs of leakage. In
the case of pears, about every second tin leaks more or less;
with plums, about every tenth tin is faulty; and, with peaches
and apricots, the leaks are few in number. As the processing is
the same in all cases, one is driven to the conclusion that the
juice of pears and, in less degree, that of plums, has a solvent
action upon the luting jelly, and that another mixture should be
employed.
414 THE SPKIVGING OF TINS OF PRESERVED FRUIT.
The experimental evidence shows that the duration and tem-
peiature of the processing are suflBcient to absolutely free the
contents from active yeasts. Their entry into the containers is
most likely to occur while the tins are cooling down or shortly
after. It is reasonable to suppose that the original air of the
tin has been partly or entirely expelled during processing, and
the vacuum created during cooling draws in the air of the factory
with its suspended yeasts and other organisms. It is entirely a
matter of chance as to the number of organisms and the activity
of the organisms that may be in the indrawn air. There
may be none, or they may be inactive, and the tin will not leak.
There may be one or more, and, if these are active gas-producers,
the tin will " spring. ' It is quite possible that some of the tins
might be so imperfectly closed that they would leak under any
circumstances, but the scarcity of leaks and springers among the
tins of stone-fruits shows that this is of very rare occurrence.
The behaviour of the pears and plums, as compared with
apricots and peaches, points to the use of a more insoluble luting
material, and, failing this method of overcoming the trouble,
there is the alternative and probably better plan of allowing the
tins to cool after processing in a current of filtered, sterile air,
and, when cold, painting the joints with a lacquer- varnish. Thus
the entry of yeasts into the tins would be prevented during the
cooling, and the varnish would make them germ-tight and germ-
proof.
415
NOTES AND EXHIBITS.
Dr. Cleland exhibited the first four parts (VoL i.) of " Epi-
graph ie Medical e : Corpus I nscriptionum ad JVlediciiiam Hiolo-
giamque Spectantium," publie par Professeur Raphael Blanchard
(Paris, 1909-15).- Also specimens of Anestellorhina augu7' (Cct/li-
phora ocea'iiice), one of the Australian Blowflies troublesome to
sheep, bred from an Agaric {Pletirotus sp.) which had probably
been infected during exposure for two days in a room — the
interest of the exhibit being that the Hies had been reared on a
purely vegetable medium. «
Mr. Steel communicated the following Note on the abnormal
gizzard of a Muscovy duck, exhibited at the last Meeting {antea,
p. 361). "On subsequent dissection of the exhibit, it was found
that, while the organ was of the usual shape and dimensions ex-
ternally, the internal cavity was only about one-third of the
normal capacity, and the gizzard- walls correspondingly thick.
The vermiform appendage was hollow, and was tightly packed
with a hard, brown substance, which broke up on treatment
with hot sodium hydrate, but exhibited no traces of vegetable
tissue, nor of definite structure, being apparently a secretion
from the walls of the appendage. The appendage was completely
isolated from thu cavity of the gizzard by a layer of white, fibrous
tissue, about \ inch in thickness, this being the thickness of the
side-walls of the gizzard at the area of attachment. 'I he speci-
men has been placed in the Museum of the Department of
Veterinary Science, University of Sydney."
Mr. Fletcher, on behalf of Mr. Musson and himself, showed
seedlings of a number of species of plants, not Eucalypts or
Angophoras — including EUeocarpus cyaneus^ Leptospermum
JlavesceiLS, Callistemon phii/olius and C. saliynus, Melaleuca
nodosa and M. yenistifolia, Trrstania iieriifoiia^ Syiicarpia
laurina, Backhousia myrtifolia, Casnariiia sp., Santalum obtusi-
folium, and several not identified — which seem to be afflicted, in
a similar manner, with composite, shoot-bearing tumours.
The President offered a cordial welcome to Mr. R. VV. Bretnall,
a Soldier-Member recently returned after service abroad.
416
ORDINARY MONTHLY MEETING.
August 28th, 1918.
Professor H. G. Chapman, M.D., B.S., President, in the Chair.
Messrs. John Hopson, Junr , D^alkeith, Eccleston, N.8.W.;
and Frederick P. Dodd, Kuranda, Nortli Queensland, were
elected Ordinary Members of tlie Society.
The Donations and Exchanges received since the previous
Monthly Meeting (31st July, 1918), amounting to 2 Yols., 44
Parts or Nos., 7 Bulletins, 1 Report, and 5 Pamphlets, received
from 37 Societies, etc., and two private donors, were laid upon
the table
NOTES AND EXHIBITS.
Mr. ^Y. W. Froggatt showed a fine series of Coeeid galls on
Eucalypts, commonly know^n as Brachyscelid galls, attributable
to members of the genus Apiomorpha: a remarkal)le gall due to
Cynips on Live Oaks in California; and an undetermined, living
snake from Moree, N.S.W.
Mr. Fletcher exhibited specimens oi Melaleuca Deanei F.v.M.,
and photographs of the largest plants seen (about 8-9 feet high,
and 4-5 inches in diameter at the base) ; and specimens of
Petrophila sessilis Sieb. — two of the rarer plants of the Lane
Cove District, to be met with only in small, isolated groups.
The Secretary communicated a letter from the Dalley Branch
of the Australian Society of Patriots at Newcastle, advocating
the reservation of Pulbah Island in Lake Macquarie. Mr. J.
Mitchell, who had visited the island, offered some favourable
remarks. Further consideration was postponed.
[Printed otf, September 4th, 1918.J
e L.S N.S.W. 1913.
G> O
O,
./^
^
o\^^
O) Q Q
O
Fig 1. A slightly ohliquc sertioM tlir..uj;li a hw\ of S,H-roln erassifolia, to sliow .levelopinent of
glandiilar hairs.
Fig. 2. Section across sheathing base of a mature leaf, to show development of glandular and
clothing hairs.
B- ^
lO
S.N.S.W. 1918.
Vig. 5. Jugate (HepialiOiE), Figs. 6-8, Freuate (Sphimjids:) Wiug-coupliug Apparatus.
Q^
-fi)^
- V
3«^
1
i ^
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s,ki^
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-j^'
— " \C ' '■ ''r~-
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ir
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fmmsmm/mmm
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P L.S N.S.W. 1918.
Fig. 1. Zaphrentis sii>. u. i? subgen. uov.). Fig. 2. SpongophyUum, (?) sp. n.
Fig. 3. fci. (?) sp. n.
P.L.s N.s.w. -rei«.
Figs. 4-5. PhiUipsastrxaffrandis, sp. u. Fig-. 6. Zaphrentis si^.
Fig. 7. Favosites, sp. n.
PL.S N.S.W. 1918
Figs. 8-9. Stroiiiatoporella loombcrensis, sp. n.
1t.
Figs. 10-11. Stroinatoiiordla bensoni, sp. u.
.L.S.N.S.W. 1013.
Fi^si. VI 16. Acttno6t.-->iiia un^trulc, Pp. u.
.L.S.N.S.W. 1911
.iP
r
Ui^.'^
7
'k/i-
Furamiuifera fium Nemiiigiia Limesluue ; ( x 5-^) .
P.L.S.N.S W. 191S.
>- «
^<r^•
/^^"W^->\
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aSX
0
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^Yr\K
- hesr
ys 1HH
r.c. .',;/. ..f ,,;,ofn.
Foraiuiiiit'era from iS'eiuiimba Limestoue.
P.L.S.N.S.W.
I I
o o Ti he
fsa.>^>»<o5pi*ra
^^Psa-^vx-^ospb^ra.
i-\r. ,,/,ofn
.Sections of Xeiuiugha Liuiestoiie.
P L.S N.S.W. 1918.
F.C. phnfo.
Chu^tftt'S stcHijonius. tii>. u. ; Loomberali J^iiuesloue.
..S.N.S.W. 1918.
i'uiutl WiiJK-tiaelieutiuii vi Llan-iala oust ,<il i>i Kl
417
ME80Z0IC IN8ECT8 OF QUEENSLAND.
No. 3. Odonata and Protodonata.
By R. J. Tillyard, M.A., D.Sc, F.L.S., F.E.S., Linnean
Macleay Fkllow of the Society in Zoology.
(Plates xliv.-xlv., and Text-fionres 11-16).
Since the publication of No. 2 of this Series (4), Mr. Dunstan
has completed the delainination of the rock taken from the
Ipswich Fossil Bed, in the course of gathering together the
material forming the third collection (3) of these fine fossils. The
whole of this collection, with the single exception of the Coleop-
terous elytra, is now in my hands for study. Amongst the very
fine material received in the last consignment are two interesting
fossil Odonata, these being the only Dragonflies found at Ipswicli
since the discovery of Mesophlehia antinodalis Till., described \w
a former paper (2).
The former consignment also contained a very fine fossil, whicli
is of such unique structure that it is not easy to place it in any
of the accepted Orders, either fossil or recent. This fossil has
been studied b}^ me for a long time; and I have definitely con-
cluded that it is best placed in the Protodonata, of which it
must form a new Suborder. The present paper deals with this
fossil and the two Dragonflies above-mentioned.
Recently, Dr. A. B. Walkom, of the University of Queensland,
Brisbane, has published his thesis upon the Lower Mesozoic
Rocks of Queensland (5), in which he discusses very fully the age
of the Ipswich Beds. There can be very little doubt that his
placing of these beds as Upper Triassic is correct, and as such I
am content to accept them. It will be more and more evident,
as the description of the fossil insects proceeds, that they consist
of a definitely Mesozoic Fauna, not unlike that of the Lias of-
31
418 MESOZOIC INSECTS OF QUEENSLAND, iii.,
England, together with a number of older forms, apparently relics
of the old Carboniferous Fauna, such as we know from Commentr}^
in France. The three insects described in this Part illustrate
this intermingling of forms very clearly.
Order ODONATA.
Suborder Zygoptera.
Family LESTID^.
Subfamily TRIASSOLESTIN.E, n.subfam.
Wings with the bases strongly reduced, petiolate; the whole
of the area between Cu and the posterior margin, below the
quadrilateral, free of veins; and, hence, no anal bridge present.
Quadrilateral narrow, placed very obliquely to the long axis of
the wing; its upper side short, its lower long, and hence its distal
angle very acute; Cuj and Cuo arising together from this angle,
Arculus complete, with Mj.^ arising from the top of the quadri-
lateral.
Genus T r i a s s o l e s t e s, n.g. (Text-figs. 11,1 2/>).
A complete antenodal cross-vein in line with the arculus; (this
is probably the second of the series). Distal side of quadrilateral
continued straight to the posterior border of the wing, by an
oblique cross-vein. M^.o arching slightly upward as each branch
is given off; M^ arises about six cells distad from the level of the
quadrilateral; Ms arises from Mj.o about four cells further distad
than Mg. Discoidal field fairly wide, with M^ and Cuj slightly
divergent; a single row of cells to beyond the level of the origin
of Ms, followed further distad by two rows. Between Cuj and
Cuo, a single row of cells at first, to about level of origin of Ms.
Cug curves strongly do\vnwards distally to meet the posterior
border of the wing; between it and the wing-border, one row of
cells basally, increasing to two distally. [Anterior and distal
portions of the wing missing, including nodus and pterostigma].
Genotype, Trlassolestps epiophJehw'kdps^ n.sp. (Upper Tri-
assic, Ipswich, Queensland).
BY R. J. TILLYARD.
19
TrIASSOLESTES EPIOPHLEBIOIDES, n.sp. (Textfigs. 1 1, \2b).
A well-preserved fragment, 18-5 mm. long, greatest width (dis-
tally) 8-5 mm.; consisting of the greater portion of the basal half
of the wing, but without the extreme base. The anterior part,
including the costa, subcosta, and ladius, missing. The presei-^ed
portion includes the quadrilateral, the media and cubitus with
their branches, and the posterior border of the wing. The anal
vein appears to be completely fused \\ ith the latter.
Text-fig. 11.
Trias.'^olesfes epioph/ebio'ide-'^, n.g. et sp., (Triassie, Ipswich, Q.). preserved
portion of wing; ( x 5). Cui, Cui, branches of cubitus; M,.o. upper,
and M4, lower sector of arculus; Ms, Z3'gopterid sector: p, petiole;
q, quadrilateral.
The appearance of the fossil may be seen from Text-fig. 11. The
veins are all beautifully preserved, but unfortunately the wing
lies upon some plant-remains, which appear to have been hard
twigs or stems, and which have caused the fossil to rest upon a
very uneven surface; with the result that, in one place at least,
the wing has become crumpled or torn transversely, so that the
continuity of the main veins is badly broken. The veins can,
however, be followed across the break, and picked up on the
other side of it. Distally also, in the region of M1.2, the wing
lies upon an uneven surface, and the position of both main veins
and cross- veins is somewhat distorted.
Eliminating the effect of these inequalities, it is not a ditlicult
420
MESOZOIC INSECTS OF QUEENSLAND, 111.
task to restore the wing to its correct form, as far as it is pre-
served for lis. The result is to be seen in Text-fig. 1 2i, to whicli
I have added the probable courses of the veins that are missing
in the fossil. For purposes of comparison, I have shown also the
N M,.3 K^
Text-fig. 12.
(rt) Epiophhhia superstea (Selj's), (Recent, Japan), forewing; h\ region of
quadrilateral of hindwing of same. [h). Triasso/e-^fes epiophlehioide><,
n.g. et sp., (Triassic, Ipswich, Q.), restored wing, with missing parts
shown by dotted lines, (c) Synlestes ireyerni Selys, (Recent, Australia),
forewing. The basal portions of all three wings shown for comparison
with one another, (all x 4). Ab, anal bridge; C, costa; N, nodus; R,
radius; sq, subquadrangle. Rest of lettering as in Text-fig. 11.
corresponding areas of the forewings of U2^iophlebia superstps;
(Selys), and Synlestes tveyersi Selys, both of which show certain
points of resemblance with this interesting new fossil.
Comparing the fossil first of all with Rpiophlebia, we note that
BY R. J. TILLYARD.
421
the fossil wing was considerably broader, and proljably also some-
what longer. The region of the quadrilateral is closely similar
in both, but the quadrilateral of the fossil wing is placed more
transversely to the wing-axis, and the lower })ortion of the arculus
is much more transversely placed than in Eplophlebia. The
quadrilateral of the fossil resembles that of the forewing of
Ejnophlebia (Text-fig. 12a) nmch more than it does that of the
hindwing (6'); but, in other respects, particularly in the strong
distal curvature of Cuo, the fossil wing more resembles the hind-
wing. A close rese)nblance to E^nophlebia is shown in the levels
of the origins of the branches of M, and in tlie shape and vena-
tion of the discoidal field and the two longitudinal areas of the
wing lying below it.
The fossil, however, differs entirely from Epioph/ebia, and,
indeed, from all known Lestidie^ in possessing a broad and abso-
lutely free area between Cu and the posterior border, below the
quadrilateral. As the wing is excellently preserved in this
region, there can be no doubt about this character, which places
the fossil, in this respect, as more highly specialised by reduction
than any known Zygoptera, except only the subfamily Froto-
tieurince of the family Ayrionid(e.
Where the nodus was placed, we cannot tell for certain. But,
if we were agreed that the comparison with Ejnophlebia would
hold also for this character, we might expect that it was situated
somewhere near the extreme distally preserved portion of the
fossil, i.e., considerably distad from the level of the origin of Ms.
The comparison with Synlestfs is by no means so close, as can
be seen from Text-fig. 1 'Ic. The wing of Synlet^tps is exceedingly
narrow, and difiers from that of the fossil in having Ms arising
from Mg close to its origin, in the very narrow (juadrilateral, in
the upward arching of Cuj from the distal angle of the quadri-
lateral, in the consequent narrowing of the discoidal field, and in
the length and straightness of Cu . But, in its extreme reduc-
tion of the anal bridge (Ab), Synlestes shows the nearest existing
approach to the high specialisation seen in the fossil; and, on
this account, the comparison is of value.
422 MESOZOIC INSECTS OF QUEENSLAND, iu.,
While placing this fossil as more closely allied to Epiophfebia
than to any known Dragonfly, fossil or recent, it seems to me
that the characters exhibited by it require its elevation to sub-
family rank. But, if the area below the quadrilateral had pos-
sessed an anal bridge, I should have been content to place it in
the subfamily EpiojMehiiiuf.
The Lestid affinities of the fossil appear to me to be beyond
doubt. As it appears that some students of the Odonata have
been unwilling to accept my removal of Epiophlehia to the family
Lesticke^ I should now like to draw their attention to this fossil,
which was unknown at the time that I made this suggestion.
The combination of Epiophlebiine characters from the quadri-
lateral distad, with the extreme reduction of the base, which
characterises this fossil, should surely be regarded as a further
link in the chain of evidence for the Lestid affinities of
Epiophlebia.
Type, Specimen No. 205rt, (Coll. Queensland Geological
Survey).
Horizon: Upper Triassic, Ipswich, Q.
Odonata iNCEKTiE sedis.
Genus Perissophlebia, n.g. (Text-fig. 13).
Between C and R, beyond pterostigma, an irregular double row
of cells. Between R and Mj, beyond level of pterostigma, at
first an irregular triple row of cells, followed by a more regular
double row. Below Mj, a strong convex sector is developed
towards the apex of the wing, three rows of cellules from it.
This is probably Mja- Close below this sector, and only a single
row of cells' width from it, there is a parallel concave sector.
Both the above sectors converge towards M^ as they approacli
the apex of the wing. Some three or four rows of cells below
the concave sector, there is a divei-ging convex sector, which is
probably Mo.
[Only a fragment of the wing is preserved, embracing a por-
tion beyond the pterostigma].
Genotype, Perissophlebia midtiseriata, n.s^. (Upper Tri-
assic, Ipswich, Q.).
BY R. J. TILLYARD. 423
So little of the wing of this fine Dragonfly is preserved, that it
would be inadvisable to name it, were it not for the very remark-
able characters exhibited by the preserved portion of the wing.
It may be stated as a general rule, both for recent and fossil
Odonata, that the spaces between C and R, on the one hand,
and between R and Mj , on the other, from the level of the ptero-
stigma to the apex of the wing, are only filled by a single row of
cells. As exceptions to this, we may note the Liassic genus
Petrothnnis Handl., in which the distal portion of the space
between C and R is filled with three rows of minute cells, and
the basal portion of the space between R and Mj carries two
ro%\ s of cells near the pterostigma, decreasing to one row only
towards the apex; and the very densely-veined Aeschnidiidce, in
which the upper of the two spaces in question has only one row
of cells, but the lower has two.
Hence we see that, on this character alone, Perissophlehia
stands as the most densely veined of all known Odonata.
A second character, that is very exceptional in Odonata, is the
arrangement and direction of the longitudinal veins lying below
Mj. These are so strongly developed that they would appear, at
first sight, to be portions of main veins. But a little considera-
tion of the position <jf the pterostigma shows us that the pre-
served portion of the wing represents only a very small piece of
a very large wing. The lowest of the three sectors seen below
M] in the fossil is concave, and may very probably be M.^; the
other two are almost certainly not l)ranches of M, but intercalated
sectors. Of these latter, the upper one, being strongly convex,
very probably corresponds with the postnodal sectur, M,a, found
in recent Zygoptera and in Petcdura; while that below it might
be either a concave supporting sector, or a branch from Mo itself.
In any case, the directions of these three veins, two being parallel
to one another but converging towards Mj distally, and the third
being parallel to Mj, are quite exceptional in the Odonata.
It is clear then that, in spite of the small amount of the wing
preserved, the characters there shown are quite marked enough
to warrant the i-ivini; of a name to the fossil.
424
MESOZOIC INSECTS OF QUEENSLAND, 111.,
Pekissofhlebia multiseriata, 11. sp. (Text-fig. 13).
Total length of fi-agnieiit, 8*5 mm.; greatest breadth of same,
8*5 mm.
A small portion of the pterostigma preserved appears to point
definitely to the fact that it was sharply angulated at its upper
distal angle; a character that is exceptional amongst fossil
Odonata.
Most of the fragment is very well preserved, with the cross-
veins clearly visible; but, in the lower portion, they are partly
obliterated.
Text-%13.
Perissophlehia mnltheriata, ii.g. et sp., (Triassic, Ipswich, Q.K Preserved
portion of wing, ( x 7{.). Mi, postnodal sector.
The distance from the tip of the pterostigma to the apex of
the wing, and the great width of the wing beyond the level of
the pterostigma, both point to the fact that it is a fragment of
a very large wing, probably as large as, or larger than, that of
Petalura yigmUea Leach.
Type, Specimen No.203 «^7> (part and counterpart). (Coll.
(Queensland Geological Survey).
It is impossible to place this fragment definitely; but I would
BY K. J. TILLYAKD. 425
suggest a likely relationship with Fpfrofheinis ^iv<inlnris Haud-
lirsch, which is itself unplaced, owing to absence of the basal half
of the wing. A figure of this latter fossil is given in Handlirsch's
Atlas (1), Plate xlii., fig. 9. Jt is a much smaller wing than that
of Fei'itisoph/ehia, and comes from the Upper Lias of Dobl)ertin,
Mecklenberg.
Order PROTODONATA.
Suborder Aeroplanoptera, subordo nova.
Insects with strongly veined wings, traversed by numerous
parallel longitudinal veins, with a network of irregular cross-
veins, well developed near the base only; in the rest of the wing,
the cross-veins tend to become weak, and almost disappear dis-
tally. Costal border strongly developed, coriaceous at base.
Radius with a single sector. Media with many dichotomous
branches. Cubitus two-branched.
This Suborder is at once distinguished from all the rest of the
Protodonata (which may be termed the Suborder Meganeuro-
ptera, from the dominant family) by the close, parallel veining
of the wings, the dichotomous branchings of the media, and the
two-branched cubitus. In this latter character, it agrees with
the Order Odonata; whereas the rest of the Protodonata have
only a simple cubitus.
Family AEROPLANID^, fam.nov.
Wings excessively elongated and narrow, the parallel longitu-
dinal veins strongly developed and close together. Subcosta
weak, in a deep groove just in front of the exceedingly strongly
developed radius. The single radial sector arising from near the
base by a double root. Media seven-branched in both wings, but
the method of branching is different in fore- and hindwings.
Radius and media just fused at bases. Cubitus with its two
branches arising close to the base and diverging gradually. A
single anal vein present, below which is developed an anal field
with irregular veining, supported by a curved recuri-ent sup-
plement.
426 MESOZOIC INSECTS OF QUEENSLAND, Hi.,
Genus Aero plan a, n.g. (Plates xliv.-xlv., Text-figs. 14, 15).
Characters of the family, with the following additions. First
and second origins of Rs arising not far apart, the portion
between them somewhat curved concavely to R. First dichotomy
of M at same level as first origin of Rs. In forewing, the
anterior branch of M gives rise to thi-ee branches, by two dicho-
tomies which take place at some distance from the first; the pos-
terior branch also gives rise to three branches, which arise much
closer to the original forking. The middle of these three last
branches forks again, a little further distad. Thus there are
seven branches in all, three arising from the anterior branch, and
four from the posterior. In the hindwing, the anterior branch
of M remains simple for a considerable distance, but finally
divides into two only; the posterior branch, on the other hand,
soon divides again into two, of which the upper branch gives
rise to three branches by two separate dichotomies, the lower to
two only. Thus there are also seven branches of M in the hind-
wing, but of these only two arise from the anterior branch, and
five from the posterior.
The two branches of Cu diverge very slightly from one another
at first, both tending to diverge also from the radius, in the same
manner that the media does. A little further distad, close to
the level of the first origin of Rs, Cuo begins to curve strongly
away from Cuj, forming a wide bend, and later on again ap-
proaches Cu], to run parallel to and beneath it.
Just distad from the first dichotomy of M, in the hindwing
only, a weak, curved supplement runs back obliquely to join Cuj,
resembling in form the anal supplement below A, but on a
smaller scale.
(Jenotype, Aeroplana lulraljilis, n.sp. (Upper Triassic,
Ipswich, Q.).
Aeuoplana mirabilis, n.s}). (Plates xliv.-xlv.. Text-figs. 14, 15).
Total length of fossil as preserved, from base of hindwing to
end of isolated piece of forewing, 41-5 mm. Total length of
preserved portion of hindwing, 26mm.; greatest breadth of same,
5 mm.
BY R. J. TILLYARD.
427
The insect lies upon the somewhat uneven surface of a dark
grey shale, in such a position that it is crushed down somewhat
dorso-laterally, the hindwing apparently covering the thorax, and
having its costal margin passing under the posterior border of
the forewing. This latter wing, however, lies at a slightly higher
level; so that, in the process of delamination, its posterior border
was lifted off, exposing the whole basal portion of the costal
Text-fig U.
Ai'ropkuia mirabilis, ii.g. et sp., (Triassic, Ipswich, Q. ). Sketch of the
whole fossil in situ, ( x 2) to show the relative positions of the different
parts. A, A, principal portions of the fore- and hindwings, figured in
Plate xHv., fig 12; B, distal portion of forewing, figured in Text-fig.
lo. To the left of the wings are shown the crushed portions of the
head and thorax. The fossil remains are left unshaded, the rock
itself is lightlj' shaded, and the deep break between the parts A and
B is shown in black. The rock around B has been partly cut away
with a penknife.
margin of the hindwing. Around the bases of the wings there
are scattered a number of chitinous pieces, some belonging to
the head, and some to the thorax of the insect. These are so
much crushed that they cannot be recognised witli certainty; but
there are two very concave impressions close together, one circular
and the other an irregular oval, which may represent portions of
the compound eyes. Below the hindwing, there are two narrow
elongated fragments, one about 14 mm. long, the other about
428 MESOZOIC INSECTS OF QUEENSLAND, Hi.,
11 mm., which appear to represent tlie broken-off costal borders
of the other pair of wings.
]Mr. Dunstan tells me that, in the search for this fossil, the
rock broke transversely across the two wings, leaving a consider-
able gap between them. On the other side of this break, Mr.
Dunstan prospected with his penknife, and was fortunate in
picking up a further (more distal) portion of the forewing, lying
exactly in line with the more basal portion. This part was
glued into position on gardboard with the rest of the fossil, as
shown in Text-fig. 14. Mr. Dunstan assures me that this part is
not, at the most, more than a sixteenth of an inch out. If that
be so, as it certainly would appear to be from the shape of the
w4ng, then it is clear that the wings of this insect were exces-
sively long and narrow-, being in shape not unlike the planes of
a modern aeroplane. This character suggested to me the name
of the genus.
The general character of the wings of the fossil can be well
seen from Plate xliv., fig. 12. The hindwing is very complete as
regards its basal portion, and exhibits a large number of in-
teresting characters. We may most profitably study this wing-
first.
The costal space appears to be strongly convex, and was pro-
bably formed of very hard chitin, since the few cross-veins in it
are indistinct, and the whole area appears to have been coriaceous
in texture. The convex curvature of this area ends posteriorly
in the deep groove of the subcosta, behind which the radius rises
strongly up again as a convex \ein. The two separate bases of
Pvs are both strongly marked \eins, but the base of the media
appears as a very weak vein diverging from R. The bases of Cu
and A are strong veins.
In tlie whole of the basal region of the wing, up to a level
just beyond the second origin of Rs, the cross- veins are strongly
marked and easily followed. The meshwork here is of a typical
Paheodictyopterous type, the cells formed being very irregular in
shape. The development of the recurrent supplements between
M and Cu, and again between A and the posterior border, in the
BY R. J. TILLYARD.
429
hindwino-, are clearly interestino- attempts to strengthen the wino-
in the region of greatest curvature of the main veins.
Beyond the level of the origin of Rs, the ci-oss-veiiis liecome
weaker, and are ditHciilt to make out. In the figure in Plate
xliv., I have indicated every possible cross-vein of which I can
find traces, by examination in lights of different directions and
intensities. 8ome of these may not be true cross-veins, while
tliere may be others present whicli I have missed.
Text -fig. 1.-).
Ai'/'oplana nv!rahilis, n.g. et sp., (Triassic, Ipswich, Q.). Distal portion of
forewing beyond the transverse fracture of the rock shown in Text-fig.
The preserved part of the forewing is very similar to the hind,
except for the different manner of branching of the media,
which has been already mentioned. Careful measurements of
corresponding portions of fore- and hindwings show that theii-
widtlis are almost the same, the forewing being slightly wider
than the hind, as judged by the perpendicular distance between
Rs and Cuj. But this may have been offset by a somewhat wider
anal area in the hindwing.
The isolated distal portion of the forewing is shown in Text-
fig. 15. The main veins are well-preserved, yet there are few
indications of cross-veins. It seems probable that the cross-veins
became weaker and weaker towards the distal end of the win^-
and finally became obsolescent near the tip, so that most of them
are not preserved in the fossil. It will be seen that there are
430 MESOZOIC INSECTS OF QUEENSLAND, iii.,
two bifurcations in this distal part of the wing; these have to be
taken into account in the restoration of the fossil, which I have
attempted below.
Type, Specimen No. 126^. (Coll. Queensland Geological
Survey).
Horizon: Upper Triassic, Ipswdch, Q.
Restoration of the Wings ^>/ Aeroplana mirabilis, n.s]!.
(Plate xlv., figs. 13-1 4).
In attempting to restore the complete wings of this remarkable
fossil, I have made two assumptions which appear to be fairly
well justified, viz., that the shape and venation of the fore- and
hind wings were approximately the same (with the principal
differences preserved to us in the fossil as it stands), and that
the isolated portion of the forewing, shown in Text-fig. 15, lay
close to the apex of the wing.
Even with this latter proviso accepted, and taking full ad-
vantage of the possibility of an error of one-sixteenth of an inch
in Mr. Dunstan's placing of this fragment, it will be apparent
that the length of the wings, in comparison with their breadth,
is enormous, the ratio being approximately 7:1.
In filling in the gap between the basal half of the forewing
and the distal preserved portion, it is necessary to bear in mind
the possibility of one or more veins having reached the costal or
posterior border, respectively, in the interval. To determine
this, I examined the veins in the distal fragment carefully, with
the result that I found the uppermost to be more strongly built
than the others. This should, therefore, be the radius, as is also
evident from its level on the wing itself. Hence, if we assume
that there have been no other dichotomies of veins in the missing-
part of the wing, the veins below R in this part of the wing
must be, in consecutive order, Rs and the seven branches of M.
Of these last, it will be seen that M^ and M7 branch again. There
remain, then, as veins that have reached the posterior border of
the wing before this level, the two branches of Cu, and A.*
" This interpretation assumes no further dichotomies of M in the missing
portion of the wing.
BY R. J. TILLYARD.
431
On the above suppositions, we may restore the two wings as
shown in Plate xlv., fios.13-14, the missing portion of each wing
being made approximately the same as the corresponding portion
present in the other.
The total length of the wings, as restored above, would be
about 46 mm. each. If, then, the insect were of slender build,
with a narrow thorax, the total expanse of the wings must have
been about 96 nnw., or nearly four inches. This is very small for
a Protodonate; but the fossil beai's the marks of a reduced type.
Affinities of the Fossil.
This fossil is certainly one of the most puzzling of all the
Ipswich Insects. As far as I know, there is nothing in the
whole range of insect- venation, either fossil or recent, that shows
any close relationship to it. In finally determining to place it
in a new Suborder within the Protodonata, the following points
have weighed w^ith me:
(1). The differences in the branchings of the media, in the two
wings preserved, are suflicient proof that we have a fore- and a
hindwing preserved, not two forewings or two hindwings.
(2). Which wing is the fore and which is the hind, is nof
capable of absolute proof. But, unless the insect is altogether
twisted out of shape, the more anterior one should be the fore-
wing, as I have assumed it to be.
(3). The close similarity evident between fore- and hindwing
in venation, as well as in shape, puts several Orders out of court at
once; notably the Orthoptera, the Perlaria, and the Homoptera."*^
(4). In considering the type of venation exhibited, there would
appear to be characters belonging to three Orders, viz., the Pro-
todonata, Planipennia, and Mecoptera. These characters had
to be carefully weighed, with reference to known types of each
Order.
" We might note here the existence, in Australia, of Homoptera belong-
ing to the subfaniil}' Derhiiice of the family Fit/goridce, with the forewing
narrowly elongate as in Acroplana; also, a similar elongation occurs in
both wings in the genus StenobieUa of the family Tt-icJiomatidcf, in the
Order Planipennia.
432 MESOZOIC INSECTS OF QUEENSLAND, iii.,
(5). In the above compaiison, due allowance must be made for
reduction of the original venation, owing to the extreme narrow-
ing of the wing.
Taking, then, the Mecopteran characters first, it will be seen
that these may be practically summed up in the dichotomous
nature of the branchings of the media. But, in the Mecoptera,
the radial sector undergoes a similar series of dichotomous
branchings. We must assume, then, that this type of branching
is not in itself evidence of Mecopteran affinities, unless accom-
panied by other definite characters. That this is certainly so is
shown by the 'Avchsiic Pf^i/chopsis illidypi, which belongs to the
Planipennia, but which has, nevertheless, a media dichotomously
branched, in a manner not unlike that of the fossil.
The Planipennian affinities suggested are the general appearance
of the wings as being those of a "lacewing," with their numerous
parallel longitudinal veins supported by, for the most part,
delicately formed cross- veins; the great strength of tlie radius,
and the close, parallel situation of the subcosta in a deep groove
in front of it; and also the correspondence between the branch-
ings of M in the fossil and in Psychopds il/idgei, already men-
tioned .
The principal characteristic of all known Planipennia, however,
is the large number of descending branches of the radial sector.
It is these veins that form, in this Order, the majority of the
parallel longitudinal or oblique veins traversing the wing. But,
in the fossil, Rs is unbranched. This alone seems quite sufficient
to prevent us placing the fossil within the Planipennia. Further,
in all archaic Planipennia, the series of cross- veins between C and
Sc is oblique; whereas, in the fossil, those that can be seen are
at right angles to the main veins, as in Odonata and Protodonata.
A further argument against both Mecopteran and Planipennian
affinities for this fossil is the absence of any signs of macrotrichia
upon the main veins. Even the strongly built radius, which is
exceptionally well preserved, shows no signs whatever of having
carried these structures.
The Odonata themselves cannot be brought into consideration.
BY R. J. TILLVARD.
433
since the fossil evidently lacks the essential discoidal cell, not to
mention a nodus, which is almost certainly absent also. We are
therefore driven back to consider the claims of the Protodonata.
The strongly built, probably coriaceous, border is already
known in the Carboniferous genus Meganeura. The new fossil
also resembles this genus in the manner in which M appears
basally as a very weak vein diverging gently from the strongly
built R. The Protodonata, like the Odonata, are notable for
the unbranched condition of Ks. Now, in Jleganeiira, there are,
in the forewing, f^'o radial sectors arising fairly close together,
and running closely parallel for a considerable distance. If,
owing to the narrowing of the wing, in the course of e\'olution,
Text-tig. U).
Diagrams to show the structure of tiie radius and media in Protodonata:
a, suggested ancestral condition, from which there can be derived: — h,
the condition found in Ai'.roplana mirahilis, n.g. et sp., bj' fusion of
the two radial sectors (Rsj, Rso) near their bases; c, the condition
found in the forewing of Meijaneura monyl Brongn., with radial
sectors still separate, but anterior branch of media (M,) eliminated;
and d, the condition in the hindwing of the same species, witli the
ladial sectors eliminated but the media not reduced.
these two sectors were to fuse together, we might expect to get
some such formation as we find in this new fossil, viz., a single
Rs with two origins. In the hindwing of Meyaneura, however,
we find that Rs is entirely absent. But there is a pecidiar
anterior branch of M, which takes its place, and which branches
dichotomously exactly in the same way that the corresponding-
branch of M in the hindwing of the new fossil does. All
32
434 MKSOZOIC INSECTS OF QUEENSLAND, iii.,
conditions, viz., that in the forewing of Mp(janevra, tliat in the
hind wing of the same genus, and that in tlie hind wing of Aero-
plana, can be easily obtained by reduction from the type of
venation shown in Text-fig. 16, a, as may be seen from b, c, and d
of the same Text-figure.
Tlie rest of the media in Jleyanenra differs widely from that
of Aerophvtia, as is only to be expected when one considers the
difference in the shape of the wings.
The cubitus in J/ef/cmeuirt (and probably in other Protodonata
so far known) is simple. But it is a sinuously curved vein,
resembling the Cuo of Aet'oplana in its general course basally.
Both genera have only a single main anal vein. But in Mfga-
neura, as in other Protodonata, both Cu and A give off numerous
descending branches, which are absent from A'eroplana.
AVe can only conclude that there is a certain amount of
evidence of affinity betw^een J/egauf^ura and An'opJaim, though
that evidence is by no means strong. The numerous points of
difference are so evident that it will scarcely be necessary to
enumerate them. There are, then, only two courses open to us:
either we must place Aeroplana in the Order Protodonata, as a
very highly specialised member, far removed from any known
type of that Order; or we must make a new Order for it. In
choosing the forme'r alternative, while at the same time empha-
sising the unique character of the new fossil by placing it in a
new Suborder, Aeroplanoptera, I have taken a course which will
serve to indicate the degree of affinity of the fossil to the rest of
the known Protodonata, while leaving it open to any other ento-
mologist, if he considers this affinity insufficiently established, to
raise the new Suborder to the status of an Order.
In conclusion, it seems to me quite possible that this extra-
ordinary fossil may have held much the same position in the
Order Protodonata, with respect to the Megmipuridfp, that the
advanced Agrionidfa hold to-day, within the (^rder Odonata,
with respect to such gigantic, archaic types as Fetalnra. If
that be so, its existence in the Upper Triassic — at a time when,
it is U) be presumed, the rest of the Order had died out — need
BY R. J. TILL YARD. 435
occasion US no surprise, but is rather to be expected; just as we
may expect that the more advanced ])ragoiiflies of to-day will
remain in existence, loni;: after Petahira has become extinct.
BIBLIOGRAPHY.
Handliksch, a. — Die Fossileii Insekten, etc., arid Atlas. Leipsic,
1908.
TiLLYARD, R. J. — '* Mesozoic and Tertiary Insects of Queensland and
New South Wales."' Queensland Geological .Survey, Publication
No.2."i.3, IDie;. (Me.sophleJna, p.24).
"Mesozoic Insects of Queensland. "" No. 1. Planipen-
nia, Trichoptera, and the new Order Protomecoptera."' Proc. Linn.
Soc. N. S. Wales, 1917, xlii., Part 1, pp. 175-200.
"Mesozoic Insects of Queensland. No.2. The Fossil
Dragonfly ^Esdinidiopsis ( .Eschna) flindersien.s'ts (Woodward), from
the Rolling Downs (Cretaceous) Series. Op. cit., 1917, xlii., Part 4,
pp. (576-692.
Walkom, a. B. — " The (Geology of the Lower ^fesozoic Rocks of
Queensland, with special Reference to the Distribution, Fossil Flora,
and their Correlation with the Lower Mesozoic Rocks of other parts
of Australia." Proc. Linn. Soc. X. S. Wales, 1918, xliii., Part 1,
pp.37-ll."5.
EXPLANATION OF PLATES XLIV.-XLV.
Plate xliv.
Fig. 12. — Ai'i'opldiia niirahilia, n.g. et sp. Principal preserved portions of
the wings; ( x 7).
Plate xlv.
Figs. 18-14. — Airoplana mi nihil i-<, n.g. et sp. Restoration of fore- and
hindwings; ( x 4K Fig. 13, forewing; fig. 14, hindwing.
lA, analis; C, costa; Cu^, Cu2,the two branches of the cubitus; Mi-M-,
the seven principal branches of the media; R, , main stem of radius; Rs,
radial sector; Sc, subcosta.
Postscript {added Jul u 1 ith, I'.JhS). — The following important
changes of nomenclature have to be made: —
Jhsopanorpa Tillyard, 1918 (Type, J/. irianamaUeusis Till-
yard, Upper Triassic of Glenlee, N.S.AV., Mecoptera,) becomes
Mps<>panorpode>^, n.n.; preoccupied by Mesopauorpa Handlirsch,
436 MESOZOIC INSECTS OF QUEENSLAND, ill.
1908 (Type, J/, harfum/i Bi-auer, Redtenb. c^- Gaiigll)., Middlf^
Jurassic of Ust Balei, Siberia). The family MesopanorjndfF
Tillyai"d becomes tlie family Mpscqxinorjtodidfc^ ii.n. {Set" Till
yard. These Proceedings, 1917, xlii., Pt.4(1918), pp.746-7).
Creagris Hagen, 1860 (Type, C. plumhfus Oliv., Neuroptera
Planipennia, Europe) becomes Creo/pon, n.ii.; preoccupied by
Cvpayris Nietner, 1857, a genus of Coleoptera, Fam. Canihidm.
I am indebted to Mr, T. G. Sloane for this discover}-. (aS'^^
Tillyard, These Proceedings, 1918, xliii., Pt.l, p.l22). The
tribe Creayrini becomes the tribe Creoleonfini, n.n. — R.J.T.
Postscript {add^d August 7th, 1U18). — The title of Part ii. of
this series of Studies should be amended as follows:—
For ^^uEschnidiojJsis {.Eschna) fiiiidersienms ( AVoodward)", write
'■^^Eschnidioi^sis flindersiensis (Woodward), [^^Eschna]^ It was
never intended to convey the suggestion that ^Eschna is a sub-
genus of ^Eschnidiopsis, and hence the order of the names must
be altered as here shown. — R.J.T.
437
THE CARBONIFEROUS TRTL0RTTE8 OF AUSTRALIA.
By John Mitchell, late Principal, Technical College at
Newcastle, New South Wales.
(Plates xlvi.-liii.)
Introduction.
For many years, m\' collection has contained a number of
Carboniferous Trilobites new to science. When the opportunity
of describing tliem presented itself, my first intention was to
confine myself to these specimens. On subsequent consideration,
it occurred to me that palaeontoloi^ical knowledge would be
increased by extending the scope of my paper so as to embrace
all previous work on the Carboniferous Trilobita of Australia,
to describe all new forms that might be available from other
sources, and review, redescribe, and redetermine, where it might
seem necessary, those species referred to by previous authors.
It will be found, in the course of this paper, that, in a number
of instances, my views are widely divergent from those of pre-
vious authors; and, while regretting this divergence, it has not
arisen through rashness or want of reflection on my part; and I
may candidly affirm that my inability to reconcile the deter-
minations of previous authors with my own, in some cases, was
the only unpleasant part of the work expended on this paper.
My aim is to clear the literature on the Australian Carboniferous
Trilobita, to date, of errors which have undoubtedly become in-
corporated therein. This aim is one not likely to be fully
realised, but it will be my endeavour at least to lessen them,
and, if I succeed in doing this, I shall be satisfied. That the
present paper is free from errors is not one of my expectations,
for with reference to my conclusions in respect to F. woodivardi,
I am not by any means satisfied; and I shall await with some
anxiety, the discovery of better petrifactions of the remains of
this species, to make certain what its complete form actually
33
438 CARBONIFEROUS TRILOBITES OF AUSTRALIA,
The discovery of the genus Brachymetopus in the Mount
Morgan district will be of much assistance in establishing the
geological age of that area, and also in correlating its sedimentary
rocks with those of other localities. For reasons which will be
obvious, the name Phillij^sia dubia has been discarded by me.
To Messrs. W. S. Dun, Palaeontologist, Department of Mines,
Sydney, B. Dunstan, Chief Government Geologist, Brisbane, and
H. A. Longman, Director of the Queensland Museum, Brisbane,
I am indebted for specimens placed with me for inspection and
description, and for which I tender to them my thanks. To Mr.
R. Etheridge, J. P., Director and Curator of the Australian
Museum, and to Mr. Dun, my thanks are tendered for affording
me access to literature. I am very grateful to General A. W.
Vogdes, of San Diego, U.S. A, Bibliographist and Authorit}^ on
Palaeozoic Crustacea, for supplying me with notes on the classifi-
cation, and a list of the Carboniferous Trilobites described to
date. Lastly, I wish to express my thanks to Mr. F. K. Cowper
Heed, M.A., F.G.S., Acting Keeper of the Sedgwick Museum,
Cambridge, England, for supplying me with a number of liis
valuable papers, from which I have received much help.
Bibliography of Australian Carboniferous Trilobites.
(I). — 1847. Prof. McCoy (^.im. Mag. Nat. Hist., xx., p.231,
PI. xii., fig. 1) described Brachymetopus strzehckii and referred to
the occurrence of the genus Fhil/ipsia in Australian Carbonifer-
ous rocks.
(2). — 1872. R. Etheridge, Senr., (Quart. Journ. Geol. Soc,
Vol. xxviii., p 338, PL xviii., fig.7) described Griffithides dubins,
which, from the description, would appear to be an abnormal
species.
(3). — 1877. De Koninck (Foss. Pal. Nouv. Galles du Sud,
Pt. 3, p. 348, etc., PI. 24, figs. 8, 9, 9a, 10, 10a, 106, and 10c) de-
scribed and figured Griffithides (PhiUipsia) elchwaldi Fischer,
Phillipsia (Griffithides) seminifera Phillips, and Brachymetopus
strzeleckii McCoy.
(4)._1892. R. Etheridge, Junr., (Geo), and Pal. Queensland
and New Guinea, pp. 214-216, PI. xxi., figs.U-H; PI. viii., figs.
BY JOHN MITCHEJ.L.
439
5, 6; and PI. xl., figs. 4, 5, 6) described the following: — PluUipsid
dubia, P. woodwardi Eth. fil., PhiUipsia{h sp. ind., for wliich he
suggested the specific name Griffithides fieininiferus vai*. (lus-
tralasicd, from Queensland. Also in the same year [Mem Geol.
Surv. N. S. Wales, Pal., No.5, Pt. ii., pp. 126-1 30, PI. xxi., figs.
1-4, 5 (and Text-fig. f)), 6, 7,8; Pi. xxii., figs. 12-15], the same
author described PhiUipsia dubia, P. grandts, J'hillipsia spp.
ind. (two, a and b), and Grijfithides, sp. ind.
(5). -1903. F. R. C. Reed, M.A., F.G.S. (Geol. Mag., Dec. iv.,
Vol.x., No. 467, pp. 193-197) redescribed BracJtymetopus strzeleckii
McCoy, and added valuable remarks on its generic and specific
position.
(6). — 1917. R. Etheridge, Junr., (Geol Surv. of Queensland,
Publication No.260) made additional reference to the occurrence
of PhiUipsia grandis in Queensland.
(7). — Besides the above descriptions and determinations, R.
Etheridge, Junr., referred a fragment of a pygidium from West
Coerdawandy and the Yaltra Mtns., on the Gascoigne River,
W^est Australia, to his species PhiUipsia grandis (MS.).
Critical Review of the Carboniferous Trilobites of
Australia, previously described.
Order TRILOBITA.
Family PROETID^.
Genus Brachymetopus McCoy, 1847.
1. Brachymetopus strzeleckii McCoy, Ann. Mag. Nat. Hist.,
XX., 1847, p.231, PI. xii., fig.l.
Since this species was described by McCoy, no specimens of it,
or of any other member of the genus, appear to have been found
by collectors, though on the occasion of the first find at Dun-
vegan, N. S. Wales, quite a number seem to have been obtained.
Genus P h i l l i p s i a Portlock, 1 843.
To Prof. McCoy we owe the first reference to the occurrence
of this genus in Australia {loc. cit. antea).
440 CARBONIFEROUS TRILOBITES OF AUSTRALIA,
2. Phillipsia eichwaldi De Kon., (non P. eichwaldi Fischer)
Foss. Pal. Nouv. Galles du Sud, 1877, Pt.3, p.348, t.24, f.9.
Under the name of Griffithides {Phillipsia) eichwaldi, De
Koninck described a trilobite from Colo Colo, N.S„W., and de-
termined it to be P. eichivaldi Fischer; but, as the writer points
out further on, this determination was evidently incorrect. Mr.
Etheridge, Junr., expressed a similar view (Etheridge, Junr., Mem.
Geol. Surv. N. S. Wales, Pal. No.5, Pt. ii., 1892, p.l24). There
is little doubt that the presence of this trilobite in Australia
remains yet to be established.
3. Phillipsia dubia Etheridge, Junr., (non Griffithides diihins
Etheridge, Senr.) Geol. and Pal. Queensland and New Guinea,
1892, pp.214-215, PL vii., fig.5; PL xliv., fig.4.
Phillipsia dubia Etheridge, Junr., (non Griffithides dnhiiis
Etheridge, Senr.) Mem. Geol. 8urv. N. S. Wales, Pal. No. 5, Pt. ii.,
1892, p.l26, PL xxi., figs.1-4, PL xxii., figs.l2, 13.
The above two forms were considered bv Mr. Etheridge, Junr.,
to be specifically identical with Griffi'thides dnhiiis Etheridge,
Senr.; and the latter to be generically misplaced. The writer
regrets that, with this determination, he cannot agree with Mr.
Etheridge. It appears to him that neither of the above trilobites
is specifically identical with Grijfithides duhius Eth. Senr., nor
with each other. Mr. Etheridge, Junr., wlien describing the
trilobite from Binge Berry, Rouchel Brook, New South Wales
iloc. cit. antea) pointed out that it diftered from the Queensland
form in having eighteen to twenty axial rings in the pygidium
instead of eight to ten, which is the number in that of the
Queensland form. This alone is suflficient to separate them
specifically; but the head-shields of the two also differ. Both
forms are here treated as separate species. The writer's view
regarding the Queensland Phillipsia dubia is, that it cannot at
present be generically or specifically joined with Griffithides
dubius; and that the latter species must still be recognised.
Only the discovery of a number of specimens of this form, show-
ing the stages of development from the immature to the mature
state, can satisfactorily establish the proper position generically
and specifically of Griffithides dubius Eth. Senr.
fiY JOHN MITCHELL. 44l
4. Phillipsia grandis Etheridge, Jurir., Geol. and Pal. Queens-
land and New Guinea, 1892, p.215. Etheridge, Junr., Mem.
Geol. 8urv. N. S Wales, Pal. No.o, Pt. ii., 1892, Text-6g.5, and
Pi. xxi., fig. 5. Etheridge, Junr., Geol. Surv. Queensland, Pub-
lication No.260, pp.11-12, PI. iii., fig 3.
It is unfortunate that Mr. Etheridge placed tliese two forms
specifically together. They are quite difterent, and are so
treated by the writer.. In describing them, their differences
will be fully shown. As the Queensland form was the one for
which the specific name grandis was first suggested by Mr.
Etheridge, it is proposed that it should continue to be known by
this name. To tlie New South Wales one, a new specific term
will be given.
The name grandis is quite appropriate for either of the forms;
because their pygidia are larger than those of any other known
Carboniferous trilobite.
5. Phillipsia woodwardi Etheridge, Junr., Geol. and Pal.
Queensland and New Guinea, 1892, p. 215, PI. 7, figs. 11, 13; PL
44, figs. 5, 6.
This is a singular species, to which some reference will be
made later on.
6. Besides the foregoing species of Phillipsia, which have been
specifically determined, Mr. Etheridge described two pygidia
from Binge Berry, Rouchel Brook, and near Paterson, respect-
ively, and referred them to this genus, but did not give them
specific rank.
7. (a) Phillipsia sp. ind. (a), Etheridge, Junr., Mem. Geol.
Surv. N S. Wales, Pal. No. f., Pt. ii., 1892, p. 1 29, PL xxi , figs.
6-8.
(b). Phillipsia sp. ind. (6), Etlieridge, Junr., op. cit., p. 129,
PL xxiL, fig. 14.
Both of these jvill be redescribed and named.
Genus G R i f f i t h i d e s Portlock, 1843.
8. (^iRiFFiTHiDES DUBius Etheridge, Senr., Quart. Journ. Geol.
Soc, 1872, xxviii., p.338, t.l8, f.7.
442 CARBONIFEROUS TRILOBITES OP AUSTRALIA,
The writer is of the opinion that this species should still be
recognised. Its description is the first record of the genus in
Australian rocks.
9. Gkiffithides skminiferus De Koninck (non Pliillips,
species) Foss. Pal. Nouv. Galles du Sud, 1877, Pt.3, p.348, t.24,
f.9, 9a.
That this determination on the part of ])e Koninck was in-
correct, there is really no doubt. It is fully dealt with in the
observations on the relationship of Phi/lipsia collitisi to other
species, described further on.
10. Gkiffithides seminifeuus var, austkalasica Etheridge,
Junr., Geol. and Pal. Queensland and New Guinea, 1892, p. 216,
PI. vii., fig. 14.
The material used by Mr. Etheridge for his description of the
above is before me, and, after close examination of it, I am
doubtful of the correctness of Mr Etheridge's conclusions. The
pygidia, by which his conclusions were greatly influenced, have
each thirteen and eleven axial and pleural divisions respectively;
and, in this respect, agree with pygidia }>laced by me with his
P. woodwardi. All these pygidia agree in tlieir ornamentation.
As regards the cephalon associated on the specimen with these
pygidia, it does not appear to differ in any essential from two of
those included by Mr. Etheridge in his F. ivoodwardi, except
that, in the latter, the granulation has been worn off by weather-
ing. On the glabella of the former, it is true, no anterior or
ftiesial glabellar furrows are visible, but this glabella is a very
imperfect intaglio. If it should ultimately be shown that this
fossil is a good species, it will not, even then, be closely related
to Gr. semitiiferns, because it possesses supplementary basal
lobes, and the latter does not. This difference certainly places
them specifically apart. It may be stated that, so disposed was
I to the opinion that this was a separate species, that I had
written a description of it under a new name, but not being able
to discern any difference in the pygidia now under discussion,
and those I have placed with F. woodtvardi, I deferred final
judgment.
BY JOHN MITCHELr.. 443
11. Griffithides sweeti Etheridge, Junr., Proc. Linn. Soc.
N. S. Wales, 1894, p. 528, PI. xxxix., f.3.
The writer lias not seen the original specimen. Judging from
the description and illustration, it is a very unusual type. See
description further on.
12. Griffithides sp. ind., Etheridge, Junr., Mem. Geol. Surv.
N. 8 Wales, Pal. No.5, Pt. ii., 1892, p.l30, PI. xxii., figs.lo, 16.
It is not improbable that this may be identical with Phillipsia
collhisi.
From, the foregoing review, the following conclusions have
been drawn, viz. : —
i. That, of the nine species of Australian Carboniferous trilo-
bites previously described and named, only the five following are
worthy of recognition.
1. lirachymetojnis strzeleckii McCoy.
2. Griffithides dubius Eth. 8enr.
3. Griffithiiies sweeti Eth. Junr.
4. PhiUipsia icoodfvardi Eth. Junr., Geol. et Pal. Queens-
land and New Guinea, p. 215, PI. 7, figs. 11, 13.
b. PhillijJsia yrandis l^lh. iuin\, oi which the type is the
Mt. Morgan form. Mem. Geol. Surv. N.S. Wales, Pal.
No.5, Pt, ii., 1892, p. 128, PI. xxi., fig.5.
ii. That the two determinations made by the late Prof. i)e
Koiiinck are incorrect, and scientifically valueless.
Family PROETID^ Steiiib.
Genus Phillipsia Portlock, 1843.
Dr. H. Woodward's* description of the genus is as follows: —
"General form oval; glabella with nearly parallel sides, marked
by either two or three pairs of short lateral furrows; the posterior
angles, forming the basal lobes, always separated by a circular
furrow from the rest of the glabella; eyes large, reniform, surface
delicately faceted; cervical furrow deep; free cheeks separated
from the glabella by the axal (facial) suture, which forms an
acute angle with the circular border of the cheek in front of the
* Moil. Brit. Carb. Trilobites, 1883-4, p. 11.
444 CARBONIFEROUS TRILOBITES OP AUSTRALIA,
glabella; whilst the facial suture cuts obliquely across the pos-
terior margin, just behind the eye, leaving a small pointed por-
tion fixed to the glabella b}' the neck lobe; angles of the cheeks
more or less produced, mart^in of head incurved, forming a
striated and punctured rim. Thoracic segments nine in number,
the axis distinctly marked off from the side-lobes or pleurae by
the axal furrows; the abdomen or pygidium usually with a
rounded border, the axis composed of from 12 to 18 coalesced
segments."
Phillipsia collinsi, sp.nov.
(PI. xlvi., figs. 1-5; PI. xlviii., figs. 8, 9; PL lii., fig. 4).
Sp. CVtars. — Complete form elongately ellipsoidal.
Cephalon sub-semielliptic, moderately inflated, densely tuber-
culated; glabella sub-oblong or sub-bellshaped, narrow, upper
surface flat, steep and straight sided, densely crowded with pro-
minent tubercles of variable sizes; medially the tubercles are
arranged in four sublongitudinal rows, those on the anterior pari
are subconcentric; front gently rounded and bearing a concentric
row of tubercles; lateral lobes and furrows distinct, the basal
pair altogether circumscribed, and bearing a varying number of
tubercles, one usually much larger than the others; limb narrow,
witli a gentle upward turn, smooth or very finely granulate;
neck-furrow distinct, narrow, and deep behind the basal lobes;
neck-ring strong, moderately arched and tuberculate, its lateral
extensions being narrow; fixed cheeks very small, eye or palpe-
bral lobes bounded by tubercles; eyes apparently crescentic; free
cheeks unknown. Axial furrows distinct; facial sutures ante-
riorly very gently sinuate (almost straight) and close to the
axial furrows; posteriorly straight, and almost at right angles to
the axial line, and at last cutting the fixed-cheeks with a sharp
backward turn.
TJiorax oblong, moderately inflated and granulated; axis pro-
minent, the rings centrally and at the bases mildly curve forward;
bases feebly tuberculated, width equal to that of one side-lobe,
median sulci of the rings narrow; pleura? convex, medial furrows
of each segment wide and shallow, and reaching the distal end.
BY JOHN MITCHELL. 44 5
which is rounded; each posterior segmental ridge bears a row of
tubercles, as du also the axial rings; axial furrows distinct.
Pygidinvi semi-ellipsoidal, moderately tumid and strongly
tubercuJate; proportion of length to width equals 3 : 4 approxi-
mately; axis has fourteen rings, prominent, dorsally depressed,
sides steep, contracting very gradually posteriorly to about half
the anterior width, terminating bluntly and somewhat overhang-
ing the border, the rings bearing tubercles which vary in number
with their spread from two or more on the posterior ring, to
eight on the anterior ones, and not exactly forming longitudinal
rows; axial furrows pronounced; pleurae very convex, arching
from the axial grooves at first gently and then steeply to the
narrow furrow separating them from the border, where all the
segments, except the anterior pair, end, segments ten in number,
each succeeding one having a gently backward trend, and dimin-
ishing in length till the last two pairs are represented by mere
tubercles. Each segment bears a row of tubercles along the
posterior ridge varying in number with its length, from one to
ten, and perhaps in some instances more; medial segmental
furrows wide and shallow, reaching to the faint furrow within
the border; the border itself is fairly wide, and thickened, and-
in normal specimens, has the same convexity as the pleura] ribs;
it bears a row of conspicuous tubercles usually placed in line
with each rib, and three to five behind the axis, these tubercles
being occasionally elongated or double; when the thickened por-
tion is removed, the undersurface shows concentric striations,
which is a common feature in the Proetidte. Only the first pair
of the pleural segments interrupts the continuity of the l)order.
Ohs —This species is (juite singular. Its outstanding features
are:— 1. The narrow, straight-sided glabella. 2. 'J'he crowded
and conspicuous tuberculation uf the glabella. 3. The equality
of the lengths of thepygidium and thorax. 4. The width of the
thorax, which equals half the total length of the complete indi-
vidual. 5. The closeness of the eyes to the axial groove. G. The
straightness of the anterior branches of the facial sutures, and
their parallelism to the axial furrows.
Dimensions: total length of an individual not quite mature.
446 CARBONIFEROUS TRILOBITES OF AUSTRALIA
one inch: widtli, half an inch; cephalon, one-quarter inch: thorax
and pygidiuH), each three-eighths of an inch.
In the proportions of length to width, and the character of
the pvgidial granulation, this trilobite agrees with P. gemmu-
lifera Phillips, with which it also agrees in the equality of the
lengths of the thorax and pygidium. In other specific features,
thev differ. In the cephalons and pygidia of F. collinsi and F.
truncdtuld, there is much common to the two. They agree in
possessing narrow, straight-sided and tul)erculate glabella, eyes
close to the axial furrows, and similarly tuberculated pygidia:
but the tuberculation on the glabella of the former is much more
pronounced than it is on the latter, and the sinuate course of the
anterior branches of the facial sutures of the latter is more pro-
nounced than that of the former. Thepj^gidium of F truncatula
has seventeen or eighteen annulations in the axis, and fourteen
divisions in each pleura ; but in F. collinsi^ these parts
number fourteen (or thirteen and a terminal piece), and ten
respectively. In the former, the pygidium has no border, and
the pleural ribs extend to the periphery; on the other hand, the
latter species has a very distinct pygidial border, which, except
in the case of the anterior pairs, the pleural ribs do not cross.
The tubercles of the pygidial axis of the former are arranged in
longitudinal rows; this is not quite so with those of the latter.
Of the North American Phillipsif^, the nearest relative to ours
appears to be P. insignis, which apparently has the same number
of divisions in the axis and pleurae of the pygidium, as are in
the similar parts of P. collinsi. The tuberculation of the axial
rings is much alike in the two species. In other respects, they
differ rather widely. The only Australian Carboniferous trilobite
fragments bearing any resemblance to P. collinsi are two pygidia
described and figured, but not named, by Mr. Etheridge, Junr.*
The one pygidium he placed in the genus FhiUipsia {loc. cit., PI.
xxii., fig. 14). It agrees with F. collinsi in possessing fourteen
rings in the pygidial axis, and in having a tuberculate test; but
differs in having a continuous, smooth, flattened border, and
'" R. Etheridge, Junr., Mem. Geol. 8urv. N. 8. Wales, Pal. No.5, Pt. ii.,
1892, pp. 129-130, PI. xxii., figs. 14, 15.
BY JOHN MITCHKLL. 447
twelve pleural segments, and less disparity between length and
width of the pygidium.
The pygidium {loc. cit., PI. xxii., fig. 15) placed by Mr Ether-
idge in the genus Grijfithides, bears a resemblance to the present
species in the character of its border and granulation, and in
having the same number of pleural segments; but its axis has
less rings, and ends in a sharp point.
Lastly, referring to De Koninck's P. {Grijjithides) seinhiifera
{noil Phillips sp.)* from Colo Colo. The dimensions, number of
rings in the axis of the pygidium, character of the granulation
generally on the pygidium and thorax agree closely with similar
features of F. collinsi: but the two forms widely differ in the
outlines of their glabellse and pygidia, if De Koninck's figures
are to be relied upon, which is doubtful, as they do not agree
with the text. In the former, the glabella is represented as
being conical in outline; and, in the latter, it is said to have an
anterior width slightly less than that of its base. Whatever
Phillipsia [Grijfithides) seminifera De Koninck, may have been,
it is practically certain it was not Grijjithides seminiferus
Phillips; but it may have been identical with the present species.
Named after Mr. Collins, C.C.M., Lecturer in Coalmining and
Mine Surveying at the Newcastle Technical College.
Loc. and H or. — Glen AVilliam Road, one mile from Clarence-
town, Parish Parr, County Durham. Lower(?) Carboniferous.
Philijpsia coulteri, sp.nov.
(Plate xlvi., figs 6-10;.
Sjo. Chars. — Complete form subelliptic.
Cej^halon sub-semielliptic, fairly inflated, finely granulated :
glabella subrectangular, rounded in front, lateral furrows and
lobes not visible, owing to the loss of the part bearing them, and
the exposure of the hypostome; neck-furrow shallow, its lateral
extensions across the cheek.s shallow and wide; neck-ring narrow;
axial furrows faint. Fixed cheeks small. Free cheeks relatively
large, strongly inflated, steep between the eye and border-furrow,
which is wide and shallow, outer edge of border only moderately
* Pal. Foss. Nouv. Galles du Sud, 1877, p.'267, PI. xxix., figs. 9, 9m.
448 CARBONIFEROUS TRILOBlTfiS Of AUSTRALIA,
thickened. Eyes prominent, large, crescentic and faceted.
Genal angles rounded. Hypostome alate, shield like, and striated
in tiie fashion common to Phillipsise.
Thoi'dx 11 mm. wide and 9 mm. long, appearing to have been
finely granulated, with a few scattered tubercles; straight-sided.
Axis very strongly and acutely arched, rings, centrally and
basally, strongh^ inclined anteriorly, each one bearing along the
medial line a rath--r prominent tubercle, giving the axis through-
out a serrated character: spread equal to that of one side-lobe,
mildly tuberculate, height and width diminishing gradualh* pos-
teriorly, axial grooves faint. Pleurae gently rising from the
axial grooves to the fulcra, thence steeply deflected, mesial fur-
rows of the somites shallow and wide, and reaching the extremi-
ties, which are rounded: along the fulcral line each rib bears a
tubercle on the posterior ridge, and there is evidence of the
presence of general fine tuberculation or granulation.
Pygidium semielliptic, length 8 mm., width 10 mm. Axis
very prominent, the first ring having a medial forward trend,
rings eight or nine, counting the end-piece, which is buttress-
like, and ends at the border in a rather fine point, and bears a
subconspicuous tubercle. Pleuriie with seven divisions, all rather
inconspicuous in the specimen (decorticated) serving for this de-
scription, moderately convex; mesial furrows very shallow and
reaching to the faint furrow within the border, each segment
bearing a tubercle at the fulcral angle, and showing traces of
other tuberculation, as do also the axial rings; the border is
relatively wide, steep, gently thickened, and bounded inwardly
by a faint furrow, and is crossed only by the first pair of ribs.
(9^^^ —Though the glabellar furrows and lobes are not visible
in the specimen described, the shape of the glabella and character
of the hypostome leave no doubt as to its being a Phillipsia, and
the specimen is one of the finest of this genus yet discovered in
Australia. It difiers so widely from other Australian species
that there is no need to detail its relationship. It bears no close
relationship to any of the American Phillipsiai referi-ed to in
Brigadier General A. W. Vogdes' Monograph.*
* " The Genera and Species of North American Carb. Trilobites,"'
Amialb N. Y. Acad. Sci., 1888, vi.
BY JOHN MITCHELL. 449
The species has been named after Mr, E. Coulter, of Katoomba,
an enthusiastic student of geology, who discovered and presented
the specimen to the author.
Loc. and /To?*. — Stroud-Dungog Road, near Dungog, Parish
Howick, County Durham. Lower Carboniferous.
Phillipsia brkvickps, sp.nov.
(Plate xlvi., figs.ll, 12; and Plate li., fig.2).
S]). CAars. — Complete form unknown.
Cephalon apparently subsemicireular, finely granulated. Gla-
bella straight-sided, rounded in front, very gently convex,
highest just in front of the neck-furrow, smooth, though a lens
shows evidence of microscopic granulation, two pairs of lateral
furrows visible, basal and mesial. 'J'he former very shallow and
circumscribing the basal lobes, which are large and pyriform;
the other narrow; neck-furrow narrow and shallow; neck-ring
relatively wide, decidedly arched back ward ly and lobed at its
l)ases; frontal limb narrow and gently recurved. Fixed cheeks
very small, eye-lobes small, and abutting the axial furrows.
Facial sutures anteriorly rather straight and close to the axial
furrows; eyes of moderate size only, judging by the size of the
palebral lobes.
Thorax unknown.
Pygidium sub-semielliptical, moderately convex, very fineh'
granulated; width 10 mm., and length 7 mm. respectively. Axis
prominent, consisting of thirteen rings, and a very narrow end-
piece, spread equal to that of each side-lobe, contracting very
gradually posteriorly, and terminating short of the border,
rounded and bluntly, with half its anterior width; some of the
rings bear very fine tubercles, barely distinguishable without
the aid of a lens; side-lobes moderately convex, with ten, or
doubtfully eleven, pairs of segments, of which the medial furrows
are distinct, and, in the case of the four anterior pairs, at least
reach the edge and interrupt the narrow border; a few of the
posterior segments bear very small tubercles at their junction
with the border, and three similar tubercles occur behind the
axis, on the axis itself centrally the posterior ridges also bear
fine tubercles.
450 CARBONIFEROUS TRILOBITES OF AUSTRALIA,
6)6s. — Aswill be apparent from a comparison of the above
description with that of P. derbieitsis Martin, the two fossils
agi-ee in several important features. For example, in both the
glabella is parallel-sided and practically smooth, anterior facial
sutures rather straight and close to the axial furrows, eyes reni-
form, pygidial axis with thirteen rings, and ten segments in the
pleur?e; and the pygidial borders obsolescing before reaching the
articulating face; in the equality of maximum widths of the axes
and pleurae of the pygidia (this is only true in the case of the
above species when the pleurae are measured obliquely, and not
the actual horizontal widths: if measured in the latter way, the
axis is wider than one pleura) and the faint tendency to orna-
mentation on these parts. Against these resemblances, there
are the following differences to be considered. The length of
the glabella of the local species, measured from the neck-furro\v
to its front, is equal to its width between the axial furrows
across the basal lobes; in that of J\ derbiensis, these dimensions
are respectively about 4 : 2. The eye-lobes are shorter, and the
frontal border wider, neck-ring more intensely arched posteriorly
in the local form than in the British one. The distinct lobation
of the bases of the neck-ring, relatively short cephalon, and the
almost equality of the lengths of the cephalon and tail are very
marked features in the local species, separating it from the other.
Then tliere is the distinct, though fine, granulation, more or less
tuberculation of the pygidium of the form now described, as
against a practically smooth one in the other. Further, judging
from the anterior pair of pleural segments of the pygidium, the
thoracic pleural ribs were not imbricated, in the case of the local
species. Although undoubtedly closely related to each other, I
have concluded that the differences between them are sufficient
to justify their separation specifically.
The glabellie of the above species and Proetus inissourieiisis
are much alike. I am indebted to J^rigadier General Vogdes'
paper [op. cit., antea) for the references made to American Car-
boniferous trilobites.
Lvc. and Uor. — Neighbourhood of Port Stephens, probably
from near Bulladelah. Carboniferous.
BY JOHN MITCHELL. 451
Philltpsia proxima, sp.nov.
(Plate xlvi., figs.lf), 16).
Sp. Chars. — Pygidinm semielliptical, fairly convex, practically
smooth, so fine is the granulation. Length 12 mm., width 15
mm. Axis strongly convex, rings twenty, tapering posteriorly
very gradually, and ending bluntly at the border, with a spread
of about half that of the anterior ring; axial furrow shallow, but
distinct. Pleurje gently convex, consisting of twelve pairs of
ribs, the last one or two very shoit and faint, all terminating at
the border, which is continuous to the articulating face and sub-
depressed: medial furrows of the segments, except in the cases
of a few of the posterior pairs, well defined, anchylosing ridges
also fairly prominent; furrow separating border and pleural ribs
distinct.
Ohs. — This p3"gidium agrees with that of PhiUipsia eiehivaldi
Fischer, in the continuous smooth and somewhat depressed
border, and the character of the pleural segments. The most
important differences between them are— (1) the pygidium of
F. eichivaldi is just as wide as long, is distinctly granulated, and
the distal axial end does not seem to be prominent. The
pygidium above-described has a length only four-fifths of its
width, and is practically smooth. The total length of the pygidia
of mature individuals of the former, as shown by the beautiful
figures in Dr. H. Woodward's work* is 9-5 mm., that of the
latter is 12 mm. The large number of rings in the axis of ours
places it far apart from the other. Named because of its close
resemblance to P. eichivaldi Fischer, in shape.
Loc. and H or. — Glen William Road, near Clarencetown, Parish
Parr, County Durham. Lower Carboniferous.
Phillipsia('?) robusta, sp.nov.
(Plate xlvii., figs.l and 8).
Phillipsia graridis Eth. fil., Mem. Geol. Surv. N. S. Wales, No.
5, Pt. ii., 1892, p.l28, text-fig.5 {non PI. xxi., fig.5).
Sp. Chars. — Cephaloji and thorax unknown.
Pygidium subsemicircular, strongly convex, very finely granu-
* Mon. Brit. Carb. Trilobites, 1883-4, PI. iv., figs. 9 and 1.3.
452 CARBONIFEROUS TRILOBITKS OF AUSTRALIA,
late, greatest width 44 mm., length 26 mm., (the width here given
is what the writer considers to be the normal, and is calculated
from the width of the left side-lobe and half the spread c)f the
axis, because the right side-lobe is distorted). Axis fairly pro-
minent, evenly arched without any decided backward or forward
inclination of the annulations, of which there are fourteen: the
furrows of these are wide and V-shaped in section, ridges high,
edged, and bore some small tubercles, bases not tuberculate.
Axial furrows shallow. Pleurae very convex, arching strongly
from the axial furrows, and showing no decided fulcra; segmental
divisions twelve pairs, which reach to the border, and are only
wently directed posteriorly, except in the case of the last two
pairs; their mesial furrows are V-shaped, wide, and reach to the
border, ridges high and chisel-edged, and show no sutures, hence
the pleural segments in this oygidium were very completely
fused: some of the ridges exhibit evidence of very fine and sparse
tuberculation, anterior pair facetti. Border continuous, wide,
steep, and its undersurface concentri -ally striated.
Qf)s. — 11iis pygidium was described and figured by R. Etheridge
Junr.,* and considered by him to be identical with a pygidial
fragment from near Mt. Morgan, Queensland, and for which he
had previously suggested the name Fhillipsia gi-andis.j Mr.
Etheridge figured this Queensland specimen, when dealing with
some New South Wales trilobites.| The writer is unable to
accept Mr. Etheridge's determination of the identity of these
two pygidia, for the following reasons. 1. The part of the
pleural lobe preserved on the Queensland pygidium shows fifteen
segments, and, when complete, without doubt had several others,
as against twelve in the New South Wales one. 2. Though the
Queensland tail, when complete, was apparently longer than
that of the New South Wales specimen, the greatest Midth of
its pleural lobes was only 10 mm., as against 15 mm. for that of
" Mem. Geol. Survey N. S. Wales, Pal. No.o, Pt. ii., 1892, p. 128, text-
fig. 5 {uoii PI. xxi., fig. 5).
t Jack and Etheridge, Cleol. and Pal. of Queensland and New Guinea,
1892, pp.2irj, 216.
X Mem, Geol. Survey N. S. Wales, Pal. No.5, Pt. ii., 1892, PI. xxi., fig.o.
BY JOHN MITCHELL 453
the latter. 3. In the Queensland one, the pleural segments
along the ridges are separated by very distinct sutures; in the
other, this is not so, 4. In the Queensland one, the border is
not nearly continuous, for the three anterior pairs of pleural
segments reach to the edge of the lobes, and these are the only
ones which are perfect on the portion of the left and pleural lobe
of the specimen. 5. The mesial furrows and ridges of the pleural
segments are not alike in the two. 6. In the Queensland tail,
there were not less than sixteen or more axial rings, but fourteen
only in that of the other. 7. The axial grooves are deeper in
the specimen from Queensland, than in the one from New South
Wales. The two agree in the character of their granulations;
but that of the Queensland one is the coarser.
That the Queensland pygidial fragment represents a new
species of either Phillipsia or Griffithides. there is no doubt;
and, to it, my proposition is to apply the name Phillipsia grandis,
first suggested for it by my friend, Mr. Etheridge.
P.i'ohnsia differs from all species of this genus, or of Griffith-
ides known to me. Mr. Etheridge, too, noted (loc. cit.) how
different it was from foreign Carboniferous trilobites.
Loc. and Hor. — Swain's Conditional Purchase, seven miles
south-east of Carroll, County Buckland (D. A. Porter). Car-,
boniferous.
Phillipsia(?) stroui>e;nsis, sp.nov.
(Plate xlvi., fig.14; Plate li., fig.ll),
Sp. Chars.-^Cephalon and thorax unknown.
Pygidium semielliptic, densely and finely granulate, moder-
ately convex; greatest width 17 mm., length 15 mm. Axis only
moderately convex. Consists of twenty annulations, a few of
the anterior of which have a very gentle forward inclination,
centrally and basally ; contraction posteriorly very gradual,
terminating bluntly, and rounded at the border, with half the
anterior width (3 mm.), medial furrows of the rings wide and
shallow; anchylosing ridges not very prominent, but distinctly
granulated, anterior spread slightly less than that of one side-
lobe. Axial furrows faint. Pleurae moderately convex, seg-
ments fifteen, the last faint, anterior pair faceted, all finely and
34
454 CARBONIFEROUS TRILOBITES OF AUSTRALIA,
distinctly granulate, reaching to the border, and gently inclined
backward (falcate). Medial furrows wide and shallow, reaching
the border, ridges low. Border continuous, narrow anteriorly,
gently increasing in width posteriorly, flat, densely and finely
granulate.
O65. — This is a fairly large pygidium, as the dimensions given
will show; but it is, in this respect, exceeded by both of the
forms placed by Mr. R. Etheridge Junr., under the name of P.
yrandifi; but it has no specific relations with either of them. In
the number of axial rings, it agrees with P. elongata, but not in
other features. It is different from all the British Phillipsise
the writer has been able to compare it with; and the same is
true as far as the North American species are concerned.
Log and Hot. — Stroud, County Gloucester, N.S.W., associated
with Orthis resupiiiata and Spirifer striattis. Lower(?) Car-
boniferous.
Phillipsia superba, sp.nov.
(Plate xlviii., fig. 15: Plate Hi., fig.3).
Complete form unknown.
Sp. Chars.— (7</?Aa?o?z semicircular or very nearly so, microscopi-
cally granulate, practically smooth; length 6 mm., width just on
double its length. Glabella subfiddle-shaped, orsuboblong with
a very gently rounded front, practically smooth, very slightly
convex, and decidedly contracted at the middle; three glabellar
furrows (doubtfully four), anterior pair or pairs very short, mesial
pair falcate and very faint, posterior pair faint at their origin;
inwardly wide and deep. Anterior and mesial lobes ill-defined,
the latter indistinctly separated from the basal pair, which are
small, reniform, and prominent, standing out from the posterior
part of the glabella, like tubercles. Neck-furrow distinct, with
a decided central and basal forward curve, its lateral extensions
straight and well-defined, joining with the lateral furrows of the
free cheeks within the genal angles. Neck-ring strong, convex,
centrally and basally anteriorly inclined, bases also gently tuber-
culate; lateral extension narrow and not prominent. Axial
grooves relatively narrow, deep, and sinuate. Limb narrow,
with gently thickened and mildly raised margin. Facial sutures
BY JOHN MITCHELL. 455
anteriorly gently sinuate, and posteriorly shortly obliquely
curved. Fixed cheeks very small, palpebral lobe almost linear,
but convex. Free cheeks relatively fairly large, subdepressed,
border wide, depressed; margin very slightly thickened, furrow
shallow, linear; bands just within the border, and the similar
bands under the shallow, wide furrows beneath the eyes, bear
rows of granules. Genal angles apparently rounded. Eyes
relatively large, reniform and faceted.
Obs. — This cephalon is clearly separated from all other known
Australian Phillipsise. In the shape of the glabella and possibly
in the possession of four glabellar furrows, relative size and
situation of the eyes, it bears likeness to P. eichivaldi; but it
differs from that species in having a nearly semicircular cephalon,
rounded genal angles, and in the absence of distinct cephalic
granulation, and of genal spines. Also, it resembles P. derhiensis
Martin, in the form of the glabella, small free cheeks, and char-
acter of its facial sutures, small palpebral lobes, rounded genal
angles, and widely differs in other respects. No North American
Phillipsi?e known to the writer, have any close relationship with
this one.
Loc. and Hor. — Supposed to have been obtained near Dungog,
N.S.W.
Phillipsia(?) waterhousei, sp.nov.
(Plate xlviii., figs.16-18: Plate Hi., fig.7).
Complete form oval.
Sp. Chars. — Cephalon subsemicircular, greatest length and
width approximately 2 mm,, and 4 mm., respectively; finely and
densely granulated. Glabella subcylindrical, narrow, very convex,
anteriorly sloping to the border rather steeply, basal glabellar fur-
rows small; the anterior and mesial pairs are visible on the cover,
basal lobes small, neck-furrows wide and deep, lateral extensions
similar; neck-ring strong, very convex, its lateral extensions also
strong; frontal limb narrow and close to the glabella. Facial
sutures, anteriorly, only gently diverging laterally in their course,
posteriorly short and oblique. Fixed cheeks small. Free cheeks
moderately large and strongly inflated, borders relatively wide
and depressed; genal angles produced into spines apparently ex-
456 CARBONIFEROUS TRILOBITES OF AUSTRALIA,
tending almost to the distal end of the thorax. Eyes crescentic,
relatively large, faceted and depressed. Axial grooves shallow.
Thorax apparently has the normal number of somites, eight
being plainly recognisable by the aid of a lens, the whole surface
finely granulated. Axis strongly and rather acutely arched, as
wide anteriorly as one side-lobe; posteriorly diminishing gradu-
ally, the fourth ring appears to be much stronger than the others,
and resembles in this respect the neck-ring; all the rings are
directed anteriorly, and bear a row of very fine granules on the
ridges. Side-lobes strongly convex, ridges and valleys of the
segments strong and deep respectively, ridges bearing rows of
fine granules; segmental extremities faceted. Axial furrows
narrow.
Pygidium subtriangular, fairly inflated, granular throughout
in a way similar to the thorax; axis prominent, diminishing
rapidly posteriorly, and terminating at the border with a moder-
ately fine point; the rings appear to be eight in number. Side-
lobes convex, made up of six pairs of segments, apparently.
Border narrow, steep and continuous.
Obs. — In so small a specimen, it is a difficult matter to clearly
distinguish its features; but, under a lens, they become discerni-
ble. The glabella in shape slightly resembles that of a Griffith-
ides; and had it not been for the presence on it of three pairs of
glabellar furrows, I would have placed it in this genus. It is
the smallest and most perfect Carboniferous trilobite belonging
either to Phillipsia or Griffithides obtained in Australia. It
may represent only an immature individual, but still mature
enough to have all its chief parts developed.
In the shape of the glabella and of the pygidium, this species
is not unlike Phillipsia leei Woodw., and P. 7ninor Woodw.
Dedicated to J. Waterhouse, M.A., Government Inspector of
Schools.
Log. and Hor. — Probable Paterson or Dungog District, N.S. W.
Carboniferous.
Phillipsia elongata, sp.nov.
(PL xlvii., figs.3-5; PI. xlviii., fig.14; and PI. 1., figs.4-7)
Phillipsia duhia Eth. fil,, Mem. Geol. Surv. N. S. Wales, Pal.
BY JOHN MITCHELL. 457
No. 5, Pt. ii., 1892, pp.126, 127, PI. xxi., figs. 1-4, and PI. xxii.,
figs. 12, 13.
Complete form elongately oval.
Sp. Chars. — Cephalon sub-semielliptic, smooth, only moder-
ately inflated, greatest length 12 mm., width between the genal
angles 18 mm. Glabella long, narrow, sub-bellshaped, convex
and prominent, highest between the anterior lateral furrows, and
from thence sloping to the front, which is gently rounded; lateral
furrows distinct, first pair short, linear and gently curved,
middle pair linear with a posterior curve, posterior pair wide,
shallow and circumscribing the basal lobes, which are relatively
small and suboval. Neck-furrow shallow, its lateral extensions
shallow. Neck-ring narrow, with a strong forward inclination,
its lateral extensions also narrow. Frontal border fairly wide,
especially at the antero-lateral angles. Facial sutures strongly
sinuate anteriorly, and posteriorly obliquely crossing the fixed
cheeks nearly in a line with the outer edge of the thorax. Axial
furrows deep. Fixed cheeks small. Eye-lobes small, crescentic
and elevated. Free cheeks relatively large, depressed, border
wide, space between the border and furrow beneath the eyes
short and steep; and immediately under the eye is a groove.
Eyes large, subcrescentic, very distinctly faceted, and wider
behind than in front. Genal angles spinate, the spines evidently
long. Thorax with the normal number of somites; length two-
thirds of the greatest width, sides subparallel, surface smooth or
very linely granulate. Axis prominent, rings with a distinct
forward inclination centrally, and showing slight indication on
the ridges of granulation, spread equal to one side-lobe; axial
furrows faint; side-lobes sloping gently from the^axial furrows
to the fulcra, thence sloping fairly steeply. Mesial furrows of
the somites distinct, and reaching to the extremities, the anterior
somites shorter than the posterior ones.
Fyyidiuin sub-semielliptic, convex, greatest width nearly equal
to the greatest width of the thorax, and consequently just on
one-third greater than the thoracic length. Axis very prominent,
tapering gradually, and ending just within the border at half
the anterior width, rather prominently and rounded; annula-
458 CARBONIFEROUS TRILOBITES OP AUSTRALIA,
tions twenty, strongly arched; axial furrows distinct; side-lobes
strongly convex, their greatest width equals the anterior spread
of the axis; segments fourteen, all terminating at the inner
boundary of the narrow border.
Ohs. — This very fine trilobite was described by Mr. K. Ether-
idge,* and was determined by him to be identical with his P.
dubia,j remarking, however, that the form now dealt with
differed from the Queensland one in the possession of eighteen
to twenty annulations in the pygidial axis, while the former had
only eight to ten. This of itself appears to the writer a sufficient
difference to justify the separation of the two forms specifically,
for, in no instance, has he found any Phillipsia, or indeed any
meinber of the Proetidse, presenting such a wide variation in the
axial annulations of its pygidium as is exhibited by these two
forms. But there are other differences between the two forms
of even greater specific importance than this one. The eyes
and front limb of the glabella of P. elovigata are relatively very
much larger than those of P. stanvelleiisis [P. dubia Eth. til.).
In the former, the facial sutures are more sinuate, the free
cheeks more depressed, eyes more squat on the cheeks, lateral
furrows of the free cheeks wider than they are in the latter. The
neck-ring and thoracic axial rings in the former are non-tuber-
culate basally; in the latter, these are tuberculate; the glabella
of the former, too, is relatively longer and straighter- sided than
is the case in the latter. The former has a relatively narrower
pygidial border, and is of a much larger growth than the other.
The two forms, in many of their features, belong to the same
typical group of Phillipsise, in which the glabellte diminish in
width anteriorly, and are sub-bellshaped, and the facial sutures
strongly sinuate in front of the eyes. The British P. Lruncatula
is a good representative of the group.
Log. and Hor. — Binge Berry, Rouchel Brook, Hunter River,
County Durham: (IjAllyn River, half a mile north-east of Gres-
ford; County Durham (Cullen).
* Mem. Geol. Survey N. S. Wales, Pal. No.5, Pt. ii., 1892, pp. 126-8.
t Geol. and Pal. Queensland and New^ Guinea, p. 215, PL 8, fig. o.
BY JOHN MITCHELL. 459
Phillipsia rockhamptonensis, sp.nov.
(Plate xlix., figs. 9, 10; Plate 1., figs. 8, 9).
Phillipsia dubia Etheridge, fil., Geol. and Pal. Queensland and
New Guinea, 1892, pp.214-215, P1.44, fig.4.
P. woodwarai Eth. fil., op. cit, p. 215, PL 4 4, fig. 6.
Complete form oblong-ovate.
*S'^. Chars. — Cephalon sub-semicircular, appearing to have been
very finely granulated. Glabella subcylindrical or conical,
widest across the basal lobes, two lateral furrows visible on the
left side, basal pair wide, deep, and very distinctly joining the
neck-furrow, basal lobes small, very prominent and sub-pyriform;
neck-furrow deep centrally and shallowed laterally by a very
mild swelling behind the basal glabellar lobes, which can barely
be said to be supplementary lobes, its lateral extensions shallow;
neck-ring stronger and more convex than any of the axial rings,
very faintly tuberculate at its base, front limb narrow, the
margin being upturned and gently thickened ; axial grooves
narrow; fixed cheeks small; eye-lobe relatively large and gran-
ulated; free cheeks fairly large, only moderately tumid, depressed
but posteriorly high and sloping steeply into the posterior
furrow, distinctly channelled under the eyes, lateral furrows
wide, lateral borders of moderate width, mildly tumid, under-
surface striated; facial sutures not distinguishable anteriorly,
and posteriorly obliquely crossing the fixed cheeks rather close
to the axial grooves; eyes of medium size, reniform, faceted,
prominent and vertically narrow, genal angles bearing Ion"-
spines.
Thorax oblong, length equal to the width of the axis and one
side-lobe, granulations microscopic, somites of the normal num-
ber, axis strongly convex, wider than one side-lobe, diminishing
posteriorly very gently and uniformly, anterior width in the
specimen under notice being 5 mm., and the posterior width
4 mm.; each individual ring has a decided anterior trend cen-
trally and tuberculate bases, ridges and valleys narrow, and on
the former the articulating seams are plainly visible; side-lobes
convex, steeply deflected from the fulcra, segmental ends strongly
faceted, axial grooves narrow.
460 CARBONIFKKOUS TRILOBITES OF AUSTRALIA,
Pygidiuin sub-semicircular or sub semielliptic, length 7 ram.,
greatest width 12mm., convex, distinct indications of granulation
present; axis prominent, strongly convex, diminishing rapidly
posteriorly, ending prominently near the margin of the border
with about one-fourth of its anterior spread, its rings are twelve;
axial grooves distinct; side-lobes convex, consisting of eight
pairs of segments, the last two pairs being rather indistinct, first
pair faceted; border of moderate width, gently tumid, continuous
to the anterior pair of segments; undersurface striated.
Obs. — This fossil was included by Mr. Etheridge with his
species Phillipsia diihia {op cit.}, but not separately described.
For so placing it, there appears to me to be insufficient reason.
That the trilobite here described is not identical with Mr.
Etheridge's P. duhia {P. stanvelleiisis mihi) may be made plain
by an explanation of their differences.
(1 ). The free cheeks and eyes are different, as are their glabella?;
for though the glabella of the present species is not perfect,
there is sufficient of it preserved to show that, anteriorly, it
neither contracts nor droops as does that of P. duhia [P.
staiivellensis).
(2). The neck-furrows of the two also differ.
(3). The axis of the former is wider than one side-lobe; in the
latter, the axis is narrower than one side-lobe.
(4). The axis of the former is much less prominent than that
of the latter.
(5). Tn the former, the length of the thorax is much greater
than its cephalic or pygidial length; in the latter, both the
cephalon and pygidium are longer than the thorax.
(6). In the latter, the anterior pleural segments of the pygidium
show distinct bifurcation; this is not visible in the former.
(7). The genal angles of the former are strongly spinate; those
of the latter are not yet proven to be so.
(8). The middle lobe of the one is wide and moderately convex;
that of the other, narrow and very convex.
(9). The angulation of the free cheeks of the former posteriorly,
and their steep slope into the furrow are very marked, but this
is not so ill the latter.
BY JOHN MITCHELL. 461
The above differences clearly separate the two species. From
all other Australian species, it is easily distinguished.
Loc. and Uor. — Rockhampton district, Corner Creek, Great
Star River, Queensland. Carboniferous.
Phillipsia stanvellensis, sp.nov.
(Plate xlviii.,%s.lO-13; Plate li., figs. 8-10).
Phillipsia dubia Eth. fil., Geol. and Pal. Queensland and New
Guinea, 1892, pp.214-215, P1.8, tig.o.
Complete form elongately ellipsoidal.
Sp. Chars. — Cephaloii sub-semielliptic,. strongly inflated, prac-
tically smooth, length 6 mm., width between the genal angles
9 mm. Glabella bell-shaped or subconical, liigh posteriorly, with
a steep anterior droop to the front limb, microscopically gran-
ulated, three pairs of lateral furrows, basal pair shallow, the
others faint; basal lobes very prominent, relatively small and
round, mesial pair subquadrate and about equal in size to the
basal pair, front pair well-defined. Neck-furrow shallow cen-
trally, but rather deeper behind the basal glabellar lobes, its
lateral extensions across the fixed cheeks shallow; neck ring
narrow, strongly arched, and its bases tuberculated, lateral ex-
tensions weak; front limb narrow, gently thickened, and turned
almost on to the front of the glabella; facial sutures anteriorly
only moderately sinuate, posteriorly oblique and crossing the
fixed cheeks in a line with the fulcra of the thoracic segments.
Free cheeks relatively large and steep, border moderately wide
and strong, lateral furrows linear and faint. Eyes of moderate
size, densely and finely faceted, subcrescentic, and rising steeply
from the shallow grooves of the cheeks just below them. Genal
angles acute, but apparently not spinate.
Thorax: length two-thirds of width approximately, and
anterior and posterior widths equal; number of somites small,
smooth. Axis very prominent, rings arched forward both cen-
trally and basally, bases also tuberculate, centrally each ring is
rather acute, which gives to the axis when viewed sideways a
mildly serrated aspect; ring-furrows rather deep, ridges high and
narrow (in decorticated specimens), greatest width 3 m-m., and is
462 CARBONIFEROUS TKILOBITES OF AUSTRALIA,
wider than the pleurse if measured horizontally. Axial grooves
linear. Side-lobes between the axial grooves and fulcra almost
horizontal, thence nearly perpendicular ; somites apparentl}^
faceted, medial grooves very shallow, ridges feeble.
Pygidiuni semielliptic, length 6 mm., width 7 mm., smooth.
Axis very prominent, made up of twelve if not thirteen annula-
tions, but not more than ten usually visible owing to the slender-
ness of the posterior ones. Posteriorly it contracts gradualh',
and ends a little short of the border with about half the anterior
width; in some specimens, the rings centrally show emargination
and other indications of having borne fine tubercles. Axial
grooves faint Side-lobes convex, consisting of eight pairs of
segments whose valleys and ridges are inconspicuous, and the
ends, as they approach the border, bifurcate, or, more correctl}^,
the sutures widen out and make the segments appear to have
free ends; the individual segments have little or no backward
curve, and the last two pairs are indicated merely by a very faint
ridge and tubercle; border continuous and relatively wide, con-
centrically and finely striated underneath, only slightly thickened
and separated from the segments by a fine suture.
06s. — This beautiful trilobite was originally described by K.
Etheridge, Junr., and with it was joined Griffithides dubiiis Ether-
idge, Senr., for reasons not explained. All the evidence avail-
able leads me to conclude that the two are not specifically the
same, and to regret that this very typical Phillipsia above-de-
scribed should have been in any way linked with Griffithides
diihius Eth. Senr. An examination of the figure given of this
latter trilobite reveals that the glabella has three continuous
lateral furrows, and a shape altogether unlike the former species;
and, further, the latter is represented as having thirteen or four-
teen axial rings, and an equal number of pleural segments in its
pygidium; while the former has only twelve and eight, respect-
ively, in the similar parts of the pygidium. These differences
place the two apart.
As may be observed (antea, under the description of P.
elonyata mihi), I have had to disagree with Mr. Etheridge in
determining that, and the above form to be specifically the same,
for which divergence of opinion reasons are given.
BY JOHN MITCHELL. 4€3
P. stanveUensis is so different from all foreign and other Aus-
tralian species that it is not necessary to detail its relationships
with them.
The outstanding features of the species are. 1. The dunce's
hat-shaped, prominent, smooth and anteriorly drooping, and
narrow glabella. 2. The slightly elevated glabellar basal lobes.
3. The moderate size, and fine but distinct faceting of the eyes.
4. The very prominent axis. 5. The shallowness of the medial
furrows, and slenderness of the ridges of the pleural segments.
6. 'J'he bifurcation of the pleural segments of the pygidium. 7.
Twelve and eight divisions in the axis and pleural segments,
respectively, of the pygidium. 8. The equality of the lengths of
head and pygidium.
Loc. a7id Hor. — Corner Creek, Great Star River, Queensland.
Carboniferous.
Phillipsia grandis Eth. til.
(Plate xlvii., fig.2; Plate 1., tigs.1-3).
Phillipsia grandis Eth. fil., Geol. and Pal. of Queensland and
New Guinea, 1892, pp.215-216.
Etheridge, Junr., Mem. Geol. Survey N. S. Wales, Pal. No. 5,
Pt. ii., 1892, PI. xxi., fig.5 (non text-fig.5, p.l28).
Etheridge, Junr., Geol. Survey of Queensland, Publication No.
260, pp.11, 12, PI. iii., tig.3.
Complete form unknown.
Sp. Chars. — What appears to be a portion of a cephalon of
this species is preserved on a rock-fragment from near Mt.
Morgan, Queensland, associated with a very nearly perfect
pygidium, free cheeks, a small portion of the anterior of the
glabella, and the greater part of the hypostome (m situ), but so
little of the glabella remains that none of its features can be
recognised; except that it was sparsely granulated, sloped gradu-
ally anteriorly, and was of unusual size. The hypostome was
large, striated, and apparently granulated; free cheeks very large,
strongly granulated, the granules in many instances joining and
producing rugosity; posterior furrows very wide and shallow;
lateral border very wide; and the undersurface bearing six or
seven concentric stride, posterior border narrow; the genal angles
464 CARBONIFRItOUS THILOBITES OF AUSTRALIA,
appear to have been spinate. The length of the cephalic shield
was approximately 30 mm.
Thorax: greatest width approximately 45mm., length 27mm.,
finely granulated. Axis moderately convex, and had a spread
of 15 mm., and hence equal to one side-lobe. These lobes were
fairly convex, greatest width 15 mm., and the mesial furrows of
each pair of the segments were wide and shallow along the
articulating faces (ridges), the sutures are plainly visible in
decorticated specimens.
Pygidiuyn: greatest width 41 mm., length 34 mm.; serai-elliptic,
moderately convex; axis mildly convex, tapering very gradually
posteriorly, ending bluntly at the inner edge of the border, much
narrower than one side-lobe, practically two-thirds as wide as
one side-lobe, the width being 11 : 15, rings apparently eighteen,
strong, densely and finely granulated, valleys narrow : axial
furrows narrow and distinct; pleurae moderately convex; ribs,
fifteen pairs, reaching to the thickened outer margin of the
border, and having a decided backward curve in the portions
traversing the border; mesial valleys fairly deep, ridges strong,
and bearing two or more rows of closely placed granules, and
articulating sutures plainly visible^ each succeeding pair gradu-
ally inclining posteriorly, border wide but ill-defined, not con-
tinuous and concave, margin thickened and upturned.
Q})s — The first fragment of this gigantic Carboniferous trilo-
bite was briefly described by Mr. Etheridge, Junr.,* and for it
he suggested the specific name gratidis. Subsequently,! he de-
scribed a pygidium from New South Wales, which he placed
specifically with the Queensland pygidial fragment. Very
reluctantly, I have to disagree with this latter determination of
Mr. Etheridge, and am compelled to give the New South Wales
fossil separate specific rank. A recent discovery of a nearly
perfect tail, portions of a cephalic shield, and thorax, in the Mt.
Morgan area, Queensland, has much simplitied the task of
separating the two forms. This recently discovered specimen
* Geol. and Pal. Queensland and New Guinea, 1892, pp. 215-216,
t Mem. Geol. Survey N. S. Whales, Pal. No.5, Pt. ii., 1892, p. 128, text-
fig. o, and PI. xxi., fig. 5.
BY JOHN MITCHELL. 465
was submitted by Mr. Dunstan, Chief Government Geologist,
Department of Mines, Brisbane, to Mr. Etheridge, who described
it briefly,* placing it with P. yrandis. This same specimen has
been used for the above description. It will be seen that, in
the description of the parts of the fossil, we are not in agreement,
our dimensional estimates, strangely, differing rathei- widely.
The dimensions arrived at by myself are :~
Cephalic shield, length 30 mm.
Length of thorax, 20 mm.
Greatest width, 45 mm.
Length of tail, 34 mm.
Greatest width, 44 mm.
Total length of trilobite, 84 mm., or nearly 3i inches, A giant, cer-
tainly, among Carboniferous trilobites.
The reasons for separating the pygidium from Swain's Selec-
tion, near Carroll, N.S.W., from the Queensland forms, are fully
entered into under the re-description of the New South Wales
form, under the name of PJiillipsia robusta mihi.
The distinguishing features of the present species are — 1. Its
great size. 2. The largeness of the free cheeks, their ornamenta-
tion, and great width of their borders. 3. The relatively narrow
pygidial axis. 4. The mild tuberculation of the bases of the
thoracic and pygidial axial rings. 5. The slight tuberculation
of the pleural segment at the inner margin of the border, and
their claw-like shape, as they cross the border. 6. The plainly
visible sutures along the articulating ridges of the pleural seg-
ments, and the fine and close granulation of these ridges. 7. The
strongly ridged and furrowed somites of the pygidium.
As Mr. Etheridge has pointed out, it is uncertain still whether
this trilobite belongs to the genus Phillipsia or to Griffithides,
but the evidence available indicates the former.
Loc. and Uor. — Crow's Nest, near Mt. Morgan, Queensland.
Carboniferous.
Phillipsia woodwardi Eth. fil.
Phillipsia woodwardi Eth. fil., Geol. and Pal, Queensland and
New Guinea, 1892, p.215, PI, vii., figs.ll, 13; PI. viii., fig.6; and
PI. xliv., figs.5-6.
* Geol. Survey Queensland, Publication No.260, 1917, pp. 11-12, PI. iii.,
fig. 3.
466 CARBONIPKROUS TRILOBITES OF AUSTRALIA,
Oriffithidea seminiferus vslv. ausiralasica Eth. fil., (in part) op.
cit., p.216.
(PL xlvii., fig.9; PI. xlix., figs. 1-8, 13-14; PI. 1., 6gs.lO-ll; PI. li.,
figs.12-14; PI. lii., fig.l; PI. liii., figs.8-9).
Here is Mr. Etheridge's description of this species (loc. cit.): —
"/S/>. Char. Glabella round, without any lateral inflection of the
margin, moderately convex in the middle line, and a little arched
posteriorly; neck furrow strong and deep, with more or less com-
plete basal furrows; anterior furrows present, but faint; anterior
border thick and upwardly turned, leaving a wide depression
between it and the front of the glabella."
In his observations, Mr. Etheridge remarks: — "The pygidium
referred to is much larger than that described as P. dubia, but
otherwise resembles it." In the text, there is nothing to show
what pygidium is here referred to, for no description of it is
given; but no doubt he refers to the pygidium illustrated in
Plate xliv., fig. 6; and if so, the determination, in my opinion, is
incorrect. This pygidium belongs to the trilobite described by
me as Phillipsia rockhamptonensis, and is shown in PI. 1., fig.8.
The following is a more amplified description of the type-
cephalon of the species. Outline subsemicircular, surface gran-
ulated. Glabella wider across the basal lobes than it is long,
very convex both transversely and longitudinally, the front lobe
(the portion anterior to the anterior pair of the lateral furrows)
semicircular, highest between the mesial pair of lateral furrows.
Three pairs of lateral furrows present, anterior and mesial rela-
tively wide, shallow and straight, basal pair wide and deep;
anterior and mesial lobes narrow, basal pair large, subfusiform,
much lower than the glabellar centre, overhanging the axial and
neck-furrows; limb very large, border strongly thickened and
upturned, furrow wide and deep. Neck-furrow deep; neck-ring
strong, convex, sharply curving anteriorly, as it approaches the
axial furrows; sides gradually thickening towards their origin.
Frontal limb very wide, border strongly thickened and upturned,
and separated from the glabella by a wide and pronounced
furrow, strongly expanded antero-laterally. Facial sutures
anteriorly directed outwards at an angle of about 25°. Fixed
BY JOHN MITCHELL. 467
cheeks appear to have been small, and strongly tumid. Axial
furrows deep. Other parts not known.
As Mr. Etheridge pointed out, this head-shield bears a strong
resemblance to that of some Proeti. The enormous limb is its
most striking feature. The discovery of a complete specimen
will be of much scientific interest. It may be noticed here that
neither of the figures, PL vii., fig. 13, PL xliv., figs. 5, 6 {loc. cit.)
are correct representations of the fossils they are intended to
depict. For example, in the case of the first of these, the
glabella is much too blunt in front; and supplementary lobes,
which are present on the fossil, are not shown in the figure.
With reference to figs. 5 and 6 of PL xliv., in the case of fig. 5,
the glabella is represented as of a battle-axe shape, instead of
being rather quadrate, or only gently rounded in front, and the
glabellar basal lobes are exaggerated in size. In fig. 6, the axis
is shown terminating much too short of the border, and, in the
pleural lobes, too many segments are shown. Photographs of
the original specimens, from which the drawings of the figures
above referred to were made, will be found in the Plates of this
paper.
At present, I hesitate to recognise the portion of a cephalon
determined by Mr, Etheridge to belong to this species, and re-
presented in PL vii., fig. 13, for the following reasons. The
glabella of this specimen is much more convex and narrower in
front than the type-specimen. Further, this glabella is just as
long as wide across the basal glabellar lobes. On the other
hand, the typical glabella of the species is much wider across
these same lobes than it is long. The glabella in question is not
correctly represented by the figure above referred to, as will be
seen from the photos of it now produced.
Since Mr. Etheridge described this species, much additional
material has been secured, and this has been placed with me by
Mr. Dunstan, Chief Government Geologist, Brisbane, for classi-
fication. Included in the collection is a large number of pygidia,
all possessing similar specific features; and these were collected
from the same localities from which all the known cephalic frag-
ments of the species have been obtained. In pait, owing to the
468 CARBONIFEROUS TRILOBITES OF AUSTRALIA,
association in this way of these heads and tails, and because the
great convexity of the pygidia referred to, which would indicate
the possession of a very convex cephalon by the species they
belong to, and, further, because no other head-shields reasonably
preserved occur in association with these remains, with whicli
they appear to have near relationship, I liave determined to place
them with Mr. Etheridge's }\ tvoodwardi, and, for this reason,
the species is redescribed hereunder.
Complete form not yet certainly known, but it may be assumed
to have been oblong-ovate.
Cephalon sub-semicircular, highly inflated, surface granulated
throughout. Glabella subquadrate, wide and mildly rounded in
front, strongly convex, highest between the mesial lobes, sloping
strongly anteriorly, and overhanging the furrow of the limb in
front; three pairs of glabellar furrows present, the anterior and
mesial pairs rather wide, shallow and straight; basal pair wide
and deep; anterior and mesial lobes narrow; basal pair large,
subfusiform, much lower than the glabellar superficial centre,
overhanging the axial and neck-furrows latero-posteriorly; limb
very large, border strongly thickened and upturned, furrow wide
and deep; neck-furrow wide and deep; neck-ring strong, convex,
bases strongly curving anteriorly under the glabellar basal lobes;
supplementary lobes present, though tjot conspicuous in the most
typical specimen. Facial sutures anteriorly directed outwards
at an angle of about 25°. Fixed cheeks would appear to have
been of moderate size, and posteriorly strongly inflated. Free
cheeks fairly large, strongly and thickly granulated, especially
on the part between the lateral and ocular furrows; this part,
too, is high, and rather acutely angulate postero-laterally, flat
superficially, laterally and posteriorly steeply falling into the
lateral and posterior furrows, which are wide and deep; border
much thickened and upturned, striated, posterior border (exten-
sion of the neck-ring) strong, genal angles apparently blunt;
eyes of moderate size, apparently crescentic and faceted, some-
what sunk into the ocular furrow.
I'horax : for certain not known, but the following is the de-
scription of one attached to a pygidium identical with a number
BY JOHN MITCHKLL. 469
of others assumed to belong to this species : suboblong, gi-aii-
ulated, longer than the p\^gidium, all the segmental ridges appear
to have borne tubercles; number of segments normal. Axis
prominent, ring-bases non-tuberculate, with a strong anterior
inclination centrally, ridges and valleys well defined, apparently
as wide as one side-lobe. Side-lobes strongly convex and steep
between the fulcra and margins, all parts densely granulated,
segments anteriorly angulate at the fulcra; mesial furrows wide,
ridges relatively narrow and faceted, and mildly thickened
posteriorly.
Pygidivm very strongly convex, length to width approximately
2 : 3, granulated; axis prominent, evenly arched, anterior spread
about equal to the greatest width of one side-lobe, posteriorly
diminishing in width very gradually, and ending prominently
and bluntly a little short of the bordei-; number of rings thirteen,
and each bore a row of small tubercles quite conspicuous on
unweathered specimens; axial grooves wide and deep. Side,
lobes convex, consif^ting of eleven pairs of segments, the lidges
of each pair bearing rows of small tubercles varying from two or
three on the shorter, to eight or perhaps nine on the longer ones.
Border continuous, mildly thickened, convex and steep.
Ohs. — At the outset, I admit that the pygidia here joined with
Mr. Etheridge's tj'pe-cephalon of the species may yet prove to
be a wrong conclusion; but should it happen so, T am inclined
to believe, too, that the heads (PI. li., figs. 1*2, 13) will also prove
not identical with the type one; but should that happen, I am
fully satisfied that these latter heads, and the tails here dealt
with, will prove to ijelong to the one species. The heads now
joined with the typical one possessing the very characteristic
frontal limb, cannot, for certain, in the absence of this limb, be
said to belong to the species.
While completing the above description, the specimen No.712,
belonging to the Queensland Museum, Brisbane, is before me;
and the trilobite fossil remains on it consist of two pygidia, one
thorax and pygidium conjoined but incomplete, a fragmentary
head in intaglio, and a part of a free cheek. (On the same
specimen, occurs the tail of a Brachymetopus, the first recorded
35
470 CARBONIFEROUS TRILOBITES OF AUSTRALIA,
from Queensland). These fossils weie those which chiefly served
Mr. Etheridge to found his varietal species Griffithides aemini-
ferus var. australasica on. After a close study of these remains,
I am compelled to dissent from his conclusions concerning them.
They cannot belong to the above species, because the glabella
possesses supplementary basal lobes, a feature by some palaeon-
tologists thought to be of generic significance, and which G.
seminiferus does not possess. Again, these pygidia have thirteen
rings in their axis, and eleven pairs of segments in the pleursp.
In these parts, G. seminiferus has twelve and nine respectively.
These differences alone would render the specific identity of the
two forms invalid, but these are not the only differences between
them. As far, then, as the fossils under present consideration
afford evidence, Mr. Etheridge's varietal species fails. The part
of the cephalon on this same specimen of rock is very imperfect,
but it shows, on the glabella, similar but clearer granulation,
because the granules are not worn off by weathering, similar
glabellar basal and supplementary lobes lo that and those
respectively found on the cephalons here determined to belong
to Phillipsia vjoodwardi. The only features not visible on the
glabella of the fragmentary cephalon, to make its identity with
the latter certain, are the mesial and anterior glabellar furrows;
but these are barely visible on some glabellse of P. woodwardi.
Lastly, the pygidium represented in Plate viii., fig. 6 (oj). cit.),
which was considered by Mr. Etheridge to be a normal pygidium
of his P. duhia (P. stanvellensis mihi) is here joined with the
present species by me. It is undoubtedly inseparable from the
other pygidia now joined with the species above described, and
certainly is not at all closely related to the pygidia of P. stan-
vellensis mihi [P. duhia Eth. fil., op. cit., PI. viii., fig.o).
Mr. Etheridge has already called attention to the Proetus-like
aspect of the cephalon of the present species. The singular form
of this part of the fossil makes it unnecessary to point out in
detail the features which separate it from other species of
Phillipsia.
Log. and Hor. — Stoney Creek, Stanwell; Crow's Nest and
Trilobite Ridge, Mt. Morgan, Queensland, etc. Associated wit |i
BY JOHN MITCHELL. 471
Brachymelopus dniififani. Middle or Lower Carboniferous pro-
bably.
Phillipsia morganensis, sp. nov.
(PI. xlix., figs.ll, 12; PI. li., 6g.l).
.S';;. CAa/-6\ — 6V/?/irt/o?i sub-semicircular, very finely granulated,
greatest length and width 8 mm., and 16 mm., respectively.
Glabella subconical or sub-bellshaped, mildly convex, finely and
densely granulated, longer than wide ; three pairs of lateral
furrows distinctly shown, basal pair very deep and wide, anterior
and mesial pairs short and only gently curved posteriorly:
anterior and mesial lobes of moderate size, basal pair fairly
large, with small complementary lobes present; neck-furrow deep
and wide, the extensions laterally also fairly deep and wide;
neck-ring strong, moderately convex, strongly curving anteriorly
at its bases; facial sutures anteriorly very straight, posteriorly
short and oblique, frontal limb narrow and close to the glabellar
front; fixed cheeks very small, with small eye-lobes. Free cheeks
rather large, only moderately tumid, finely granulated, both
borders and lateral furrows large, the former being moderately
thickened, finely granulated on the upper and striated on the
undersurface, the latter being shallow, ocular furrow wide and
shallow. Eyes of moderate size, finely faceted, vertically narrow,
and decidedly higher behind than in front. Genal angles obtuse.
Hypostome shield- like, narrow, not alate, posteriorly glossiform
and corrugated, no striae visible, widest at two- thirds of its length
from the front edge.
Thorax not known in a complete state, but one, on which a
pygidium and a free cheek rest, shows the following features —
moderately convex, finely granulated. Axis moderately convex,
diminishing in width and height gradually posteriorly, wider
than one side-lobe, rings strong, valleys narrow and having an
anterior inclination, bases non-tuberculate, axial furrows linear.
Side-lobes not well shown, but the segmental ridges and furrows
were strong and deep respectively.
Pygidium widely triangular, mildly convex and granular,
length 7 mm., and greatest width 12 mm. Axis mildly and
evenly convex, anterior width equal to that of one side-lobe,
472 CARBONIFEROUS TRILOBITES OF AUSTRALIA,
diminishing gradually in width and height posteriorly, and end-
ing short of the margin inconspicuously with about half its
anterior width, rings apparently eleven; ridges not prominent,
furrows also faint, some of the anterior ones have centrally a
backward trend. Axial furrows linear. Side-lobes convex, con-
sisting apparently of nine pairs of segments (eight only are
visible on the best available specimen), front pair faceted, ridges
and furrows fairly distinct, and having successivelv onlv a
moderate backward trend.
Obs. — This trilobite belongs to the ty^pe oi Phillipsia which
had the glabella narrower in front than posteriorly, which seems
a common characteristic among the Australian members of the
genus, and, in this respect, shows a closer relationship to the
typical genus of the Proetidse, than do the Phillipsiie of Europe,
and, perhaps also, America. A singular feature in the present
species is the form of the hypostome assumed to belong to it.
None similar to it has come under my notice. The one nearest
to it is one figured by Dr. H. Woodward* with other illus-
trations of Phillipsia eichivaJdi, but evidently not belonging to
that species. 'J he straightness of the anterior branches of the
facial sutures, and their mild outward divergence as tiiey reach
the frontal limb, are striking features of the species. A study
of the description and illustrations of the species will render its
separation from other Australian Phillipsise simple. In contour,
the glabella is not unlike what that of the glabella placed with
P. woodtvardi (PI. xlix., fig. 7) would be, were it not so convex;
but, in the former, the glabella is longer than it is wide across
the basal glabellar lobes; in the latter, these dimensions are
equal. No foreign species has come under my notice witli which
it seems necessary to compare it.
Loc. and Hor. — Trilobite Ridge, Mount Morgan, Queensland.
Carboniferous (Lower'?).
Phillipsia connollii, sp.nov.
(PI. li., figs.4-7).
Phillipsia grandis Eth. fil. (MS.).
Only an incomplete pygidium known.
* Mon. Brit. Carb. Trilobites, 1883-4, PI. iv., fig. 7.
6Y JOHN Mitchell. 4/3
Sp. Chars. — Pyyidiuiu semielliptic, moderately convex, and
microscopically granulated. Axis very prominent, consisting of
ten, possibly eleven rings, not as wide as one side-lobe, evenly
arched, and slightly flattened dorsally, ending at the border very
prominently and rather acutely, ridges and valleys of the rings
quite distinct. Axial grooves shallow. Side-lobes mildly convex,
consisting of eight or nine pairs of segments, -which are only
very gently directed backward, and terminating at the border,
except in the case of the anterior pair, which encroaches on it.
Mesial furrows and the ridges of the segments moderately defined.
Border proportionately wide and continuous, mildly convex,
horizontal, and apparently wider laterally than behind, separated
from the ribs by a shallow furrow. Greatest width 13 mm.,
length 9 mm., anterior width of axis 3 mm.
Obs. — Thispygidium was placed by Mr. Etheridge, Junr., with
his Fhillipsia grandis, but to this species I find it has no close
relationship. In its wide border, and the number of axial rings
and pleural segments, it bears strong resemblances to our
Gri(tithides conveoncaudatus, but its axis is much more prominent
and relatively narrower, than is that of the latter. The axial
and pleural divisions are more clearly defined in the former than
they are in the latter; the border in the former is ilat, and in the
latter very steep. The shape or outline of the pygidia of these
species is very different. In number of axial and pleural
divisions, this species is related to both i". cordteri and /-*. staii-
reUensls, but not otherwise. In possessing very prominent and
relatively nai-row axes, P. stanvellensis and the present species
agree. I am not aware of any foreign Carboniferous trilobite
having a pygidium closely resembling this one.
Named after Mr. Connolly, wlio discovered it.
Loc. and llor. — Gascoigne River, AN'est Australia (Connolh').
Carboniferous.
Phillipsia kouchklensis, sp.nov.
PkUiipsui sp., ind. {a), Eth. til., Mem. Geol. Surv. N. S. Wale.s,
Pal. No.5, Pt. ii., 1892, p.l29, PL xxi., tigs. 6, 7.
(PI. xlviii., figs. 4, 5, 6, 7).
474 CARBONIFEROUS TRILOBITES OF AUSTRALIA,
Cephaloii and thorax unknown.
Sp. Chars. — Py^ic<t?AWi sub-semicircular, width 15 mm., length
11mm.; smooth and moderately inflated. Axis moderately
convex, tapering very gradually, and terminating bluntly with
a spread rather greater than half that of the first ring; anterior
width about equal to that of one side-lobe; consisting of eighteen
rings, some of which show traces of having been very mildly
tuberculated; axial furrows faint. 8ide-lobes convex, with no
decided fulcra, segments thirteen, or doubtfully fourteen, mildly
recurved, not crossing the border; border of moderate width,
steep, smooth, and separated from the pleural segments by a fine
suture.
Ohs. — This pygidium was briefly described by Mr. Etheridge
{loc. cit.), but not specifically named, though he stated that it
possessed certain features which clearly indicated that it was
a new species. It differs from all other species of Phillipsia and
Griflithides known to me.
Loc. and IIor.~ Binge Berry, Rouchel Brook, near Muswell-
brook: County Durham.
Phillipsia dungogensis, sp.nov.
Phillijjsia sp. ind. {b), Eth. fil., Mem. Geol. Surv. N. S. Wales,
Pal. No.5, Pi. ii., 1892, p.l29; PI. xxii., fig.14.
(Plate xlvii., figs.6, 7).
Complete form unknown.
Sjy. Chars. — Fygidium sub-semicircular, very moderately con-
vex, finely granulated throughout, greatest width 22 mm., length
16 mm. Axis mildly convex, consisting of fourteen annulations,
the furrows and ridges of which are respectively shallow and
low, diminishing very slightly in width posteriorly, the end
being wide and rounded, bases of the rings not tuberculate,
anterior width less than one side-lobe: axial furrows fairly dis-
tinct. Pleui-ie gently inflated and sloping from the axial furrows;
segments twelve pairs, which are very completely fused, the four
anterior pairs reaching to the outer margin, medial furrows wide
and shallow, ridges not prominent and showing no sutures :
border narrow and continuous.
BY JOHN MITCHELL. 475
Obs. — This is another of the pygidia described by Mr. Ether-
idge, Junr., {loc. cit.) but which he did not specifically name.
It resembles no other Australian Carboniferous form at all
closely; and, so far as I have been able to ascertain, it stands
apart from the pygidia of foreign species of the genera Phillipsia
and Griffithides.
Loc. and Hor. — Greenhills, near Dungog; County Durham.
Carboniferous.
Genus Griffithides Portlock, 1843.
Geol. Report, Londonderry, &c., p. 3 10,
The following is Portlock 's original description of the genus —
''Cephalothorax semi-oval, longitudinal: glabella strongly marked
and gibbous, rounded in front, narrowed posteriorly into an ob-
solete neck with a furrow more or less distinct on each side.
Cheeks: triangular spaces very slightly convex. Wings either
ending in an angle posteriorly or prolonged backwards in a
tlattened spine. Eyes near the axis, not large, lunate, smoothC?).
The minute neck tubercle sometimes present. '
" Thorax. — The pleuripedes are compound, in number nine, or
with the neck segment ten."
^^Fygidium. — Fully developed and strongly resembling that of
Phillipsia."
General A. W. Vogdes {loc. cit.) gives the following brief
summary of the generic features of this genus: — 1. Glabella
short, tumid. 2. No short lateral furrows on the glabella. 3.
Basal lobes distinct. 4. Eyes small, placed close on the glabella,
reniform. 5. Axis of pygidium has 10 to 17 segments.
Griffithides convexicaudatus, sp.nov.
(PI. xlvi., tig.l3; PI. xlviii., figs.1-3; PI. lii., figs.5, 6;.
Sp. Chars. — Complete form suboval with straight sides
Cephalon sub-semicircular, apparently finely granulated and
strongly inflated. Glabella subpyriform, tumid; basal furrows
faint; basal lobes relatively small; neck-furrow shallow, its lateral
extensions deeper, and communicating with the lateral furrows
of the free cheeks; neck-ring wider than the axial rings of the
476 CARBOMFEtlOtS TKILOfilTES OF AOsTRALtA,
thorax, its lateral extensions narrow but prominent. Axial
grooves faint Fixed cheeks very small and high; palpebral lobe
high, narrow, convex, and very faintly separated from the basal
lobe. Free cheeks iiigh, sharply rising from the narrow lateral
furrows, lateral border narrow, thickened and raised. Eyes
reniform, short fore and aft, high and prominent, separated from
the upper and flattened part of the cheeks by a faint sulcus, ap-
parently faceted. Facial sutures anteriorly follow the course of
the axial grooves, posteriorly oblique passing out in a line with
the fulcra of the thoracic pleurae. Limb narrow and pressed
to the front of the glabella. Genal angles bear very short spines.
Thorax fairly convex, finely granulated, rectangular, length
practically two-thirds of greatest width, anterior and posterior
widths approximately equal, somites nine. Axis prominent,
widely and evenly arched transversely, width or spread through-
out about equal, the last two rings only being slightly contracted,
a little wider than one side-lobe, centrally the rings have a gentle
forward arch, and bear a row of inconspicuous granules, bases
non-tuberculate: axial furrows shallow. Side-lobes sloping very
gently from the axial furrows to the fulcra and thence fairly
steeply, median furrows of each segment narrow and shallow,
but reaching just to the margin, ends faceted.
Fygidiiun sub-semicircular, evenly and very convex, finely
granulate, length equal to length of thorax, and to about three-
fourths of its own greatest width (9:13): axis convex, bearing
eleven annulations, diminishing posterioily very gradually in
width and prominence, ending bluntly and rounded a little short
of the border. Its length equals seven-ninths of the pygidial
length, annulations faint, especially towards the distal end.
Side-lobes strongly convex, anteriorly having a spread approxi-
mately equal to that of the axis, possessing eiglit ribs, only the
first three pairs being at all conspicuous: all, except the first
pair, stop at the faint furrow separating them from the wide,
convex, smooth border.
Obs. —The individual, whicli served for the above description,
measured 25 mm., of which the cephalon was 7 nnn., the thorax
and pygidium 9 mm. each; width of thorax 14 mm. The speci-
fiY JOHN MITCHELL. 477
men was not quite complete, being minus part of the right front
quarter of the cephalon.
This fossil agrees very closely with one described by De
Ivoninck from the Upper William River,* and determined by
him to be Grijfithides (Phillipsia) eichwaldi. 'Jhe total lengths
of the two fossils exactly agree, as also do the widths of the
thoraces; but there are some discrepancies in the dimensions of
the separate parts. De Koninck gives 10 mm. and 8 mm., re-
spectively, for thorax and pygidium of his specimen; whilst 9mm.
is the length of each of these parts in the specimen under
review. The two hav^e nearly the same number of annulations
in the pygidial axis, and probably the same number of pleural
ribs, a similar wide pygidial border: also the same relative
length to width of their pygidia, and an identical frontal pro-
jection of the glabella on to the border. In the characters of
granulation and of thoraces and pleural ribs of the pygidia, thev
also agree. If, at this, the discussion of their relationship
stopped, the identity of the two would have to be accepted as
conclusive, as I believe it actually to be. But against these
agreements have to be placed some important differences, which
make their specific identity difHcult to reconcile. For instance,
T>e Ivoninck states that his specimen has only eight thoiacic
segments, that tlie anterior annulation of the thoracic axis has a
widthf of 3 mm., and not one of the annulations has a width
less than 1 mm. The present specimen has the normal nine
somites, and the widest annulation of the thoracic axis does not
exceed 1 mm. De Konincks figure of his specimen shows it to
have had a \'ery globular frontal glabellar lobe, and, in this, differs
from the one above described. Again, De Konincks text does
not agree with his illustiation. The former indicates his speci-
nien to have eight thoracic somites, and the pleural lobes to have
a width equal to that of the thoiacic axis; but his figure shows
nine somites, and pleural lobes much narrower than the axis.
These important discrepancies make it impossible to accept his
" Foss. Pal. Nouv. (dalles du Sud, Pt. i., 1870, pp.278-9, PI. xxiv., tig.8.
I The writer assumes the width of tlie axial rings to be their measure-
ment alonj.; the lonifjtudinal line of the axis.
478 CARBONIFEROUS TRILOBITES OF AUSTRALIA,
description as having any scientiHc value, for there is no evidence
to show whether his text or figure is correct, the tj^pe specimen
having been destroyed in the Garden Palace fire of 1882.
The next consideration is whether the present fossil, or even
De Koninck's, is specifically identical with P. eichwaldi Fischer.
Accepting the types given by Dr. H. Woodward* as fully
reliable, the author does not believe it a difficult task to prove
the aegative.
1. Dimensions. — The relative lengths to widths of the parts of
the author's fossil and that of P. eichwaldi Fischer, are, respect-
ively, for cephalon 14 : 28 and 17 : 27; thorax 17 : 28 and 17 : 31 ;
pygidium 9:13 and 1:1.
The dimensional differences, as these measurements show, place
the fossils apart; but these are not nearly so important specifi-
cally as the following. The pygidium of F. eichwaldi is semi-
elliptical, and the length equal to the width: that of the author's
is sub-semicircular; that is, if the centre of the junction of the
second axial ring with the third be taken for centre, the portion
of the pygidium posterior to this forms a semicircle; and the
length is only two- thirds of the width, approximately. There are
up to sixteen rings in the axis, and twelve to fourteen pleural
divisions in the pygidium of P. eichwaldi; while, in the local
one, these divisions are eleven and eight, respectively. Then, in
the former, the pygidial border is depressed; in the latter, tlie
curve of convexity of the pleurae continues uninterruptedly across
the border to the outer edge. In the former, also, the genal
spines are long; in the latter, very short, only extending past
the first thoracic segment. These differences are sufficient to
prove that the author's specimen is not F. eichwaldi Fischer, and
the same conclusion may be drawn in respect to De Konincks
fossil.
In several respects, this species resembles G'rij/ithides ylobiceps
Phillips; but the proportionate lengths of the thorax and pygidium
of the latter are different from those of the former; as also is the
proportionate length of the cephalon to these same parts. In
the former, there is no lobe connecting the eye-lobe with the
* Mon. British Trilobites, Pt. i., 1883.
BY JOHN MITCHELL. 479
glabella, though, indeed, tlie eye or palpebral lobe is very close
to, and only faintly separated from, the basal lobe of the glabella.
The eye, too, is relatively larger, the pygidial border is wider,
and the glabellar frontal lobe less globular ; the free cheek-
border and glabellar limb narrower, and the eyes less prominent
and more posteriorly situated in the former than in the latter.
Tiie British species, too, was much larger than the New South
Wales one.
The specific name was chosen for it because of the strong con-
vexity of its pygidium.
Log. and I/or. —Glen WilUam, about two miles from Clarence-
town, Parish Parr, County Durham. Lower Carboniferous.
Griffithides sweeti Eth. fil.
Grijfithides siveeti Eth. fil., Proc. Linn. Soc. N. S. Wales, (2),
Vol. ix., Pt.3, pp.5-28-9, PI. xxxix., fig.3, 1894.
(PI. liii, figs.1,2).
Through the courtesy of Mr. G. Sweet, of Melbourne, the
type-specimen of the species is before me, and I regret that,
owing to the incompleteness of its cephalon, it fails to elucidate
certain difficulties met with in considering the thoraces and
pygidia from the Mt. Morgan area, assumed to belong to
PhiUipaia woodtrardi Eth. fil., or to settle whether G. siveeti Eth.
fil., and that species are not identical. All the numerous pygidia
obtained from the Mt. Morgan area, except those which belong
to P. siaitvellensis, P. rockltatnptonensis, and P. inoi-yaiietLsis.
have thirteen axial, and eleven pleural divisions; and, aftermost
careful and repeated inspection, 1 have to conclude that Mr.
Etheridges G. sweeti has the same number of divisions in the
axis and pleurie of its pygidium; also the same kind of pygidial
border and furrow: and, in fact, its pygidium cannot be sepa-
rated from pygidia which are the most plentiful in the Mt.
Morgan distiict, and which I have tentatively considered to
belong to P. woodwardi, or at least to those cephalons represented
on PI. li., figs. 12 and 13, l)ecause these, too, were the most
plentiful cephalons occurring in the same area. As regards the
head-shield of G. sweeti., sufficient of it has not been conserved
480 CARBOKiFEKOUS TRlLOBlTES OF AUSTRALIA,
to enable one to say positively whether its features place it with
the Phillipsia or the Griffithides type; but, except for the un-
certainty of the presence on its glabella of the normal lateral
furrows of the Phillipsian genus, it does not differ from the
head-shields above referred to, which are considered to belong to
P. looodwardi Eth. fil. Further, it will be found that G. sweeti,
as far as present evidence admits, cannot be separated from
Grlffitliides semi)iifenis var. australasica Eth.fi]. This, I think,
will be con(;eded after the description and figures of the latter
have been studied in conjunction with my remarks on the latter,
under P. iroodivardi: and an examination of Plate li., fig. 14^
which is a photo of Mr. Etheridge's type specimen, which shows
two tails, the greater portion of a thorax and tail conjoined, an
intafylio of a portion of a head, and, between the two tails and
on the left top corner, is a view in relief of this intaglio.
We are up against a problem here, wliich can be solved only
Ijy the discovery of better material: and when it is solved, it
seems to me that a new genus or subgenus will be needed for
the reception of tliis trilobite with the ten thoracic somites.
The following is a fuller description of the species than is
given bv Mr. Etheridge It will be seen fronj the photograplis
of this species now given, that it was not as correctly figured as
it might have been.
Complete form suboval.
^}i. Chars. — Cephalon incomplete, apparently granulated
throughout, with grHiiules of uniform size. Glabella incomplete,
moderately tumid, mesial and anterior furroAvsnot visible (though
there appears to be a faint trace of the mesial pair): basnl
furrows deep, joining the neck furrow: basal lobes fairly large,
suboval: supplementary lobes of moderate size, and suboval :
neck-furrow wide and fairly deep: neck-ring stronger than any
of the axial rings of the thorax. Fixed and free (;heeks absent.
Thorax consisting of ten segments, finely and evenly gran-
ulated: axis prominent, diminishing posteriorly very little in
prominence, and barely at all in spread: each ring, except the
last, has centrally a slightly forward direction, the last ring is
stron<ier than the others, non-tuberculate; axial furrows shallow.
BY JOHN MITCHELL. 481
Side-lobes, between the axial furrows and fulcra, rising verv
gently, and thence are depressed almost at right angles, seg-
mental ends strongly faceted, and the ridges and vallevs strong
and deep respectively, segments strongly angulate at the fulcra,
the whole thoracic surface was finely granulated.
Pygidium subelliptic, finely granulated, strongly convex: axis
prominent, rings thirteen, the anteiior one being somewhat over-
lapped by the last thoracic one, decreasing in spread gradually and
ending prominently at about half of its anterior spread, a little
short of the border; axial furrows distinct, side-lobes stronglv
convex, granulated, consisting of eleven segments, each verv
gently and increasingly curving posteriorly, segmental ridges
strong, valleys deep; -border entire, steep, very finely granulated,
middle thickened and relatively wide, separated fi-om the pleural
segments by a narrow furrow, accentuated by punctations at
each segmental end.
Ghiffithides dubius Etheridge Senr.
Griffithldes dubius Etheridge Senr., Qiuu't. Journ. Geol. Soc,
1872, Vol. xxviii., p.338, PI. xviii., fig.7.
Fhillipsia dubia Eth. fil , Geol. Pal. Queensland and New
Guinea, 1892, pp.2U, 215, PI. 7, fig.l2.
(PI. liii., fig,7).
The original description is as follows: — 'vHody elongated, oval,
length about twice the width, sides parallel. Axis width of
pleurje. Thoracic segments 10 to 12. Pygidium rounded,
margins entire; axis composed of ten segments, not extending
quite to the posterior margin. Cephalic portion much crushed;
glabella small and round anteriorly, furrows indistinct. Owing
to the crustaceous test being removed, we have no means of
arriving at the condition of the original ornamentation; there
are, however, indications of tubercles upon the axis of the
pygidium."
"Zoc. Don River, Queensland. Form. Carboniferous."
The whereabouts of the type-specimen I have not been able to
discover, and, consequently, cannot add anything to the above
description.
482 CARBONIFEROUS TRILOBITES OF AUSTRALIA,
Tn the possession of ten or more segments in the thorax, it
resembles G. sweeti Eth. fil., and, certainly, in this respect, is
not a normal Grijfithides or Phillipsia. This peculiarity, too,
very decidedly separates it from P. stam'ellensis mihi {P. dnhia
Eth. .fil.).
Family PROETID^.
Genus B r a c h y m e t o P u s McCoy, 1 847.
Ann. Macr. Xat. Hist., xx., p 229, PI. xii., figs. la, \h.
McCoy's description of the genus is as follows : — ^'•Gen. Clear.
Cephalothorax truncato-orbicular: limb narrow, produced back-
wards into flattened spines: glabella smooth, cylindrical or ovate,
about twice as long as wide, not reaching within about its own
diameter of the front margin: one pair of small, basal, cephalo-
thoracic lobes, or none. Eyes reniform, in the midst of the
cheeks (1 smooth); eye lines unknown. Surface strongly gran"
ulated: one tubercle on each side of the anterior end of the
glabella, the marginal row and a circle round each eye being
larger than the rest. Body-segments unknown. Pygidium
nearly resembling the cephalothorax in size and form, rather
more pointed, strongly trilobed, and with a thickened prominent
margin; axal lobe about as wide as the lateral lobes, of about
seventeen narrow segments ; lateral segments about seven,
divided from their origin, each terminating in a large tubercle
at the margin.'" Genotype, Br. slrzehckii.
In this description, characters are included which are merely
specific. H. Woodward* supplies the following amended de-
scription of the genus : — " General form elliptical: headshield
semicircular and slightly pointed, about one-third wider than
long; glabella small, somewhat elevated, one-third the width of the
entire shield and about one-half the length, having a basal lobe
on each side, but no short lateral furrows on the glabella: neck-
furrow distinctly marked, equal in width to the posterior border
of the free cheeks: eyes small, smooth, equal to half the length
of the glabella; no facial sutures visible, only the axal furrow
surrounding the glabella and the neck-furrow; free cheeks slightly
^ Mom, Brit, C^^rb, Tnlobites, 1883-4, pp. 46-7.
BY JOHN MITCHELL. 483
convex, nearly twice as long as they are broad, with no visible
suture separating them from one another in front of the glabella:
margin broad and slightly grooved, angles of cheeks produced
posteriorly into spines. The entire surface of the head covered
irregularly with a small bead-like ornamentation."
"Thoracic segments unknown, probably nine."'
" Pygidium consisting of a vai iable number of segments, from
ten to seventeen, according to species, the axis tapering rapidly
to a bluntly rounded extremity, each segment of axis ornamented
with bead-like granulations, ribs with a double furrow extending
nearly to the border which is smooth and rounded."'
This description also includes quite a number of features that
possess only specific significance.
(General A. W. Vogdes* summarises the characters of the
genus thus: — 1. Glabella short, tumid. 2. No short lateral
furrows on the glabella. 3. Basal lobes distinct. 4. Eyes small,
placed close to the glabella. 5. Axis of the pygidium has ten to
seventeen segments.
To this may be added —6. Facial sutures absent. 7. Axial
furrows continue round the glabella-front. 8. Ornamentation
bead like.
R. F. Cowper Reedf has suggested a genus or subgenus
{Brachymetopina) for the European forms without defined
cephalic margins, and non-spinate pygidial margins.
Brachymetopus STRZELKCKii McCoy, 1847.
Brachymetopus strzeleckii McCoy, 1847, ojk cit., p. 231, PI, xii.,
fig.l. De Koninck, Foss. Pal. Nouv. Gallesdu Sud, 1877, p.352,
PI. xxiv., figs. 10, 10a, b,c. Vogdes, Trans. Acad. Sc. St. Louis,
Vol. V. (1892), p.617. Etheridge, R., Junr., Mem. Geol. Surv.
N. S. Wales, 1892, Pal. No.5, Pt. ii., p.l24. Reed, Geol. Mag.,
N.S., Dec. iv.. Vol. X., 1903, pp.193-196.
(Plate liii., figs.3-6).
McCoy's brief description is as follows : — "»S'p. Cha?-. Glabella
* "North American Carboniferous Trilobites." Ann. New York Acad.
Sci., vi., 1888, p.70.
tGeol. Mag., N.S., Dee, iv., Vol. x., p.i96,
484 CARBONIFEROUS TRILOBITES OF AUSTRAfJA,
widest at the base, with one verv minute obscurely marked
cephalothoracic furrow at the base on each side; all the segments
of the pygidium with an irregularly tuberculated ridge along the
middle; lateral segments forming large tubercles where they join
the thickened limb, opposite each of which there is a short
slender spine projecting from the margin. "
The following is a fuller description of the species by F. ]>. C.
Reed {op. ci^.) :—" Head-shield semicircnlai', moderately convex,
with strong raised rounded border increasing slightly in width
towards the front, and separated off by a deep furrow. Genal
angles furnished with slender divergent smooth spines, less than
half the length of the head shield. At its base is a pair of small
nodular basal lobes, in most specimens quite inconspicuous.
Two large tubercles are situated in a line down the middle
of the glabella, followed by a similar median one on the occipital
segment. Occipital segment strong, rounded, separated off
by a deep furrow. On cheeks at anterior end of glabella is a
pair of large tubercles one on each side. No facial sutures
visible. Eyes prominent, reniform, less than half the length of
the glabella, distant from the axial furrows about one-third the
width of the cheeks, and about their own length from posterior
margin. Surface of head-shield, including glabella, border and
neck-segment, rather coarsely tuberculated. An indistinct ring
of larger tubercles surrounds the eyes, and a large tubercle is
situated at each end of eyes on innei- side. Thorax unknown.
Pygidium semicircular, slightly convex, with spinose maigin.
Axis broad, conical, about one-third the width of the pygidium
at front end, tapers rather rapidly to obtuse point, nearh^ touch-
ing the border: consists of 9-10 segments, of which eight rings
are distinct and completely tuberculated across : the 1st, 3rd,
5th, and 7th have, in addition, a large median tubercle. Lateral
lobes consist of six (\ seven in some) pairs of pleurae, of which
the last pair is very small: each pleura is gently curved and is
divided unequally by a strong longitudinal furrow into a broader,
raised, rounded, posterior ridge, and a narrower anterior ridge.
The posterior ridge of each pleura crosses a distinct, raised,
rounded bordei;, which surrounds the pygidium and bears a large
BY JOHN MITCHELL. 485
tubercle at the spot where it crosses, and a single median one
behind the axis. The posterior pleural ridges are prolonged into
short, recurved, equidistant, and subequal spines, projecting
beyond the margin. (In one specimen there seems to be a
median spine behind the axis. In another immature example
the anterior two or three pairs of spines are half as long as the
whole pygidium). Surface of pygidium rather coarsely tuber-
culated: the posterior ridge of each pleura bears 4-5 tubercles,
and the anterior ridge 5-6 smaller ones. The axial rings bear
each 5-7 tubercles."
" Dimensions: —
Length of head-shield ... ... ... .. 8 'Omni.
Width of head-.'^hield ... ... ... ... 4'omm.
Length of pyoidiuni ... ... ... ... 2*5 mm.
Width of pygidium 4 "0 mm.
For affinities, see op. cit.
To quote Reed further, he remarks that " McCoy gave as
generic characters the circle of tubercles round the eyes and pair
of large tubercles at the front end of the glabella, but these may
well be considered as of lower classificatory value, and likewise
the relatively greater length of the glabella as compared with
the European species. It does not, however, seem possible to
regard tlie peculiar pygidial characters in the same light, though,
as Vogdes (Trans. Acad. So. St. Louis, Vol. v, (1892), p, 617)
.says, we have many other genera of trilobites with spinose and
non-spinose representatives. The fewer number of segments in
the p\'gidium, and the rai.sed spinigerous boi'der separate it from
all the European forms. "
" The genus or subgenus Phcetonides, as now understood, is
partly distinguished for analogous reasons from the typical
Proetus; and it seems open to question whether the European
species of Brachymetojnis should not be regarded as constituting
a distinct group or subgenus for which the name of Brachymeto-
pina may be suggested."
Personally, I do not think that pygidial characteristics alone
are sufficient grounds upon which to found even a 5^ubgenus
Loc. and Hor. — Dunvegan, Burragood, and Glen William,
36
486 CARBONIFEROUS TRILOBITES OF AUSTRAF.IA,
N.S.W. (Etheridge, Junr., Cat. Austr. Foss., Camb., 1878, p.41).
Carboniferous.
Bkachymetopus dunstani, sp.iiov.
(PI. xlix., figs.15, 16: PI. lii., figs.l, 2).
Complete form unknown.
Sp. Chars. — Cephalon : length and width 3 mm, and 4 mm.,
respectively; semielliptic, surface covered with tubercles of vary-
ing size, only mildly convex; glabella conical, densely and evenly
granulated, very mildly convex, basal furrows well defined and
joining the neck-furrow, basal lobes small, greatest width about
one-fourth that of the cephalon, length about half that of the
cephalon; the surrounding axial furrow relatively deep, cheeks
gently convex, bordering the furrow surrounding the glabella;
they bear eleven tubercles of uniform size, and follow a course
inside of the eyes to the posterior margin: from the front of the
eyes there branches from this main bead-like chain of tubercles
two other sets (one on each side) of five each, which are arranged
along the outer boundary of the eyes; the one or two tubercles
in front of, and the one in line with, the back of each eye, appear
to be larger than the others of these sets: besides these rows
there are a few tubercles bordering the inner edge of the
marginal furrow, and the thickened border bears a chain of
these small bead-like tubercles of uniform size, and about twenty-
five in number, the rest of the surface being finely granulated.
The eyes are small, crescentic, close to the glabella and posterior
margin; neck-furrow shallow, as are also its lateral extensions:
neck-ring fairly robust and granulated, lateral extensions rela-
tively strong and narrow; border furrow wide and deep: border
strongly tumid and tuberculate; angles, if not spinate, are acute.
06s. — The above Brachymetopus is the first and only specimen
of the genus collected from the Carboniferous rocks of Queens-
land. It occurs at Trilobite Ridge, Mt. Morgan, Queensland,
associated with other genera of trilobites. In several features
it resembles lir. strzeleckii Mc(Joy, the prototype, and agrees with
this species and Br. Muccoyi in possessing a well-defined cephalic
marginal border, bounded inwardly by a well-defined furrow.
BY JOHN MITCHELL. 48?
The tuberculation, too, on this border is similar in the three
species. Dimensionally, this species agrees closely with Br.
strzeleckii, judging from the dimensions given by Reed,"^ so also
does it in part in the character of the ornamentation, but in
part in this respect, it differs very widely, as the description
shows.
The chief differences between our species and Br. strzeleckii
are — 1. The character of the ornamentation. 2. The more conical
shape of the glabella in ours, and the absence of the large longi-
tudinally placed tubercles on this part. 3. The more acutely
rounded cephalon, particularly at the front. 4. The eyes are
situated closer to the posterior border, and perhaps to the
glabella. 5. The lateral extensions of the neck-ring are narrow
and prominent. Without doubt, the two forms are closely
related.
Since writing the above, I have found, on some of the Queens-
land specimens from Rockhampton and Mt. Morgan districts,
several pygidia and a portion (cheek) of a cephalic shield, which
belong to the genus Brachymetopus. One of these pygidia,
which is fairly well preserved, I am assuming to be specifically
identical with the cephalon above described. Its description is
as follows.
Pygidium semicircular, only mildly convex, strongly tuber-
culate. Axis moderately prominent, contracting gradually pos-
teriorly, ending short of the border bluntly, with less than half
its anterior width: it bears five longitudinal rows of tubercles,
the individuals of the middle row being much larger than those
in the rows on each side of it. There are at least twelve or
thirteen rings in the axis, and a central tubercle on ten of these
can be clearly seen; this applies also in the case of each row
immediately adjacent to the central one. Axial furrows deep.
Side-lobes consist of six (? seven) pairs of segments, the ridges of
which are very prominent, and bear several relatively large
tubercles and some of smaller size: each of the posterior pair
merely consists of a tubercle, and each ridge of the anterior pairs
bears several tubercles varying much in size, the larger being
'■' Geol. .^Ja.^, X.S., Dec. iv., Vol. x.. IIMKS. pp. llKMUfJ.
488 CARBONIFEKOUS TRILOBITES OF AUSTRALIA,
situated near the axial furrowy and border; the ribs apparently
extend into spines beyond the border, mesial furrows of the ribs
deep. Border of moderate width, and apparently only mildly
thickened.
This pygidium, though it presents in a general way a
strong resemblance to that of Br. strzeleckii, exhibits features
which help to separate the two forms. The chief differences
between their pygidia are — l. The axis of Br. dunstani consists
of twelve or thirteen rings, as against ten in the other. 2. Five
longitudinal rows of tubercles traverse the axis of the former,
while seven rows occur on the latter on the same part. 3. A
large tubercle is situated centrally on each axial ring of the
former, but only on every alternate axial ring in the latter.
4. There is a smaller number of tubercles on each pleural seg
ment of the former than on the similar segments of the latter.
The pygidial ornamentation of our species agrees closely with
Br. lodiensis Meek.
Dedicated to Mr. B. Dunstan, Chief Government Geologist,
Brisbane, whose palyeontological discoveries have been numerous
and important.
Loc. and Hor. — Trilobite Ridge, Mt. Morgan, Queensland,
associated with P. ivoodivardi. Lower or Middle Carboniferous.
Specimens Nos.F1031, F 1007, Geological Museum, Dept. of
Mines, Brisbane; and No.712, Queensland Museum, Brisbane.
Brachymetopus, sp. indet.
\\\ addition to the head and tail, which served for the descrip-
tion of Br. daiisfaui, there occur on specimen No. 712 of the
Queensland Museum, Brisbane, no less than three pygidial frag-
ments of Brachymetopi, besides the pygidium which served for
the foregoing description of Br. dunstani. In the case of two,
one is partially superimposed on the other, and the tliird is
almost touching these. On specimen F 1007 is a portion of a
right cheek, which presents features evidently unlike either Br.
strzeleckii or Br. dunstani. Two of the fragmentary pygidia
also appear to be new, and maybe specifically identical with the
individual to which the cheek-fragment belonged. The material.
BY JOHN MITCHELL, 489
liowever, is too fragmentary to describe and figure. The occur-
rence of three pygidia in a launch indicates that they may be
plentiful, and careful search may bring better specimens to light.
KXI^LANATION OF PLATES XLVI.-LIII.
Plate xlvi.
PliWiima colfinsi Mitchell.
I^'ig.s. 1, 2, o, 4. .">. — Fin. 1 i« a photo ( x S) made froni the specimen repre-
sented in Fig.'i. The glabellar features are plainly visible. Figs.
.'i, 4, T) represent pygidia. and shoM" their characteristic featui-es
very well. Figs.4 and "> x | (about). (Coll. Mitchell).
PhUlipsia coulteri Mitchell.
Figs. (). 7, S. *•. 10. — Different views of one individual. Fig.H x 4, and Fig.
10 nearly natural size. In Figs. 8 and 9, the mild dorsal serration is
visil)le. In Fig. 10. the hypostome is plaiidj^ seen, and the latlier
large tubercle of the terminal axial ring. (Coll. Mitchell).
PJi///ip-^ia hftricejjs Mitchell,
Figs.ll. 12.— Cephalon and tail (Colh Mitchell).
Grijfit hides conrexicaiidatus Mitchell.
Fig. 18.— Dorsal view. (Coll. Mitchell).
PJii//ip)<ia xfroHdeii.-!(s Mitchell.
Fig. 14. — The specific features clearly represented; ( x :]). (Coll. Mitchell).
Phillipxiu proxlnm ^litchell.
Figs. IT), 16. — ^Two views of a pygidium, the lattci' alxjut naUnal size.
(Coll. Mitchell).
Plate xlvii.
PhiUipuKt rohnsta Mitcliell.
l''ig. 1. — The pygidium originally described l)y Mi'. W. Flhcridge. Junr..as
/'. <irandi-<. II shows the important features rather clearly-. (Coll.
< Geological and Mining Museum, Dept. of Mines, Sydney. No. 14'.»2).
Pl,llli)>s;n (iruiidi.s Ftli. Hi.
l''ig.2. — Piioto of the specimen originally descrihetl hy Mr.' H. Flhcridge.
Junr. , and for which the specific nam3 r/y-rt/Af//.*.- was first suggested
by him. The figure shows how different in character arc the pleural
ribs from those of P. rohiisfa; the sutures along the aitit-ulatijig
ridges, and the crossing of the border by the anterinr ril»s. (Coll.
Geological and Mining Museum, Dept. of Mines, Sydney. Xo.l.lKO).
PhiUij,xi<i cloiKjald Mitchell.
Figs. 8, 4. — ^Casts from an almost perfect mould (jr cover. They exhil)it
clearly the features of this fine trilobite. [Coll. (reological and
Mining Museum, Dept. of Mines, Sydney. No. I.IOO, (cover)].
490 • CARBONIFKHOUS TRILOBITES OF AUSTRALIA,
Fig. 5. — The natural cast. Part of the right side of thecephalon is absent.
Shows traces of the left genal spine. (Coll. Geological and Mining
Museum, Dept. of Mines, Sj^dnej-. Fig. 3, FloOO, etc.).
Phillipsia dimgof/ensis Mitchell.
Fig. 6. — Part of a pygidium, showing the continuous border, and other
essential features. (Coll. Geological and Mining Museum, Dept. of
Mines, Sydney. Xo. 1494).
Fig. 7. — A smaller and nearly perfect pygidium; (x^). (Coll. Geological
and Mining Museum, Dept. of Mines, Sydney. No. 1494).
Phi/Npsia rohusta Mitchell.
Fig. 8. — Another view of the pygidium, only less enlarged.
Phillipsia ivoodwai^di Eth.fil.
Fig. 9. — Medial portion of a cephalon much weathered. (Coll. Geological
Museum, Dept. of Mines, Brisbane. No.F. 1017).
Plate xlviii.
All figures on this Plate x % about.
Griffith ides conrexicaiidatus Mitchell.
Figs.l, 2. — Dorsal and side-view.
Fig, 3. — Pygidium and free cheek, with the ej'e of a young individual: the
free cheek partly covering the tail. (Coll. Mitchell).
Phillipsia rouchelensis Mitchell.
Figs. 4, ."), (>, 7. — Four p3-gidia. Thej' exhibit the specific features. Figs.
o and 7 are photos of the specimens used bj^ Mr. Etheridge for his
figures (PL xxi., figs. 6-7, opj. cit, antea). (Coll. Geological and
Mining Museum, Dept. of Mines, Sj'dney. Nos. E 1495, 3534 (2),
F 1595).
I^hiUipsia colli n-'<i Mitchell.
Figs. 8, 9. — Fragmentary head-shield, and faint outline of a poorly pre-
served l)ut almost complete individual. In Fig. 8, all the glabellar
features are visible. (Coll. Mitchell).
Phillipsia staiirellensis Mitchell.
Figs. 10. 1 1, 12. — Three different views of a mature individual. The speci-
men represented by these figures was used bj^ Mr. Etheridge for his
Queensland type of P. dubia (PL viii., fig.5, op. cit.). (Coll. (geolo-
gical Museum, Dept. of Mines, Brisbane. No.F9«i9).
Fig. 13. — On this photo are the middle pait of a cephalon and a pygidium
of an immature individual. The former shows the normal and
dunce's hat-like shape of the glabella, globular basal glabellar lobes,
and the relatively strong neck -ring. The pygidium shows the iiar-
row, prominent axis, etc., characteristic of the species. (Coll. Geo-
logical Museum, Dept. of Mines, Brisbane. No.F 980).
BY JOHN MITCHELL.
491
Phillip-na eJom/afa Mitchell.
Fig. 14. — Portion of a head-shield, thorax, and tail. The markings on the
basal glabellar lobes are accidental. The normal glabellar frontal
limli and anterior courses of the facial sutures are shown. (Coll.
Geological and Mining Museum, Dept. of Mines, Sydney. No.ri498;
Pal. Mem. Pt..!. No.2, PI. xxi., fig.2).
PJullipsia superlKi Mitchell.
Fig. lo.^Cephalon minus the right free cheek. (Coll. (ieologieal and
Mining Museum, Dept. of Mines, Sydnej-. No.'24).
Phillipsia iraterhoHsei Mitchell.
Figs. 16, 17, 18. — The first and second of these are from an intaglio, and
the other is from a squeeze from it. With the aid of a lens, the
features can be made out. (Coll. Geological and Mining Museum,
Dept. of Mines. Sydney. Nos. 28 and 30).
Plate xlix.
PhiUipsia iroodnrirdi Eth.fil.
Figs l-fi. — Pygidia, all assumed to belong to this species. All show prac-
ticallj' the same features. Figs. 3 and 4 are more strongly gran-
ulated than the others; but this difference probablj^ arises from
degrees of weathering. Fig. 5 was figured by Mr. Etheridge {op. cit.
PI. viii., fig. 6) as the normal tail of his P. dubia. (Coll. (geological
Museum, Dept. of Mines, Brisbane. Nos. F 985, F 1024, F 995,
F 1026, F 968, F 993).
Figs. 7-8. — Photos of a fragmentary cephalon, about which I am doubtful
that it is rightly placed here. It is one of Mr. Etheridge's types
(Geol. and Pal. Queensland and New Guinea, PI. vii., fig. 13). (Coll.
(Geological Museum, Dept. of Mines, Brisbane. No. 967).
Phi//ipsia roekhamp)toHensi-s Mitchell.
Figs.9-10. — Two prints of a nearly complete specimen. Fig. 10 has some
of its features intensified. This specimen was figured by Mr.
Etheridge, Junr., and determined to belong to his P. duhia {op. cit.,
PL xliv,, fig, 4). This figure very indifterently represents the
original, as will be seen by comparing it with the present photos,
which show its chief characteristic features clearly. (Coll. Queens-
land Museum, No. 716).
Phillipsia morgane.nsis Mitchell.
Figs. 11-12. — Fig. 11 represents portions of a cephalon, thorax, pygidium,
and a hypostome. Fig. 12 is a rough sketch of part of a cephalon.
(Coll. Geological Museum, Dept. of Mines, Brisbane, No.F 1000).
Phillipsia woodwardi Eth.fil.
Figs. 13-14. — Photos of the type-specimen. Fig. 14 has the outline traced
in, (Coll, Geological Museum, Dept. of Mines, Brisbane. No,966).
492 CARBONIFEROUS TRILOBITES OF AUSTRALIA,
Brachymetapv.^ dnvsfani Mitchell.
Ficrs. 15-16. — Showing the cephalon and pyoidium of the species. (Coll.
(geological Museum, Dept. of Mines, Brisbane, No, F 1031; and
Queensland Museum. Brisbane, No. 712).
Plate 1.
All figures, except 2 and H, whieli are about nat. size, x #.
Phillipsia (iramlis Eth. fil.
Figs. 1. "2, 3. — Photos of a pygidium. Figs. 2 and 3 are from casts of No. 1
in Fig. 1. Besides the pygidium, parts f)f at least three free cheeks,
etc., are visible. One cheek shows within it tlie hypo-stome. The
borders of these cheeks are remarkablj- Luge. (Coll. (geological
Museum, Dept. of Mines, Brisbane. No. F 027).
Phinipsia eloiKjata Mitchell.
Figs. 4, 5, 6, 7. — Figs. 4 and 7 represent pygidia. Fig. .3 is a side-view of a
very fine specimen, and shows, besides other features, the squatness
of the eyes. Fig. (>, a pygidium and a portion of a head-shield.
(Coll. (Geological and ■Mining Museum. Dept. of Mines, Sydney.
Nos. F 14!>6, 1.-)(I0. and l.)0()).
J^hi/h'p.-n'a rockhaniptoiitnsis M itchell.
Figs.S, 9. — Two pygidia. Fig. 8 is a photo of the specimen sliown in (U'ol.
and Pal. (Queensland and New (Guinea, Pl.xliv., fig.G. Fig.9 repre-
sents a young individual. (Coll. (Queensland Museum, Brisbane,
No.833: and (Jeological Museum. Dept. of Mines, l>risbaiie, No.
F7i>2).
PhiUipsiai'i) woodwardi Eth. fil.
Fig. 10,~(3n this specimen are several pygidia and a remarkable iiypostome,
doubtfully assumed to belong to this species; and, in addition, the
greater part of a cheek of a Hrachymetopiis; the position of which
is indicated by an arrow. (Coll. (Geological Museum, Dept. of
Mines, Brisbane. No. 1007).
Fig. 11. — Glabella; one of Mr. Etheridge's types. (Coll. (Queensland
Museum, Brisbane. No. 707).
Plate li.
If not otherwise indicated, all the figures are x 'i (about).
Phillipsia mori/anensis Mitchell.
Fig. 1. — A photo, in two sections, of a specimen on which are parts of at
least three individuals, all assumed to belong to this species. The
glabellar, pygidial, ocular, and hypostomal features are all dis-
cernible, as are also those of the thorax. (Coll. Geological Museum,
Dept. of Mines, Brisbane. No.F 1000),
BY JOHN MITCHELL
49:;
rhilli)>sia brevictp-^ Mitchell.
Fi<;K.2, 3. — Photo of a specimen showinj; portions .of two cephalic-shielda
and a very perfect pyicidiiini, whose characteristics are represented.
Mark the peculiar contraction near the middle of the tail, which
suggests a short tail, and a portion of a thorax. Fig.3, a portion
of a cephalon. shoN\ inu- limb, mesial furrow on the right, etc. (Coll.
Mitchell).
riifUipsia connollii Mitchell.
Figs. 4-7- — ^A doisal and side-view of the only pygidium known. The pro-
portionately very wide border, pi^ominent axis, granulation, and
other features are faii-lj- well shown. In Fig. 6, the axial and pleural
divisions have been intensified slightl3'. (Coll. (Geological and
Mining Museum, Dept. of Mines, Sydney. No. F 1497).
Philllpsia sfanvel/ensis Mitchell.
Figs.S-lO. — Dorsal and side-view of a very perfect tail, and showing the
very prominent, mildly serrated axis clearlj'. In this specimen, all
the axial rings (12. doubtfully 13) are visible, as also are eight
pleural segments, and the steep striated border. (Coll. (Geological
Museum, Dept. of Mines, Brisbane. No.F977).
PIn/lipsia xfroiidennx Mitchell.
Fig. 11. — A medium-sized tail, having the dorsal part of the axis damaged;
but other\\ise exhibiting the normal fetitures. (Coll. Mitchell).
Phillipsia icoodicardi Eth, fil.
Figs. 12-13.— Portions of cephalons. Fig. 12 shows the glabellar furrows,
strong neck-ring, and supplementary lobes, etc., very well. Fig, 13
exhibits these features less clearly. Fig. 12 is from the same speci-
men as that figured by Mr. Etheridge, Junr., {op. cit., PI. xliv.,
fig.5). (Coll. Queensland Museum, Brisbane, No.707; and Geological
Museum, Dept. of Mines, Brisbane, No.F 1017).
PhiUipsia vmodu:ardi{''.) Eth. fil.
Fig. 14. — This photo shows the fragments of trilobite-remains, on which
Mr. Etheridge chietly founded his species G. >*emmiferus var. au8-
tralasica. On this specimen also occurs the tail of Brarhymetopus
dunstani. It remains to be proven that they are not portions of the
above species. The tails shown on this specimen have the same
number of axial and pleural divisions, and kind of granulation as
those included with the cejihalons of P. ivoodwardi. (Coll. Queens-
land Museum. Brisbane. No. 712).
Plate lii.
Phillipsia inoodwardi{'t) Eth. fil., and Brachymetopus dwnstani M'ltaheW.
Fig.l. — This represents a portion of specimen No.712 of the Queensland
Museum, Brisbane. On it is the pygidium assumed to belong to
P. iL'OodAi-a rdi , and a pygidium of Jir. davMani in front of the arrow;
( X 3).
494 CARBONIFEKOUS TRir.OBITES OF AUSTRALIA.
Brai-liyine.topus diinstani Mitcliell. etc.
Fig.2. — Enlarged poitjon of specimen No. F 1017, (reological Museum,
Dept. of Mines, Brisbane. Oh it is the intaglio of Br. dnnstani,
showing, fairly well, the chief specific features. The other pygidial
and cephalic imprints are assumed to belong to P. iwodirardi Eth.
fil. ; ( X 3).
Phi/npsia HHperha Mitchell.
Fig.3. — Head-shield ( x4) from the same specimen as PI. xlviii., tig. 15.
PliiUipskt coUinsi Mitchell.
Fig. 4. — A perfect tail of an immature individual; ( x f ),
Grijfithides convexicaudatust ^Mitchell.
Figs. 5, 6. — Tails about complete. Fig.5 is that of a young individual. Init
shows all the axial and pleural divisions.
PliUUpsia waterhoiisei Mitchell.
Fig. 7. — From a cast; it shows tlie glabellar furrows plainly: and other
features.
Plate liii.
Griffi.thidef>{1) sweet I Eth. fil.
Figs. 1,2. — Dorsal aspect, rraces of glabellar furi-ows are visible, especi-
ally on the right side generalh'. The important features are well
shown. The photos are from the type-specimen; (x2). (Coll.
Sweet).
Brachymetopus strzeJeckil McCoj'.
Figs. 3, 4. — Photos of the enlarged figures of McCoy (lor. cit.).
Figs.5, 6.— Photos of Reed's figures of the species (/or rif.).
Grijfif hides dnhins Eth. Senr.
Fig. 7. — Copy of the figure given by R. Etheridge, Junr.. (in (^eol. Pal,
Queensland and New Guinea, 1892, PI. vii., fig. 12).
Phif/ipsia iroodwardi Eth. fil.
Figs.8, 9.— Copies of Mr. Etheridge's figufes (op. cit., PI. vii.. figs. 13, 15).
Griffithidesi'i) seminiferus var. anstralasica.
Fig. 10. — Copy of Mr. Etheridge's original figure of a free cheek.
495
ORDINARY :M0NTHLY MEETING.
September 25tb, 1918.
Professor H. G. Gliapiiicui. ^[.D., B.S., President, in tlie Cliair.
The Piesident announced tliat tlie Council had (;ome to the
conclusion, that tlie proposed scheme "to create a miniature
Australia in its primary conditions," on Pulhah Island in Lake
Macquarie, as mentioned at hist Meeting, was not a matter in
wliich the Society could move with advantage, at present.
The Donations and Exchanges received since the previous
Monthly Meeting (28th August, 1918), amounting to 4 Yols., 38
Parts or Nos., 5 Bulletins, 1 Report, and 2 Pamphlets, received
from 30 Societies, etc., were laid upon the table
NOTES AND EXHIBITS.
Dr. J. B Oleland mentioned that, on two consecutive days in
August last, he had heard and seen a Little Penguin, Eiidpytula
minor, off Kurraba Point, in Neutral Ba^^
Dr. R. J. Tillyard exhibited a slide of some transverse sections
of the Saccoid Caudal Gills of the larva of the Dragoni^y Fseudo-
phcea sp., (Fam. CalopterygidcE) from Java, sent by Dr. F. Ris, of
Rheinau, Switzerland. In this preparation, the alveoli show
some remarkable structures, described originally by Dr. Ris as
'"moss-like branching tufts of standing fibrils," and considered
by him to be special respiratory organs of the gill. From the
photographs which Dr. Ris published of these sections, Dr.
Tillyard concluded that thny were artefacts, a conclusion which
Dr. Ris is unwilling to accept. He therefore sent the slide in
support of his opinion. The slide shows many alveoli in which
the fibril-tufts project considerably beyond the boundary of a
single alveolus. They were examined under the microscope by
496 NOTES AND EXHIBITS.
biologists and ph3'siologists present at the Meeting, ail of whoin
agreed tliat they were artificial products: thougli it is not pos-
sible to say definitely what caused them, in view of the fact that
no information is available as to the method of fixation used by
the collector who took the larva in Java. (These Proceedings,
1917, xlii., pp.51-54, and Pl.iv., figs.31-33).
Mr. Fletcher exhibited five portions of typical, water-storing
Mallee-roots, and one of two bottles of root-water from South
Australia, received from Mr. H. S. Crummer, through the kind
co-operation of Mr. T. Gill, of Adelaide, and Mr. G. Murray, of
Fowler's Bay, S.A.
Dr. R. Greig-Smith oflt'ered some observations, based on a pre-
liminary examination of one of the two samples of water obtained
from Mallee-roots.
497
NEW AND RARE FRESHWATER ALG^.
By G. I. Playfair, late Science Research Scholar op the
University op Sydney.
(Plates liv.-lviii.; and eleven Text-figures.)
In the following notes are embodied observations on a number
of new and rare freshwater alg?e, that have come under my notice
during the last few years. Of those mentioned, a little over two-
thirds of the new forms, and about half the others, are from
Lismore, the gleanings of 23 gatherings made during the summers
of 1916 and 1917: to be precise, from Sept. 16th, 1916, to May
Uth, 1917, and from Sept. 21st, 1917, to Feb. 10th, 1918. During
the colder months of May, June, July, and August, very little of
interest is to be found. The others are from older samples, also
locally obtained, or out of still earlier gatherings from Sydney.
In all, 66 new forms are described and figured, 28 being ad-
mitted to specific rank, 29 classed as variations, and 9 as forms;
una new genus is proposed. The figures are, of course, all drawn
from Australian specimens.
Habitats. — The districts from which the various forms have
been obtained are purely of local interest, but more general value
attaches to the character of the habitat. The numbered gather-
ings, referred to, are accordingly classified as follows. Plankton :
80, 81, 90, 99, 100, 115, 121, 136, 258, 265, 266, 311, 316, 327,
328, 332, 340«, 347, 362. Out of weeds (practically plankton):
3, 26, 50, 158, 184, 189, 193, 272, 317. Mucous strata: 306,
323, 326. Ponds, lagoons, and swamps account for the remain-
ing twenty.
One of these last, a swampy pool formed entirely by surface-
water, deserves special mention; during the two smnmers, it was
responsible for 30 out of the 64 new forms here described. The
samples which refer to it are 284, 286, 298, 302, 308, 311, 312,
38
498 NEW AND RARE FRESHWATER ALGiE,
316, 317, 327, 337, 340a, 345, 347, 352. The pool is the drain,
age of a considerable area of ground swampy after rain, and is
about 2 feet deep in the centre; in droughty weather, it is often
entirely dry."*^
On account of its unpromising appearance, and to secure ever}^-
thing that might be present, a quantity of water was taken from
a spot where there was a growth of coarse weeds, the weeds being
well shaken up, and the bottom disturbed also to some extent.
This water was strained through a plankton-net of common calico,
ground-samples from the swamp being sometimes added. Several
gallons would be treated in this way, and the last quart, or so,
of water and sediment taken home in a tin and decanted.
During the periods mentioned above, a gathering was taken,
on an average, once a month, the main portion of the water being
drawn from the same spot. The results show the importance of
a regular periodic examination of a pond. The desmids and
diatoms were fairly constant, but the Myxo2:>hyceai, Volvocacece,
and Protococcoidece presented, from month to month, an ap-
parently endless succession of forms, new, uncommon, or not
previously recorded. Rarely could any of these be noted in two
successive gatherings, and very often they were never seen again.
For instance, of Chlamydomonas, 10 different forms, new or rare,
were noted; of Carteria, 5; of Phacotus, 2; of Volvox, 2; of
Oocystis, 7; of Tetraedy^on, 7; of Geminella, 4. Yet, in two
summers, the most that could be said was, that a few of them
had been recorded a second time.
Of course the swamp, with its ever-changing conditions of life,
is the real breeding-ground of all these forms, the pool being
only the repository of them, Chlamydomonas and Gay-teria are
notoriously dependent on rainfall, and are very sensitive to stag-
nation; but I find that forms of Oocystis and Tetraedron are
just as irregular in their appearance. Probably, the reason is
* Swampy pools of this sort are the best places to search for new and
rare forms of microscopic life; permanent waters, on the other hand, can
generally be relied on to jaeld a micro-flora and fauna of unvarying
character.
BY G. I. PLAYFAIR. 499
to be found in the propagation of all tliese genera by autospores.
To l)egin with, the autospores differ sHghtly (and sometimes more
tlian slightly) from the parent-cell, and their subsequent develop-
ment along diverging lines will, no doubt, result in more oi' less
widely differing forms.
Another interesting point to be noted is, that now and again
one or other of these forms would be recorded at the scmip time
from some other pool connected with an entirely different water-
shed, or even in another part of the neighbourhood, but thev
were just as evanescent.
MYXOPHYCE^.
Syn. Phycochromojjhycefe^ Cyanophycece, Schizoi^hycpw.
Fam . CHROOCOCCACE.^J .
Genus Syxechococcus ^ag.
Syxechococcus grandis, n.sp. (Text-fig. 1 ).
Cellulse magnse, crasspe, oblongo-o vales; apicibus
late-rotundatis ; lateribus arcuatis ; cytoplasmate l^'^'^'.
jeruginoso, gi-anuloso. /l;'\^«-'*-i"4;^l
Long. 33, lat. L>3//. Lismore (345).
Of Nageli's three species (Gatt. einz. Alg., p. 5 6,
T. i.E, f.1-3), none are over 20/x long. S. major
Schroter, the nearest in size, is cylindrical. See G. Text-fig. 1."
8. West, Br. Frw. Alg., p. 347, f.l61D, E.
Genus M e r i s M o p e d i u m Meyen.
Merismopedium punctatum Meyen. (Text-fig. 2^).
In Wiegm., Archiv, 1839, p. 67, sec. Rabenhorst, Fl. Eur. Alg.^
ii., p. 57. »Syn., J/. KiUzingii Nag., I.e., p.55, T. i.D, f.2. Con-
tents generally pale blue. Usually found in our waters in isolated
sets of four cells. As figured, however, out of weeds in the river,
the cells were in large sheets but still semi-detached in coenobia
of 4.
Cell. diam. 2/x. Lismore, Richmond R. (272).
* Synechococciis grandis, n.sp., ( x 660),
500
NKW AND RARE FRKSHWATP^R ALGiE,
Var. OBLOXGUM Playf. (Text-fig. 25, c).
Biol. Richm. River, p. 135. Generally sparsely distributed in
isolated sets of 4, or more rarely of 16. The latter were frequent
in plankton from the Nymboidia River, near Grafton, where I
0181 18 1®
01 m iiii
00 80 0i8i
GO
nng
Text-fig. 2.
(a) Merismopedium punctatum Meyen, forma, ( x500); (h, c) var. oblongum
•Playf., [h X 1000, c x6fi0); (cZ, e) var. vacuolatum, n.var., ( x660);
(/) M. cyaneitm, n.sp., ( x 660); ig) M. elegans var. constrictum, n.var.,
( x 660); (A) M. duplex, n.sp., ( x 1330).
even noted a large coenobium (30 x 20/x) of four sets of 16 cells
each. The cells are irregularly oblong, the contents pale blue.
Cell. long. 2-3, lat. l-|-2/x (plerumque 3 x 2/x).
Lismore(350); Grafton (265, 266).
Var. VACUOLATUM, n.var. (Text-fig. 2c/, e).
Cellulae globoste vel oblongie; media cellula loco vacuo magno
occupata.
Cell, diam. 2/x; vel long. 3, lat.
Lismore (350).
The cells are irregularly globose or oblong, and the centre is
occupied by a large vacuole giving each cell the appearance of a
ring
coenobium
Globose and oblong cells are sometimes mixed in the same
UY G. J. PLAYFAlU. 501
Merismopedium cyaneum, 11. sp. (Text-fig. 2/).
Celluh« o vales vel oblong^e; cytoplasmate cyaneo.
Cell. long. 5-6, lat 3/x. Potts Hill (121).
Ccenobium of 4 cells measured 12 x 6/i, the cells not quite
divided; the colour of the cell-contents a strong bright blue.
Merismopedium elegans v. constrictum, n.var. (Text.fig.2r/).
Ccenobium membranaceum, extensum, e cellulis permultis, con-
fertis, compositum; cellulis oblongis medio constrictis; cytoj^las-
mate dilute Cc^^ruleo vel dilute viridi.
Ctenob. long. c. 300, lat. c. 200; cell. long. 6-7, lat. 4-5/x.
Lismore, Richmond R. (184, 193).
The crenobia were wide-spreading, containing 1024 cells (32 x
32) or so. Cells oblong, constricted in the middle, pale blue or
pale green. A few cells of the type present, oblong, not con-
stricted, 6 X 4//.
Merismopedium duplex, n.sp. (Text-fig. 2A).
Cellulfe sph£ericc\3 vel ovales, remotte ; cytoplasmate dilute
creruleo.
Cell. long. 4, lat. 3/i Coogee (4).
Coenobia small, about 30 x 25f(., consisting of 8-32 cells gener-
ally found half -divided, remote. Contents pale blue.
Fani. 08CILLAT0RIACE.*:.
Genus Oscillatoria Vaucher.
OSCILLATORIA FLEXA, n.Sp. (PI. Hv., fig.l).
►Stratum mucosum, foliaceum, saturate viride; fills arete in-
tricatis rectis, juxta apices subito deflexis, baud attenuatis,
apicibus rotundatis, vaginis tenuissimis; trichomatis dilutissime
ferugineis, ad genicula baud constrictis; articulis subquadratis
vel cylindraceis; dissepimentis fegre cernendis, haud granulatis;
cytoplasmate tenui-granuloso.
Diam. fil. 21-3, cell. alt. 2-8/x. Lismore (317).
A mucous stratum of dark green flakes in shallow water at
the grassy edge of a swampy pool, in quantity. The filaments
were very lively under the microscope, as is usual with species of
502 \EW AND RARE FRESHWATER ALGiE,
this genus; and the sheath, though very delicate, was quite dis-
tinct in broken filaments. I have always found it so, even in
the finest species of Oscillatoria. The filaments, in general quite
straight, were bent suddenly about 10// from the end, which was
not attenuated.
Oscillatoria simplicissima v. granulata, n.var. (PL liv., f.2).
Forma dissepimentis tenuissime granulatis.
Diam. fil. 8, cell. alt. 4-6 (plerumque 4/i,).
Lismore (327).
Filaments free-floating in prodigious quantities, colour pale
grey-green, contents finely granular, as also are the dissepiments.
Var. CONSTRICTA, n.var. (PI. liv., f.3).
Forma trichomatis ad genicula levissime constrictis.
Diam. fil. 13, cell. alt. 4-8/x. Lismore (350).
Filaments free-floating, more than half as bi'oad again as in
the type, with the cells slightly constricted, a mere nick at the
edge. Colour pale blue, contents homogeneous not granular.
Type in Gomont, Monog. d. Oscillariees, p. 2 19, PL vii., f.l.
Oscillatoria princeps Vauch. (PL liv., f.4).
Diam. fil. 30-57, cell. alt. 4-8/x. Lismore (350, 352).
Colour pale grey-blue or grey-green. I mention this species
for the sake of giving a figure of a piece of empty sheath show-
ing fixed dissepiments right across, with incipient intermediate
ones. In surf ace- view, the septa appeared as faint transverse
lines, but, in optical section, they showed bulging to one end, as
if the cells had been violently forced out.
Genus P h o r m i d i u m Kiitz.
Phormidium GRANDE, n.sp. (PL liv., f.5).
Stratum papyraceum olivaceum; fills rectis, apicibus rotundatis,
calyptra nulla, vaginis plerumque crassis; trichomatis saturate
viridibus interdum luteolo-viridibus, ad genicula baud constrictis;
articulis brevissimis; dissepimentis latis, baud granulatis; cyto-
plasmate tenui-granuloso.
6Y G. I. PLAYFAiR. 503
Diam. fil. 21-23, trich. 19-20, cell. alt. li-2i (plerunique 2/x).
Lismore (32.3).
An olive-green papyraceous stratum in great cjuantities on the
surface of a creek-pool, after heavy rain followed by heat. The
trichomes under the microscope were of a brilliant green colour
(very unusual in this family, but found also in 0. nigro-viridis)
or sometimes yellow-green. The tips of the filaments were some-
times rounded, but generally slightly constricted at the terminal
cell so as to be subcapitate.
Phohmidium numarium, n.sp. (PL liv., f.6, 7).
Stratum papyraceum olivaceum; filis angustissimis ai'cte in-
tricatis, vaginis tenuissimis, apicibus rotundatis ; trichomatis
dilute Ccoruleis, ad genicula valde constrictis, articulis brevissimis
disjunctis; cytoplasmate homogeneo.
Diam. fil. 2, cell. alt. J-lJ/>t. Lismore (326).
An olive-green papyraceous stratum on the curb-stone near a
down-pipe. The cells of the trichomes are disjoined and are of
the palisade-type, reminding one of a rouleau of coin, at most
quadrate before division, only to be distinctly made out with
the yV obj., the contents homogeneous, pale blue with a darker
band at the sides.
Fam. NOSTOCACE/E.
Genus A n a b ^ n a Bory.
ANABiENA AUSTRALICA, n.Sp. (PI. liv., f.8, 9).
Trichouiata angusta, recta, libere natantia, ad genicula
levissime constricta; cellulis rectangularibus, adpressis, apicibus
truncatis; cytoplasmate dilutissime cseruleo jjiene hyalino, homo-
geneo vel rainutissime granulato; heterocystidibus angustis cylin-
draceis, apicibus rotundato-truncatis ; gonidiis angustis cylin-
draceis, apicibus rotundatis, lateribus parallelis, juxta hetero-
cystides ordinatis; cytoplasmate pulchre cyaneo, granulato.
Cell. diam. 3-1, alt. 6-10, plerumque 8; heterocyst. diam, 4-5,
alt. 10^-15; gonid. diam. 5-7, alt. 19-32/x.
Lismore (327, 337).
Found free, floating. Trichomes narrow, very pale blue,
504 NEW AND UARE FKESIIWATKR ALG.'E,
almost liyaliiie, witli cells so strictly rectangular and so closely
adpressed that the trichoine has the appearance of an Oscillatoria\
the slight constriction at the joints, a mere nick at the edge, is
discernible only with the yV obj. (often the dissepiments also):
contents homogeneous or finely granular. Heterocysts narrow ,
strictly cylindrical. Gonidia one or two on either side of a
heterocyst, narrow, cylindrical; contents bright blue, granular.
Var. CONSTRICTA, n.var. (PL liv., f.lO).
Forma gonidiis in medio constrictis. Cell. diam. o, alt. (S-lO;
heteroc^^st. diam. 4, alt. 11; gonid. diam. 4-6, alt. 19-21/x.
Lismore (350).
A form with rather short gonidia somewhat constricted in the-
middle.
Anab^na variabilis var. cylindracea, n.var. (PI. liv., f. 11-1 3).
Trichomata Hexuosa, dilute cterulea, ad genicula constricta;
cellulis stricte rectangularibus, disjunctis, apicibus truncatis;
cytoplasmate minute granulato ; heterocystidibus oblongis vel
oblongo-cylindraceis, apicibus rotundatis; gonidiis doliformibus,
quadratis vel oblongis, disjunctis, apicibus truncatis.
Cell. diam. 3, alt. 4-6; heterocyst. diam. 4-5, alt. 7^-10^; gonid.
diam. 6, alt. 7-lOJ/x. Lismore (316).
Easily recognised as A. variahilis by its catenate gonidia remote
from the heterocysts, differing from the type, however, in its
rectangular cells. All parts of the trichome also seem to be
slightly more slender than usual. The gonidia were perhaps
immature, and would probably become oblong with rounded ends.
Anab^na oscillarioides var. crassa, n.var. (PL liv., f.l4).
Forma cellulis crassioribus, globosis (e divisione oblongis vel
oblongo-constrictis) ; cytoplasmate teruginoso ; heterocystidibus
interdum minoribus, sphgericis; gonidiis oblongis, apicibus rotund-
atis, cytoplasmate seruginoso granulato.
(1). Cell. diam. 6-8; heterocyst. 4; gonid. diam. 9, alt. 22-23/x.
Lismore (298).
(2). Cell. diam. 8-9|; heterocyst. 8; gonid. diam. 12, alt. 25/x.
Lismore (337).
1?Y G. i. PLAYFAIR. 505
111 this form, the cells are broader aud the heterocysts some-
times narrower than in the type (cell. diam. 4-6, heterocyst. 6-8,
gonid. 8-10/x); c/'. Hornet et Flahault, Kevis. d. Nostoc., p.'233;
Tilden, Minnesota Algtp, i., p. 193, PI. ix., f.l9.
Anab.ena laxa \ar. hortkxsis, n.var. (PI. liv., f. 15, 16).
Forma heterocystidibus cylindiaeeis : g()ni(his cvlindraceis
elongatis.
Cell. diam. 6-7, alt. 8-12; heterocyst. 7, alt. 14-16; gonid. 7,
alt. 33//.
Botanic Gardens, Sydney (158).
Cf. iSpha'rozyc/a {Anahfcua) Jaxa Habenhoi'st, Fl. I'^ur. Alg., ii.,
p. 193; Tilden, I.e., p. 192, PI. ix., t'.i8. In the type, the hetero-
cysts are spherical, and the gonidia are shorter. Cell-contents,
in our form, pale blue, in the heterocysts and gonidia a deeper
blue. Ptabenhorst, I.e., gives '■^frichomatibtis . . . liUeo-fernyineis"
and ^'sporis . . so.tjirate luteo-virldihus^'' for the type.
(jienus C Y L I X D R O S P E R M u M Kiltz.
Cylindrospekmum stagnale v. australe, n.var. (PI. liv., f. 17-21).
Forma cellulis cylindraceis, disjunctis, apicibus truncatis ;
heterocystidibus oblongo-cylindraceis vel conicis ; gonidiis ob-
longis vel oblongo-cylindraceis, apicibus rotundatis, cvtoplasmate
pulchre cyaneo vel dilute viridi.
(1). Cell. diam. 4-5, alt. 5-10 (plerumque 7-8); heterocyst. diam.
4-6, alt. 8-11; gonid. diam. 9-13, alt. 18-30/x. Lismore (302).
(2). Cell. diam. 3, alt. 4 10 (plerumque 4-5); heterocyst. diam.
3-4, alt. 7-14; gonid. diam. 7^-9^, alt. 11-12//. Lismore (306).
Cf. G. S. West, Br. Frw. Alg., p.328, f.l50; Tilden, Minnesota
Algie, i., p. 198, PI. X., f.2. The type has oblong cells; in this
form, they are quadrate or cylindrical. 1 found it once (sample
306) as a grey-green, foliaceous, mucous stratum at the grassy
edge of a swampy pool. Though the cells and heterocysts of the
interlaced trichomes were all of the same size and shape, there
were intermingled in this one stratum three distinct tyjjes of
gonidia. The most frequent answered exactly to C. stagnale;
another I have already described as C. rectangalare; and the
506 XEW AND RARE FRESHWATER ALG^E,
third seems to be a form of C. Goetzei Schmidle. T have
accorded these forms specific rank, as it is the general practice,
and cumenient for classification: besides, what should be char-
acteristic of a species in these plants, if not the shape of the
gonidia! Nevertheless, their being intermingled in one mucous
stratum indicates that they are merely allotropic forms of one
plant, brought about by different modes of growth.
Most of the trichomes in the mucus were merely interlaced,
but here and there they were closely coiled in a spiral manner
(PL liv., f.21). This is not uncommon in free-swimming filaments
of the plankton, but very unusual in mucous strata.
Cylindrospermum rectangulare Playf.
(1). Gonidia thin-walled, rectangular: heterocysts oblong or
acutely conical.
Cell. diam. 3, alt. 4-10; heterocyst. 3-4, alt. 7-14: gonid. 4,
alt. 14/x.
Lismore (306). (PI. liv., f.22).
(2). Gonidia thick- walled, cylindrical: heterocysts oblong.
Cell. diam. 4, alt. 5-10 (plerumque 7); heterocyst. 4-6, alt. 8|-
\0h; gonid. diam. 9i-12, alt. 29-38/x.
^ Lismore (302). (PI. liv., f. 23).
Cf. Frw. Algfc of the Lismore Distr., these Proceedings, 1915,
p.349, f.A.
Var. PARVICELLULA, n.var. (PI. liv., f.24, 25).
Forma cellulis minimis, plerumque adpressis, quadratis vel
diametro brevioribus, cellula apicali acute-conica; heterocystidi-
bus oblongis; gonidiis interdum lateribus paullo arcuatis.
Cell. diam. 2-2|; heterocyst. 3i, alt. 7; gonid. 10, alt. 24/x.
WoyWoy(270).
In a gathering kindly sent me by Mr. A. H. 8. Lucas. The
cells are unusually small for plants of this family: it should be
noted, too, that the gonidia are not any smaller on that account.
The walls of the latter were incrassate, rufescent and scrobicu-
late; this takes place with age, however, in the gonidia of every
species.
BY G. I. PLAYFAIR.
507
Cylindrospermum Goetzei Schm., forma. (PL liv., f.26, 27).
Trichoniata cUigusta, ad genicula constricta; cellulis quadratis
vel cyliiidraceis, disjunctis: heterocy«tidibus oblongis vel conicis;
goiiidiis laiiceolati.s vel subdoliformibus, apicibus arigustis truii-
catis.
Cell, diaiii. .'^5, alt. 4-10; heterocyst. 3-6, alt. 6-14; gonid. 7-10,
alt. 14-2:
Lismore (306).
C rectangulare Flayt, parthn, Frw. Alg. Lismore, p. 349, f. A.
Of. Schmidle, Schiz. Conj. Chlor. in Engler's Flora von Afrika,
H. xxii., p. 2 4 5, T. iv., f.5. In this, as in all other species of the
family, the cells and heterocysts may vary much in shape and
size, but the form of the gonidia is characteristic. With the
three species of Cylindrospermjim mentioned above, compare
Anah(Hna oscUlarioides (type), its var. stenosj^oi-a Born. <fe Flah.,
and A. Volzii Lemm. These show the same three allotropic
forms of gonidia in what is, biologically, one species; they are all
found here with the same (quadrate or cylindrical) cells and
heterocysts, very often also accompanying one another.
Fam. SCYTONEMACE.^.
Genus T o l y p o t h r i x Kiitz.
ToLYPOTHRix LANATA (Desv. ) Wartmanii.
(Text-fig. 3).
Diam. fil. 15-17, trich. 14; cell. alt. 4-8
(plerumque 4); heterocyst. 15 x 14//.
Grafton, Nymboidia R. (265).
Our specimens agree in dimensions
fairly well with those given by Raben-
horst, Fl. Eur. Alg., ii., p. 277 (fil. diam.
14-181-, trich. 11-15/x), but the cells are
much shorter (equal to, slightly less than,
or half as long as the diameter — Rabh.).
Hheatli colourless, J to IJ/x thick, trich-
omes constricted at the dissepiments,
cytoplasm pale green or pale grey-green,
very finely granular.
Text-fig. 3. — Tolypothrix lanata (Desv.) Wartmann; ( x 660.
508 nkw and rare freshwater alg^,
FLAGELLATtE.
Fani. HYMENOMONADACE.^-:.
Genus 8 y n u r a Ehr.
Synuka (GRANULOSA Playf. (PL Ivi., f. 1-3).
I give another figure of the cells of this species, that in Fi-w.
Alg. Lismore, PL xlv., £.3, being shown too finely granulate. The
granules are quite coarse, no more than 16 being visible at the
edge, round the anterior margin. Home, that T noted lately, had
from 1 to 8 minute dark red, or nearly black, irregular granules
at the base of the fiagella; or, sometimes, one, larger, simulating
a stigma, in the gap between the chloroplasts (PL Ivi., f.2, 3).
The chromatophores in this case were a very pale green with(jut
any trace of yellowness, and the crenobia peculiar in being oblong
(not globose and stellate), with the cells attached round a central
peduncle. The origin of these oblong ccenobia becatne clear
later, when I found a long cylindrical ccenobium (80/x long by
30/x broad, cells long. 12-20, lat. 6-10/x) with a central mucous
or membranous peduncle (PL Ivi., f. 1). The latter being gradu-
ally drawn apart by the perpetual twisting of the cells, the
c<enobium broke up into two oblong colonies. In the same way,
these probably break up into smaller groups, which become
regularly stellate by cell-division. The formation of the ccenobia
in this genus would appear, therefore, to be on somewhat the
same lines as in Anthophysa vegetans.
Genus T e s s e l l a r i a Playf., nom. emend.
Cf. Frw. Alg. Lismore, p. 3 15. I find that ^^TesselW^ is pre-
occupied by Ehrenberg for a genus of diatoms; I hav^e altered
the name of my new genus, therefore, to Tessellaria.
Tessellaria volvocina Playf.
In PL Ivi., f.I, I give a figure of a yellow-green fiagellate evi-
dently belonging to this family, which I think is very probably
a free zooid of this organism. In shape globose, truncate in
front, diam. 10/x, with two long fiagella, two small yellow-green
chromatoiDhores confined to the anterior part of the cell, and a
number of relatively large refractive globules scattered round
BY G. T. PLAYFAIR. 509
just inside the membrane: moml:)rane very delicate. One e.v.
was noted at the side, but perhaps more were present concealed
by the globules.
This organism in its early stages seems to be distinctly a
plankton-form and to have a fondness for the surface-layer of
water. Only later, when the coenohia have become heavier, do
they fall to the bottom and become ground-forms. On the
surface of a gathering which was being decanted, T was able to
descry, with the help of a Coddington lens and a ray of direct
sunlight, quite a miniature world of micro-Hora and fauna,
among which there turned out to be numbers of TpsspJlavia
ccenobia of varying sizes but all small.
Incertce sedis.
Genus Xanthodiscus Schew.
Xanthodiscus Lauterbachi Schew. (PI. Ivi., f.5, 6).
Schewiakow, Geogr. Verbreit. d. Siisswasserprotozoen, Mem.
Acad. sc. de St. Petersbourg, Ser. 7, T. xli., 1893. A rare
flagellate, about whose position and characteristics there seems
to be some uncertainty. Wille, Conj. und Chlorph., p. 21 (in
Engler k, Prantl, Die naturlichen Pflanzenfamilien) has placed it
as a doubtful genus of the Volvocacece, relying, apparently, on a
certain similarity to Ghhiinydomojias in the arrangement of the
cell-contents (fig.7D, E). T first obtained it in great abundance
out of M yriophylliinn in the Orj^han School Creek, near Canley
Vale Railway-station (March, 1909), and afterwards from Fair-
field; I have noted it also frOm Gardener's Road, Botany. Un-
fortunately, the disposition of the contents was vague, and my
lenses at that time not good enough to distinguish any details.
My recollection is, that the chromatophores were pale yellow-
green (not brown-green as described). One point, however,
which is quite certain, has not been noted either in the figure or
description (/.c), viz., that the cell-wall is in two parts, as in
Phacotns, merely agglutinated together. Tlie organism appeal's
to be a freshwater survival of a large mai'ine family of flagellates
— the Frorocentracece — distantly related to the Peridiniect,
510 NEW AND RARK FRPISHWATER AhdM,
Stein, in Naturg. d. Flagell., ii., T. i., f. 27-38, figures it as Dino-
pyxis fcevis Stein. He shows a pair of c.v. in front; the large
posterior globule, he considers the nucleus; and the smaller central
bodies, pyrenoids (two appear in his figures). Lemmermann,
however, Reise n.d. Pacific, p. 361, mentions Dinopyxis /cevis as
a synonym of Exnviella Lima (Ehr.) Schtitt, {Crypfomonas Lima
Ehr.), quoting Schtitt, G^^mnodiniacese, p. 8, fig. 9 (in Engler c^-
Prantl, I.e.). Schewiakow's genus and species, therefore, would
seem to be invalid. As a freshwater form, it is kno^n only
from Australia.
CONJUGATE.
Genus S p i r o g y r a Link.
Spirogyra Lismorensis Playf.
Biol. Richm. River, p.28, PI. iii., f. 1. Noted also from the
Nymboidia River (Grafton Water-supply), infertile. It seems,
therefore, to be a plankton-form.
Spirogyra mirabilis (Hass.) Kutz. (Text-fig. 4).
Diam. cell. veg. 15-23, alt. 230-245: cell.fruct. max. 25-42, alt.
180-190; zygosp. 21-34, alt. 44-93/x.
Lismore (308).
Cf. Petit, Spir. de Paris, p. 14, PL iii., f.3, 4; Borge (in Pascher,
Susswasserti. Deutschl.) Zygnemales, p.21, f.l7. This seems to
be the first record of the conjugation of Sp. mirahilis, as Borge,
I.e., 1913, says ^'ICopulation nicht hekaimt'' The vegetative cells
were very long, 10-16 times the diameter; there was one chloro-
plast making 3 J- 4 turns. With this species should be compared
Sp. Spreeiana Rabh. The two are so very much alike, both in
characteristics and dimensions, that, in spite of the replicate cell-
ends of the latter, I am not convinced that they are distinct
species. Moreover, Hirn, Finlandischen Zygnemaceen, p.ll, f.4,
in Sp. kuusamo'ensifi ( = a more slender form of Sj^. Sjyyeeiana)
and in /Sp. Spreeia7ia itself. I.e., p. 12, f.5, shows both simple and
replicate septa in the same filament. So also Borge, in Spirogyra
spp., Sibiriens Chloroph., p. 8, f. 2, 3 { = Sp. SpreeAana stouter
BY G. T. PLAYFAIR.
511
form. For convenience of comparison, T append a Table of the
Text-fig. 4.
Sipirotjyra mirahilis (Hass. ) Kiitz ; (a) conjugated cells ( x 330); {h, r) two
sizes of zj'gospore, {h x 660, c x 500); {d) form with broader filaments
and larger, more slender zygospore, ( x 330),
dimensions of all these forms; in the characters of the cells,
chloroplasts, and zygospores tiiey are all one.
Diam.
Zygo.
cell. veg.
cell, fruct.
diam.
alt.
.S^. mirahilis
18-27
• —42
24-29
U-2plo.
aplanc
Ours
15-23
25-42
21-34
{
44-93
2-3 plo.
Sp. Spreeiana \
18-24
30-42
30-36
2-3 plo.
(Borge, Zygnem.)i
Sp. kunsamoensis...
13-17
25-40
23-33
45-75
2-3 plo.
Spirogyra spp. . \
26-31
•'. 40-48
.30-46
(
07-118
Borge J
I
2.^ plo.
Let Sp. kuusamoensis be united with Sp. Spreeiana, and ou]
512 NEW AND RARE FRESHWATER ALGiE,
specimens agree perfectly with them in all points, as well as with
Sp. mirabilis.
SpiROGVRA NEGLECTA (Hass.) Klltz. (PI. Iviii., f.l 13).
A. — Cell. veg. diam. 46-57, alt. 53-360; cell, fruct. diam. ad
dissep. 46-57, ad inflation. 65-68, alt. 42-125/^. Zygospora^ ut
sequitur: —
a. Zygo. late ovales, long. 52-66, lat. 42-49//. (PI. Iviii., f.3).
h. Zygo. longiiis ovales, long. 64-74, lat. 45-51/j(.. (PI. Iviii., f.4).
c. Zygo. elliptic^, long. 78-91, lat. 46-51/x. (PI. Iviii., f.5).
cL Zygo. oblongse, long. 75-80, lat. 45-51/x. (PI. Iviii., f.6).
Lismore (353).
Cf. Petit, Spirog. d. Paris, p. 26, PL ix., f.l -5; Borge, Zygne-
males, p. 2 9, f.3 6. In the very short cells, there are generally 3
chloroplasts making 1 turn: in the long cells, 3-5 chloroplasts
making 2-3 turns. Three forms of chloroplast also were noted
intermixed; all, however, had the central ridge, which is said to
be characteristic of the species: —
(1). As in Petit, /.c, a narrow delicate band with minutely
irregular edges, and very small, almost indistinguishable, pyre-
noids. Only noted in uncopulated cells of fertile filaments.
(PI. Iviii., f.7, 8).
(2). A very delicate band with finely fimbriated edges; also
with very minute pyrenoids. (PI. Iviii., f.9, 10).
(3). A more robust band with edges irregularly scalloped, not
fimbriated, and with large, distinct pyrenoids. (PI. Iviii., f. 1 M3).
Four distinct shapes of zygospore were noted in this one
sample, and, in such profusion, as to be all present under the
cover-glass at one time. Inflated sporangial cells, and uninflated,
were noted in the same conjugated filament (PI. Iviii., f.l, 2).
B. — Cellular vegetativse paullo crassiores ; zygosporis paullo
majoribus; chloroplastidibus tenuibus laciniatis.
Cell. veg. diam. 62-64, alt. 70-120; zygo. long. 94-100, lat.
52-56/x. Lismore (297).
A broader specimen than usual, with slightly larger zygospores.
In gatherings of Sp. neglecta there may generally be noted a few
BY G. T. PLAYFAIR. 513
filaments of Sp. nitida, easily (listingiiishecl l)v their greater
l)i-eadth:— diam. cell. veg. 95-116, alt. 84-360; 3-5 ehloroplasts
making '!-?> turns, edges scalloped, a central ridge evident here
and there, large pyrenoids 6-10 to the half tuiii. (Lismore, 352,
353, 358). It is not unlikely that they really form one species.
Sj). FulJebonnl 8chm., Alg. aus Nyassasee, p. 76, T. iii., f.2 (fil.
diam. 40-42, zygo. 64-80 x 40//) seems to me a somewliat narrower
form of Sj). neglecta.
Var. AMYLACEA, n.var. (PI. Iviii., f. 14-1 6).
Oelluhe vegetativte chloroplastidibus amvlo suflusis, in granula
fractis; pyrenoidibus maximis, granulosis.
Cell. veg. diam. 46 54, alt. 60-260: zygo. long. 74-90, lat. 48-50//..
Lismore (271).
In this form, the chloroplasts have become impregnated with
amylum, and have broken up into minute irregulrir gi-ains. The
central ridge, however, is generally still noticeable, and the
pyrenoids are very large (up to 10-1 2/x), and gr^-nular as in
Closterium hinula. The whole of this gathering was of the same
character. The zygospores present were long-elliptic with
pointed ends, lying diagonally in short oblong cells which were
not inflated (as in PI. Iviii., f. 1). Probably, however, all the
various forms of zygospore detailed above for the type will be
found also in this variation. Chloroplasts 2-3, making 1| to 3|
turns in the cell.
Genus Mougp]otia (Ag.) Wittr.
MouGEOTiA L^TEViRENS (A.Br.) Wittr. (Text-fig. 5).
Cell. veg. diam. 27-44, alt. 255-408; zygo. long. 61-63, lat. 42-53//.
Botanic Gardens, Sydney (158); Lismore (328, 332).
Cf. Borge, Zygnemales, p.40, f.62. In our specimens, the
chloroplasts generally have a large number of p3^renoids, either
irregularly scattered or arranged in two lines at the edges. This
is the only species of Mougeotia, as fai- as I know, in which this
occurs. The pyrenoids are sometimes compai'ati\'ely small (2-4/x)
but occasionally very large, diam. 10//. The contents of the cyst
39
14
NEW AND RARE FRESHWATER ALG^,
are generally retracted into an oval (53 x 32/x), oblong (53 x 4:2/x)
or more rarely globular (diam. 42/x) zygospore.
Text-fig. 5.
Mougeotia Icetevirens (A.Br.) Wittr. ; (a) conjugating filaments with unusu-
ally large pyrenoids; {h, c, d, e) types of zygospores; (all x 330).
CHLOROPHYCE^.
Fam. VOLVOCACE^.
Genus C a r t e r i a Diesing.
Carteria rugulosa, n.sp. (PI. Iv., f.l^ 2).
Cellulse reniformes, in fronte acuminatse, a tergo concavse,
angulis posterioribus rotundatis. A vertice late ellipticse. Meni-
brana paullo rufescens, jugis granulatis convergentibus 12, longi-
tudinaliter dispositis, ornata. Cytoplasma retracto gioboso,
stigmate juxta apicem.
Long. 17, lat. 23, crass, 17/x. Lismore(311, 312).
A most curious form, noted in some numbers. The cell- wall
is much larger than the mass of cytoplasm, and is reniform or
rather broadly obcordate in shape, with 12 coarse longitudinal
ridges bearing a series of granules. The ridges converge to a
point in front and behind. The cell-wall was rufescent, and the
BY G. I. PLAYFAIR. 515
arrangement of the chloroplasts could not be discerned; a distinct
stigma near the apex was noted, but no pyrenoid.
Var. ANGULATA, n.var. (PI. Iv., f.3).
Forma ad apicem depressa, in medio depressione papilla in-
structa; lateribus inferne paullo angulatis; ceteris ut in forma
typica.
Long. 17, lat. 23/jt, Lismore. Cum pi-iori.
A cup-shaped depression with central papilla replaces the
pointed apex of the type. The sides run back to a slight angle
in the hinder portion.
Var. IN.EQUALIS, n.var. (PI. Iv., f.4).
Forma major, ovata, in fronte acuminata, a tergo late-rotund-
ata. Membrana luteo-fuscescens: ceteris ut in forma typica.
Long. 30, lat. 21-25J/X. Lismore. Cum priori.
In all specimens of this form there was a minute projection at
one side of the apex. Undoubtedly, I think, this variation is
produced by uneven growth, only one lobe of the cell-wall devel-
oping; the small subapical projection is the remains of the un-
developed lobe.
Carteria multifilis (Fresen.) Dill. (PL Iv., f.5, 6).
Cellulse parviB sph^ericse; membrana tenuissima vel interdum
crassa; papilla nulla nee cytoplasmatis projectione; chloroplastide
crasso parietali; pyrenoidi in media cellula.
Diam. cell. 8-11; pyrenoid. 3/^. Lismore (345).
Cf. Fresenius, Beitr. z. Kenntn. mikrosk. Organism., Abh. d.
Senckenb. Gesells., 1856, p.235. Goroshankin, Morph. u. System,
d. Chlamyd., ii., 1891, p. 120, reproduces Fresenius' original de-
scription of the type, the chief points of which are: — cell spher-
ical or oval, length 10-16/x, pyrenoid in the centre of the cell,
cytoplasm minutely granular, stigma between the apex and the
middle, membrane very delicate, close-fitting.
I have not seen Fresenius' figure of the type, but our specimens
seem to me in some respects to tally with his description better
than other published figures, and specially in this, that they have
516 NEW AND RARE FRESHWATER ALGvE,
a massive parietal chloroplast with a central globular locellus in
which the pyrenoid (when present) is situated.* Excellent
figures are given by Goroshankin, I.e., PL ii., f.l4, and by G. S.
West, Br. Frw. Alg., p. 18 8, f.73A, B, but in each case the
pyrenoid is eccentric. Fresenius says '■'■In der Mitte p'ln scharf
umschriehener Kern.^' Ours, on the other hand, are smaller,
(Rabenhorst, however, rejDorts specimens of 6-8/x diam.) quite
spherical and without any apical protuberance. The cell-wall,
generally very thin, is sometimes stratified. This species, con-
sidered the most common in Europe, is very rare here, in my
experience. I obtained it (the only time) from two local ponds
in October, 1917, after the spring rains, in some quantity-
Though I had assiduously worked these pools for five years pre-
viously, I had never seen a sign of it before, nor have I noted it
since from either.
Carteria bullulina, nom.nov.
Syn., Carteria multifilis, forma, Australian Frw. Phytoplankt.,
these Proceedings, 1917, p. 824, PL Ivi., f.1-4. The expression
"closely fitting membrane" {dicht anliegenden Hant) in Frese-
nius' description of C. imdtifilis rules out this form from any
connection with the latter. My description. I.e., applies to the
cell-membrane ; the mass of cytoplasm is much smaller and
attachea at the apex (fig.l). The details of the chloroplast, I
have not yet noted.
Carteria granulosa, n.sp. (PL Iv., f.7, 8).
Cellula compressa, fronte visa psene circulata, superne fissura
obliqua instructa; a latere valde compressa, superne in cornua
duo divisa, cornibus evolutis. Membrana rufescens, granulosa.
Cytoplasma retractum, ad cellulse apicem affixum, a latere com-
pressum. Pyrenoidis nulla nee stigma (in speciminibus notatis).
Long. 31|, lat. 29|, crass, 10/x. Lismore.
A compressed form, with granulate cell-wall. Above, the cell-
wall is divided into two rounded overlapping heads by a deep
* When the pyrenoid, as in this ease and manj' others that I have
noticed, is. central, it would seem to enclose the nucleus.
ftT G. I. PLAT FAIR. 517
oblique cleft. In sideview, both cell-wall and green cell are seen
to be compressed, the cell-wall cleft above and the two heads
turned outwards. No pyrenoid in the specimens observed.
Nucleus and stigma not visible, the membrane strongly rufescent.
Genus C h l a m y d o m o n a s Ehr.
Chlamydomonas reniformis, n.sp. (PL Iv., f. 9, 10).
Cellula compressa; fronte visa reniformis ubique rotundata,
pone indentata. A latere compressa, ovalis vel ovata. Membrana
tenuissima glabra, cytoplasmati arete adhaerens. Pyrenoidis
nulla nee stigma (in specimine notato).
Diam. 10^, alt. 8-|//.. Lismore (312).
A very curious form, of which I saw but one specimen. It
ver}^ much resembles Garteria rugulosa, and might possibly be a
young form of that species. The flagella (2) were very distinct
and unusually thick. On the other hand, the cell-wall is closely
adherent to the cytoplasm, perfectly smooth and hyaline.
Chlamydomonas Lismorensis v. gracilis, n.var. (Pl.lv., f.ll).
Cellula anguste cylindracea, pone rotundata, fronte conica,
lateribus parallelis. Chloroplastis parietalis ; pyrenoid e nulla
visa; stigmate distincto inter apicem et medium posito; granulis
paucis juxta apicem.
Long. 9J, lat 2J/x. Lismore (345).
The type (these Proceedings, 1917, p.827, PI. Ivi., f.l8) is ob-
long-cylindrical with rounded ends, and proportionately broader.
This form is narrower, and conical in front. Chloroplast bright
green, homogeneous, parietal; no pyrenoid seen; stigma wick-
shaped, very distinct, halfway between the apex and the middle.
Chlamydomoxas rotula, n.sp. (PI. Iv., f. 12).
Cellula oblonga (vel globosa ?) ubique rotundata; in media
cellula pyrenoide magna; chloroplastide in discos teiiues parietales
multos diviso, discis singulis ad pyrenoidem taenia chlorophyllacea
connectis; stigmate nullo viso.
Long. 19-21, lat. 15; pyrenoid. 6/y.. Lismore (345).
Easily recognised by reason of the extraordinary disposition of
518 NEW AND RARE FRESHWATER ALGM,
the endochrome. The chloroplast is broken up into a number of
very delicate parietal discs, each connected with the large central
pyrenoid by a filament narrowing from without inwards. There
are about seven discs visible round the margin in optical section.
C£. Chi. stellata Dill, Gatt. Chlamyd., p.l7, T. v., f.31-36; Wille,
Alg. Notizen ix.-xiv., p. 134, T. iv., f.4. The latter, however, has
a massive chloroplast, and differs considerably in appearance.
Chi. rotida also has a papilla or apical protuberance. A Glmo-
cystis-stage was noted, eight cells in a mucous sphere; the cells
were in every respect the same as the zoospore. I have given
this species the name '^rohda," as 1 have my suspicions that
Oocystis rotula Playf., is a young Gloeocystis-^tagQ of it.
Chlamydomonas alpina (Wille) mihi. (PI. Iv., f.l3).
Chloromonas alpina Wille, Alg. Notizen ix.-xiv., p. 152, T. iii.,
f.24-34.
Cell. long. 10 J, lat. 7/x. Lismore.
Noted both with and without a pyrenoid, the latter situated
close to the posterior end of the cell. The chloroplast-discs are
about 2-3/x in diameter. Two c.v. observed, and an oval orange-
coloured stigma in the middle of the cell (near the apex in the
type). In the specimen measured, the Hagella were very long,
quite twice the length of the cell, but I did not notice if they
were so in every instance. Noted in winter, among fungoid
growth on a rotten stick in swamp-water.
Chlamydomonas caudata Wille. (Pl.lv., f.l4).
Algologische Notizen ix.-xiv., p. 135, T. iii., f.4-11. The only
tailed species of Chlamydomonas; it is impossible to make any
mistake in the identification. Yet there was not a trace of n
pyrenoid, though the body was very transparent, and the nucleus
distinctly visible. Stigma wanting.
Cell. long. 30, lat. 18; corp. long. 20, lat. 12//. Lismore (344).
Chlamydomonas maculata, n.sp. (PI. Iv., f. 15-17).
Cellulse sphaericie, papilla nulla nee projectione apicali; mem-
brana crassa interdum lamellosa ; vacuolis contractilibus 2 ;
stigmate distincto, hemisphserico, luteo-fusco, paullo supra medium
BY G. I. PLAYFAli?. 519
posito; niicleo in media cellula. Chloroplastide valida parietali,
inaculata (superficie externa scrobiculata); pyrenoide plerumque
nulla (in speciminibns notatis).
Diam. 15-25; membr. 2-4; cell, matric. 40, aiitosp. 11-13//.
Lismore (345, 350).
Compare Chi. subcaudata Wille, Alg. Notizen ix.-xiv., p. 11 8,
T. iii., f. 12-1 8; W. & G. S. AVest, Frw. Alg. Brit. Antarct. Exp.,
p.274, PI. xxiv., f.25-29. Generall}^ spherical without apical
projection; membrane thick, sometimes lamellar in two layers.
It is distinguished from Chi. glohulosa Perty, by the massive
parietal chloroplast evenly distributed round the cell-wall except,
of course, at the apex, and particularly by the fact that the outer
surface of the chloroplast is pitted, giving it a spotted appear-
ance all over. The central locellus is globular. Nucleus central;
stigma orange-coloured, a little above the middle ; pyrenoid
generally wanting. A mother-cell, however, was noted, contain-
ing four autospores, of which one had a distinct pyrenoid; in two
others, it was faintly indicated, and, in the fourth, it was entirely
absent.
Var. OBLONGA, n.var. (PI Iv., f.l8),
Cellul* oblongi^; ceteris ut in forma typica.
Long 16, lat. 13|-/>t. Lismore (350).
Var. PLANKTONICA, n.var. (PL Iv., f.l9).
Celluhe oblongje, interdum apicem versus modice angustatse;
humili projectione apicali instructye, membrana tenuissima arete
adh^rente; stigmate nuUo nee pyrenoidi (in speciminibus notatis);
ceteris ut in forma typica.
Long. 20-26, lat. 13-17/x. Lismore (347).
This form has the same parietal chloroplast with scrobicula?
on the outer surface, but the membrane is very thin; there is a
slight apical protuberance, and, in specimens observed, the stigma
and pyrenoid were absent.
Chlamydomonas metastigma Stein. (PI. Iv., f.20).
Forma stigmate luteo-fusco, bacillari, subapicali.
Long. 26, lat. 13/x. Lismore (327).
520 NEW AND RAIiE fRESHWATER ALG/£,
Cf. Stein, Natiirg. d. Flagell., i., T. xv,, f.46 : Gorushankiii,
Morpb. 11. System, d. Chlamyd.^ ii., p. 131, PL iii., f.26: Wille,
Alg. iS'otizen ix.-xiv., p. 146, T. iv., f.22. Goroshankins figure
(reproduced by Wille) differs from the type in being a short oval:
whereas Stein's figure is a long oval, subcylindrical. Our speci-
mens agree with the type in outline and in proportions (Steiiis
figure works out at 20 x 11/x), but differs in the position of the
stigma, which is near the end in the type, but subapical in ours.
Goroshankin makes it hemispherical, here it was wick-shaped.
The upper pyrenoid was laterally disposed, and perhaps was
geminate, or becoming so; Goroshankin notes the occuri'ence of
two pyrenoid s in front.
Chlamydomonas paupercula, n.sp. (PL Iv., f.21, 22).
Cellulae longo-ovata?, subelliptica?, in medio latissimte, pone
paullo angustatsB, fronte conicae; apice obtuso paullo producto;
membrana tenuissima inconspicua arete adhterente; chloroplastide
crassa parietali, granulis sparsis superficie exteriore vel interiore
dispositis; stigmate nullo nee pyrenoidi (in speciminibus uotatis).
Long. 14-17, lat. 6-8//. Lismore.
In shape and size, very like Chi. alboviridis Stein, Naturg. d.
Flagell., i., T. xiv., Abt. vi., f,l, 2, 21; (figures 1, 2 work out at
26 X 13//, fig. 21 at 17 X 8/x). The cell is elliptic, conical in front,
the apex obtuse and slightly produced; membrane very thin.
The chloroplast is massive and parietal, with scattered granules
sometimes on the outer surface, sometimes on the inner. Though
viewed with the yV obj., the nucleus was not discernible; pyre-
noid and stigma absent.
Chlamydomonas pusilla, n.sp. (PL Iv., f.23).
Cellula3 ovata?, ubique rotundatJie; apice obtuso baud producto;
membrana tenuissima, inconspicua, arete adhjerente; chloroplast-
ide crassa, parietali; intus granulata; nucleo in media cellula;
stigmate mediano; pyrenoidi nullo viso.
Long. 30, lat. 20//. Lismore (350).
Cell broadly ovate, rounded off" everywhere, obtuse in front
and without projection. Chloroplast massive, parietal, granulate
ftY G. t. PLAYFAItf. 621
oil the inner surface, no pyienoid in the specimens observed,
stigma median. The nearest form seems to be Ghl. media
Klebs, Beding. d. FortpHanz., p.425, f.l2, (Wille, Alg. Notizen
ix.-xiv., p. 140, T. iv., f.ll) which, however, is more pointed in
front, and has a siibapical stigma and a pyrenoid at the side in
the middle.
Chlamydomoxas angulosa v. obesa, n.var. (PI. Iv., f.24).
Cellulae prje latitudine breviores, crasste, subglobosse; de singuHs
rebus ut in forma typica.
Long. 16, lat. 14/jt. Lismore (350).
Cf. Dill, Gatt. Chlamyd., p.l5, T. v., f.21-25; Wille, I.e., p.l41,
T. iv., f.1.3. Oiu" form agrees in all details with Dill's type, but
in shape it is subgiobose, almost subquadrate, instead of oval.
Dill's figure works out at 20 x 14/^..
Chlamydomonas pisiFORMis V. ocELLATA, n.var. (PL Iv., f.25).
Cellulw ambitu formte typical consimiles; chloroplastide autem
parietali; in media cellula locello magno pyrenoideiii continente:
stigmate nullo viso.
Long, 17, lat. 8i: cell, matric. long. 25, lat. 19//.
Lismore (350).
Cf. Dill, I.e., p.l4, T. 5, f. 13-19: Wille, I.e., p.l3S, T. iv., f. 8.
Only noted as autospores, 2, 4, 8 to the mother-cell. In shape,
they agree exactly with the type, even to the slight curve to one
side, and the apical projection, broad from the front and wick-
like from the side. The chloroplast, however, is parietal, and
the centre of the cell occupied by a large locellus containing a
pyrenoid (diam. 5/x). No stigma, and the nucleus not discernible.
Genus C h l o r o (i o n i u .-si Ehr.
Chlorogonium minimum, n.sp. (Pl.lv., f.26).
Cellula? minutte, gracillimae, fusiforines, pone acutisshna^ froiite
rostrat?e; membrana tenuissima, arete adhyerente; chloroplastide
tenuissima, homogenea, parietali; in media cellula locello hyalino
(hiucleum contingente); flageUis 2, ex apice oblique orientibus;
stigmate distincto, bacilliformi, subapicali; pyrenoide nulla.
522 NEW AND RARE FRESHWATER ALGA,
Long. oO, lat. 2-3/7.. Auburn; Lismore (350).
A minute and very slender species, rostrate in front and \'ery
acute behind; membrane very delicate. Chloroplast also very
delicate, homogeneous, parietal, without pyrenoids. A hyaline
space, probably containing the nucleus, in the centre of the cell.
Stigma very distinct, wick-shaped, some distance down from the
apex. Contractile vacuoles not noted. Flagella 2, going off at
right angles close to the tip.
Genus Pteromonas Seligo.
PterOxMONAs angulosa v. australis Playf.
Syn., Pt. alata v. australis Playf., Frw. Alg. Lismore, p. 335,
PI. 42, f.l4. Pt. alata Seligo, is a synonym, it appears, of Pt.
angulosa (Carter) Dang., {Cryptoglena anyulosa Carter, Ann. Nat.
Hist., 1859, T.l, f.l8a-6'). The side-view (PI. Iv., f.28) is char-
acteristic of the type and all its forms, cf. Chodat, Alg. vertes,
p. 144, fig. 68 E.
Forma. (PI. Iv., f.27, 28).
' Membrana a tergo incisa. Long. 22J, lat. 15/x. Lismore(312).
Var. 8CUTIF0RMIS, n.var. (PI. Iv., f.29).
Cellulie membrana oblonga, fronte paullo angustata; lateribus
inferne rectis, levissime convergentibus, apicem versus incurvis;
apice levissime concavo ; augulis posterioribus rectis ; basi in
medio inllata.
Long. 16, lat. 11; corp. lat. 8/v.. Lismore (344).
The enveloping membrane is oblong, with straight sides con-
\ erging slightly, and drawn together in front, the apex a little
concave. Lower angles square, and at the base convex. In all
these forms observed here, the chloroj^lasts, though very pale and
thin, showed the characteristic cup-shape. No stigma, however,
was present, nor pyrenoid.
Var. VEXiLLiFORMis, n.var. (PI. Iv., f.30).
Cellulse membrana fere quadrata; lateribus rectis, parallelis:
apice piano; angulis superioribus oblique truncatis; basi convexa;
6Y G. I. PLAY FAIR. 523
angulis inferioiibus obtusis. Pyrenoides parvse 2, pauUo supra
cellulam mediam ad latera disposit?e.
Long. 20, lat. 16, crass. <'^: corp. lat. 11//. Lismore (344).
The eavelope is quadrate with convex base, straight parallel
sides, fiat apex, and the upper angles obliquely truncate. A
specimen was noted w ith two small pyrenoids, on either side one,
a little above the centre ot" the cell. No stigma present. For
another example of a pyrenoid on each side, see Chi. longistigma
Dill, (Wille, I.e., T.iv., f.21).
Genus Phacotus Perty.
Phacotus crassus, n.sp. (PI. Ivi., f.7, 8).
Cellulae fronte visse fere circulat?e, margine aspera, ad apicem
papilla instructse; a latere paullulo compressa;, oblongae, lateribus
arcuatis, apicibus late-rotundatis, papilla instructis. Membrana
crassa, aspera. Cytoplasma in globo retractum, densum, crasse
granulosum ; vacuolis contractilibus 2 ; stigmate nullo nee
pyrenoidi.
Long. 22, lat. 19, crass. 14/x. Lismore (311).
Nearly circular in face-view, with apical papilla; oblong some-
what compressed from the side, with broadly rounded ends and
arched sides. The membrane is thick, and rough with irregular
asperities. The cytoplasm is retracted into a globe beneath the
apex, contents dense, coarsely granular, two contractile vacuoles
noted but no stigma or pyrenoid.
Phacotus glaber, n.sp. (PL Ivi., f.9, 10).
Cellulai fronte visse fere circulate, ad apicem concavie, papilla
instructse; a latere modice ovato-oblongse, ad apicem truncatse, a
tergo fere hemisphiiericse, lateribus levissime arcuatis. Membrana
crassa, glabra, dilute rufescens. Cytoplasma in globulo parvo ad
apicem retractum; stigmate nullo nee pyrenoidi.
Long. 22, lat. 21; cytopl. glob. diam. 9^/x. Lismore (312).
In face-view, nearly circular with a slight concavity, and a
papilla at the apex; ovate-oblong from the side, sides slightly
arched, apex truncate, hemispherical behind. Membrane smooth,
524 NEW AND RARE FRESHWATER ALG^t.
thick, slightly coloured. Green cell remarkably small, retracted
to the apex: no stigma, no pyrenoid.
Forma. (PI. Ivi., f.ll).
Celluhe fronte A'is?e liaiid circiilatse sed modice quadratae,
iibique rotundatse; ad apicem qiiam levissime concavpe, papilla
nulla; ceteris ut in forma typica.
Long. 21, lat. 19, crass. 14:; corp. 13/^. Lismore (312).
Rather rounded-quadrate than circular in face-view, apex very
slightly concave, no papilla; other details as in the type.
Forma. (PI. Ivi., f.l2).
Cellulye ad apicem paullo quadratai, pone circulat?e; membrana
crassa, saturate rufescente; cytoplasmate in massa ovata retracto;
chloroplastide parietal! intus granulis amylaceis magnis ornata;
vacuolis contractilibus distinctis 2; stigmate mediano.
Long. 21, lat. 20; corp. long. 12, lat. 10//. Lismore (345).
In shape, intermediate between the two forms (sicpra), being
broad and "shouldered" above, and circular below. Membrane
deeply rufescent: cytoplasm retracted into a small ovate mass;
chloroplast parietal, granulate on the inner surface with large
granules; stigma distinct, median; two contractile vacuoles dis-
tinctly visible. These details are very rarely observable in
PhacotnK.
Phacotus australis, n.sp. (PI. Ivi., f. 13, 14).
Cellulie fronte vis;e fere circulatfe, ad apicem concaxie, papilla
insti-uctte; a latere valde compressse, ellipticse, apicibus acute-
I'otundatis. Membrana tenuis, glabra, dilutissime rufescens.
Cvtoplasma in massa apicali retractum, a latere visum com-
pressum, ovatum; stigmate nullo nee p^'renoidi.
Long. 20, lat. 21, crass, c. 8//. Lismore (350).
Almost circular in face-view with a slight apical concavity and
papilla ; from the side very compressed, elliptic with acutely
rounded ends. Membrane thin, smooth, pale biscuit-colour, with
a matt surface. Cytoplasm retracted, compressed, ovate, in side-
view: no stigma nor pyrenoid.
BY G. I. PLAYFAIR. 525
Genus 8 p o n d y l o m o r u m Ehr.
Spondylomorum quaternarium Eh J.
Ehrenberg, Beob, zweier generisch. neuei' Fornien d. Frilhlings-
gewassers bei Berlin, Monatsber. d. Berl, Acad. d. Wissensch.,
1848. CJ\ Stein, /.c, T. xviii., f. 30-34. 8yn., EnrkiRia cornuta
W. & G. S. West, Frw. Alg. Biuma, p.L>28, PL xii., f.19-21: Uva
Casinoensis Playf., Biol. Richm. R., p.l08, PI. ii., f.l3.
This flagellate seems to be very little known; it is not men-
tioned either by Chodat in Alg. vertesde la Suisse, 1902; nor by
G. S. West in Br. Frw. Algae, 1904. Lemmermann reports it
from a single locality in Sweden, but it is not included in Bach-
mann's Das Phytoplankton des Siisswassers, 1911. In my former
paper, I had not the assistance of Stein's excellent illustrations;
also the figure and description of this organism published in the
Micrographic Dictionary, p.721, PI. 3, f.23, are entirely incorrect
and misleading. The figure here given agrees almost exactly
with that by Stein, I.e., f.30, save that the stigma is above, not
below% the centre.
The coenobia are generally oblong, consisting of rings of four
cells, arranged one above another. They easily break up either
longitudinally or transversely; in the latter case, the resulting
coenobia are often hemispherical or rosette-shaped. The cells,
originally globose, very soon become broadly oval or ovate; the
hiembrane is very delicate, and, by reason of their very rapid
movement through the water, it is generally drawn out at the
back into a short pointed sac, at first very indistinctly outlined.
Later, however, it becomes stouter and more evident. The chlo-
roplast is cup-shaped, generally without a pyrenoid; there is a
central nucleus, two apical contractile vacuoles, and a stigma,
median or a little above or below the midline. The internal
details, however, are not easily seen, as the cytoplasm is gener-
ally very dense. In a zooid which became detached from a
coenobium, I was able to see distinctly that there are two un-
usually long apical flagella. Noted from at least four different
spots round Lismore.
Ccenob. long. 28-40, lat. 22-27: cell. long. 10-20, lat. 6-12/x.
526 NEW AND RARE FRESHWATER ALG^,
Merrylands (8ydney) ; Casino (189): Lismore (237. 246, 307,
344).
Yar. ROSTRATUM, n.var. (PI. Ivi., f.l6, 17).
Cellulse plerumque 4-8 in fascina conjunctse, t'ronte globosoe,
pone in rostrum acutum levissime curvatum productae. Cum
forma typica atque ejusdem dimensionis.
Coenobia generally small, with a few cells in a bunch or rosette;
cells globose in front, and behind drawn out into a sharp-pointed
slightly curved beak; cytoplasm and chloroplast filling the whole
cell, only the tip hyaline. Generally no pyrenoid or stigma.
Genus Volvulina Playf .
VoLvuLiNA Steinii Playf. (PI. Ivi., f.l8).
Coenob. diam. 70, cell. diam. 15, alt. 12/a. Lismore (350).
In a coenobium of the type with very transparent pale green
cells, a minute central nucleus could be observed. The flagella
originated close together, and, at their base, two contractile
vacuoles, as in 8tein, I.e. In some cells, a large orange-coloured
stigma on the margin in front. The inv^esting membrane of the
coenobium, as usual, very delicate, and the cells, which were
nearly spherical, were pressed close against it.
Genus V o l v o x Linn.
VoLvox AUREUS var. HEMisPHiERicus, u.var. (PL Ivi., f.l9, 20).
Coenobium rauco investiente tenui; cellulis vertice visis circu-
latis, diametro inter se distantibus; a latere plus minusve hemi-
sphsericis, basi plana. Cytoplasma granulis magnis amylaceis
singulis instructum, stigmate ad marginem; vacuolis contractili-
bus 3 circa marginem; pyrenoidi nulla.
Cell. diam. ^-^|x. Lismore (311, 328).
The investing mucus of the coenobium very thin, cells quite
close to the edge. From above, the latter are circular, about
one diameter apart, with a large amylaceous granule towards the
centre, stigma at the margin, and three contractile vacuoles at
intervals round the circumference. In side-view, the cells are
more or less hemispherical, with a flat face closely adpressed to
BY G. I. PLAYFAIR. 527
the investing mucus. The connecting filament arises at the
angle. Tn the shape of the cells, this form i-ecalls VolvuHna
Steinii, which also has, sometimes, a series of c.v. i-ound the
circumference.
VoLvox Carteri stein. (PI. Ivi., f.21, 22).
Coenob. diam. ad 900; cell. diam. 4-5, inter se distant. 7-10;
zygo. immat. diam. 44-46, matur. ad 60/j-.
Lismore (311, 327, 328).
Cf. Carter, Ann. Mag. Nat. Hist., Ser. 2, Vol. iii. 1 have not
access, unfortunately, either to Carter's paper or to that of Stein
(the species is not mentioned in his Naturg. d. FlagelL); but
Lemmermann, Das Plankton schwedischer Gewasser, p. 105, in
one of his very useful little reviews of various genera, gives the
name, with the remark that the zygotes "are said to possess an
undulate membrane." By this character, I was able to recognise
the species. The ccenobium is like that of V. tertius Meyer, with
globular cells which, even under the yV obj., show no signs of
connecting filaments. It is the immature zygote which has an
undulate membrane (cell. diam. 31-33, lat. max. 44-46/>t); when
mature, the undulations have been produced into stout, slightly
curved spines (cell. diam. 36-37, lat. max. 53-60/x). Several dozen
zygotes in the ccenobium. I have the species from two places at
Lismore; it would appear to be the first time that this Volvox
has been recorded since Carter discovered it in India, fifty years,
or so, ago.
Fam. PLEUROCOCCACE.E.
Genus Elakatothrix Wille.
Elakatothrix gelatinosa Wille. (PI. Ivii., f.l, 2).
Cell. long. 15, lat. 3/x. Sj^dney Water-supply (100).
Cf. Wille, Conj. u. Chloroph., p.38, fig.l8A-E ; W. k G. S.
West, Phytoplankton Eng. Lake Distr. (Naturalist, 1909), p. 291,
f.6. A ccenobium of four lately-divided cells (8 x 2/jt), and a
single full-grown cell (15 x 3//.) noted in the filtrates of the Sydney
Water-supply. This organism bears a suspicious resemblance to
Spirotcenia acuta Hilse {see W. k G. S. West, Brit. Desm., i., PI,
528
NEW AND RARK FRESHWATER ALG.E,
iii., f. 14, 15). Known liitherto only from Norwegian and English
lakes.
Genus D a c t y l o t h e c e Lagerh.
Dactylothece arcuatum, n.sp. (PI. Ivii., f.:3, 4).
Cellulfe sparsie in muco amorpho involutse; fronte visie lineari-
ellipticie, quam levissime arcuatje, apicibus obtusis; a latere rectai,
lineari-ellipticae, lateribus quam levissime convexis, apicibus ob-
tusis; membrana tenui; chloroplastide tenuissima, parietali, minute
granulosa, interdum in medio di^•isa: pyrenoidibus nullis.
Cell. long. 14 15, lat. 3-4/x. Lismore (302).
About 30 cells in a clump, irregularly disposed, enveloped in
mucus. Cells linear-elliptic, straight in side-view but from the
front arcuate, one side slightly concave, almost tiat, the othei*
distinctly convex, ends obtuse. "Membrane thin ; chloroplast
delicate, parietal, pale green, minutely granular, sometimes
divided in the middle, no pyrenoids.
Fam. PROTOCOCCACE^.
Genus C h a r a c i u m A.Br.
Characium guttula, n.sp. (Text-fig. 6rt).
Cellulsp corpore ovato levissime curvato, supra rotundato, subter
Text-fig. 6.
(a) Characi am </Hf til/a, n.sp.; {h, c) Oh. cer'assiforme var. minimum, ii.var. :
id) Ch. ornithocephalum A.Br.; (all x 1000).
rostrato in petiolum longum tenuissimum protracto; membrana
tenui; chloroplastide interdum pyrenoidibus singulis continente.
Cell. long. max. 25-40: corp. long. 12-16, lat. 6-8//.
Lismore (284, 286).
BY G. I. PLATFAIR. 529
Cells drop-shaped, rounded above, pointed below, and drawn
out into a slightly curved hyaline beak, which merges into a long
thin hyaline stalk as in Ch. lougipps: Membrane thin, chloro-
plasts sometimes containing a pyrenoid. Noted from two spots
at Lismore, singly, or several in a clump, on filamentous algie.
Characium cerassiforme v. minimum, n.var.
Cellular minutne, supra rotundatse nee deplanata?; petioloparvo
aut nullo; magnitudine formae typicfe triens.
Long. 1418, lat. 10/x. Lismore.
Cf. Ch. ceraHdformp. Eich. cfe Racib., Nowe gat. zielenic, p.l,
T. iii., f.l2; also Ch. pyrifurmp A. Braun, Alg. unicell., p.40,
T. v.B. About one-third the size of the type, and not flattened
above. Ch. cera-^si/nrme is more globose in tlie bod}^ of the cell
than Ch. lyyviformp. On Trihoripma; (Text-fig. G/), (■).
(•HARACIUM ORNITHOCEPHALUM A.Br. (Text fig. 6^).
Long. Corp. 23, lat. 74; petiol. long. l^jx. Lismore.
Fani. HYDROGASTRACE.E.
Genus U r x e l l a, gen.nov.
Character idem ac speeiei.
Urnella terrestris, sp.unica. (Text-fig. ?«-<:•).
Cellulse primum clavatse, deinde dehiscione anguste-urniformes;
lateribus subparallelis; ore everso; basi rotundata ; aut humo
sedentes aut tubulo insequali longo erectae. Membrana tenuis,
hyalina. Chloroplastis parieialis in extrema cellula contracta :
pj^-enoidibus plurimis.
Corp. urniform. long. 00-90, lat. 12-20; tubul. lojig. ad 150, lat.
7-8/ji. Auburn.
Noted in a green growth on a garden-path (clay) in verv wet
weather. The important part of the plant seems to be the long
clavate cell which, at dehiscence, becomes a slender urn-shape,
rounded below, with sides almost parallel, and the rim everted.
This cell is sometimes seated on the ground, sometimes erected
at the end of a fairly long irregular tubule of which it really
forms the infiated head. Membrane thin and hyaline, the chlo-
40
530
NKW AND KARE FRESHWATER ALG^.,
roplasts deep green, parietal, collected at the end of the cell, and
containing several pyrenoids. R^eproduction apparently by zoo-
gonidia. The plant bears a general resemblance to Protosiphon
Klebs, but it does not form a series of t/emmce, nor does the apical
Text-fig. 7.
(a) UrndJa terresh'i.^, gen.nov. et sp., (x500); {h, c) tubular form (x 880);
id) var. ahnormi.^, n.var., (x.500); (r) akinetes ?, ( x 500).
cell multiply by division, but seems to be entirely a gonidangium
There is also a great likeness to Codiolnm grey avium A.Br., but
the latter has a thick cell-wall and a laminated stipes — ^''stipitem
elongatum hyalinnm soHdum {gelatina indurata farchi/niy' — A.
Braun, Alg. unicell., p. 20, T. i.
BY G. T. PLAYPAIR. 531
Yar. ABNORMis, n.var. (Text-fig.To?).
Cellula tota tubifoniiis, loni^-e pi'otracta noc corporo nrniformi
ill extremo instructa.
Cell. lon^. e. 1.50-200, lat. 7-10//. Auburn.
In this form, found with the t\'pe, the cell, probably bj- excess
of I'ain, is drawn out into a simple irregular tube, dehiscing at
the end. There are none of the inflations in the tube which
evidence previous attempts at forming the gonidangium. Repro-
duction probably by zoogonidia, and perhaps also by akinetes;
(Text-fig. 7/').
Fani. OOCYSTACE.E.
Genus E r r m o s p ii je r a De By.
Eremosph.f.ra viRinis v. tetraedrica, n.var. (PI. Ivii., f.5).
Forma tetraedrica, lateribiis valde arcuatis, nodibus 4 levissime
angulatis.
Cell. diam. 110//. Lismore (308).
A curious form, in which the cell looks as if it were an inflated
tetraedron. TIk^ sides are well arched, and the angles only
slightly marked.
The hypnospore of E. viridis, which was described and figured
by me in Oorf/t^fis and JiJr/miosjjhcera, p. 11 6, f.G, I have lately
noted again in a fresh gathering (316) from Lismore. Outer cell
diam. 120, hypnospore 97/^. The scrobiculations are closer even
than figured, being only about their diameter apart, and the
scrobiculate membrane is backed, apparently, by the thick muci-
laginous lining often found in Er^'mosphcera. The contents were
green, but the chlorophyll seemed to be diffused in an even
parietal layer, no distinct chloroplasts being visible.
Genus O o c v s t i s Nag.
OocvsTis ROTULA Pla.yf. (PI. Ivii., f.fi, 7).
Ccenob. diam. oO, aiitosp. diam. 12/^. Lismore.
Cf. Chodat, Entwickl. d. Erem. virid., f. 13, 21 ; Playfair,
Oocystis and Evpmosjjh., p. 130, PI. vii., f.31. Coenobium spheri-
cal, thin-walled, containing 8 spherical antospores. The latter
are thick-walled, each with a central pvrenoid and cuneate chlo-
532
NEW AND RARE FRESHWATER \LG.V..
roplasts radiating out from it, six visible in optical section. It is
very difficult to see how they show at the surface, probably as
delicate discs. I had hardly finished congratulating myself on
this find, when a Chlamydomonas wdth similar cell-contents
turned up (from a different pool, however) and cast great sus-
picion on its validity as an Oocystis; (see Chi. rotula).
OocYSTis NODULOSA v. CRASSA, n.var. (Text-fig. 8).
Forma prse longitudine crassior,
subglobosa; membrana crassa; apici-
bus et interiore incrassatis et exteriore
tuberculatis ut in forma typica.
Cell. long. 35, lat. '2^-•2^\ cell,
matric. 84-7 4/a.
Lismore (308).
Cf. W. West, New Brit. Frw. Alg.
(J.R.M.S. 1894), Pl.ii., f. 31. Our
specimens are larger than the type,
and proportionately \ broader, which Text-fig. 8.
makes them, therefore, subglobose, Oocystk nodulosa var. crassa
not oval. W. West gives long. 25-26, "-^^^r., mother-cell with two
^ J T n T >~ J.1 J- • autospores; ( x 500).
lat. 1d-1//x as the dimensions. ^
Oocystis lacustris v. natans (Lemm.) Plaj-f. (PI. Ivii., f.8, 9).
Coenob. long. 57, lat. 38; cell. long. 23, lat 121/x.
Lismore (316).
Cf. Oocysth and Eremiosph.., p. 125, i.\ih. Our specimens are
the same diameter as the type, but a trifle longer; the chloroplast
also is more reticulate. The shape of the mother- cell is charac-
teristic of 0. lacustris, as also the arrangement of the autospores,
I.e., PI. vii., f.20.
Genus Franc ei a Lemm.
Franceia oblonga, n.sp. (PI. Ivii., f.lO).
Cellulse oblongse nee ovales; membrana crassa; setis tenuissimis
ubique vestitis.
Cell. long. 15, lat. 10; set, long. 15/x. Lismore (332).
BV G. 1. PLAYrAlR. 533
Cf. Vnuicput oralis (France) Lemni. in AMlle, Conj. u. Chloropli.,
p.59, £.31, wliicli works out at 25 x 15J/x. Our form, however,
is distinct!}' oblong. The contents are deep green, showing
signs of division into four parts.
Genus L A g e r h p: i m i a Chodal.
Lagerheimia elliptica, n.sp. (PI. Ivii., f.ll).
Cellulse exacte ellipticse, graciles, utroque polo setis longis sub-
apicalibus binis instructse. Cell. long. 12 J, lat. 6: set. long. 2()/x.
Sydney Water-supply (100).
The nearest form to this, I know of, is L. geneveiisis v. (jracilU
Playf., {L. ciliata v. (jracilis) Plankt. Sydney Water-supply, PI.
liii., f. 10-1 2. The latter, however, is narrowly cylindrical, while
this form is a perfect, slender ellipse.
Genus Bernardia Playf.
Bernardia tetraedrica, n.sp. (PI. Ivii,, f.l2).
Cellulcie sphfericse, spinis longis cavis gracillimis 4, tetraedrice
ordinatis instructfe; spinis basi valde inflatis, apice obtusis.
Diam, c. spin. 30; cor^^. 5//. Lismore (351).
This species consists of a single spherical cell with (apparently)
the membrane drawn out into four long very slender hollow
spines, arranged tetraedrically. The spines are blunt at the
extreme tip, not acutely pointed, and at the base are greatly in-
flated. Chloroplast delicate, pale green, parietal, confined to the
central part of the cell. Cf. Bernardia Chodati (Bernard) Playf.,
Austral. Frw. Phytopl., PI. lix., f.5, 6.
Genus Kirchneriella 8chm.
IviRCHNERIELLA OBESA W tt. G. S. West. (PI. Ivii., f 13-16).
Cell. lat. G-10, alt. 6-8, crass. 3i-5/x. Clyde, Duck Creek(26).
Cf. W. A: G. S. West, New Brit. Frw. Alg., p. 16. Syn.,
>S''lenasfrniu ohesum W. West, Alg. Eng. Lake Dist., p. 22, PI. x.,
f. 50-52; K. mibmlifaria G. S. West, Some Critical Green Alg{«,
p. 285, PI. 20, f. 20-30. Dimensions of the type, "diam. max. 6-9/x,
crass, cell. 3-4 •2/a"; of K. subsolitaria, "diam. max. 6-7/x, crass,
cell. 2 '9-3 -2/7-." I have found this form only once.
534 New and hare freshwater alg^
Genus T e t k a e d r o n Kiitz.
Tetraedron granulosum, 11. sp. (PI. Ivii., f.17, l>^).
Cellulpe plaiise, parvse, triangulares; lateiibus rectis in medio
levissiineconNexis; angulis acutis haud iiitiatis; membrana grosse
g]-aiiulata, granulis in quincuncein ordinatis. A latere visa",
mudice cuinpressie, elliptico-lanceolata^, angulis acuminatis.
Cell, diain. 21, crass. 11/x. Lismore (302):
The specimens noted were Hat, not tetrahedral, triangular with
straight sides shghtly convex in the centre; angles acute, not
inflated; membrane coarsely granulate in (|uiiicunx. From the
side, elliptic-lanceolate with pointed ends. In the centre of the
cell might be discerned a minute nucleus enclosed in a pale brown
globule (?pyrenoid). ►Sometimes the nucleus was entirely hidden
Ijy the globule, which then had the appearance of a large brown
pyrenoid; cf. Oocyst is and Bremosj^hcera, p. 126, remarks on 0.
lacustris \. pahtdeiisis; and p. 137 on 0. oirdis y. cf/Zindracra,
where the same phenomenon occurs.
Tetraedron (iUADRATUMf.3iiN0R Keinsch. (PI. Ivii., f.lD).
Cellula plana; diau]. sine spiiiis 17/x. JJsmore.
Forma excavata, n.f. (PI. Ivii., f.2()).
Forma plana, lateribus alternantibus conca^ is, alteris levissime
convexis; angulis spinis brevibus acutis singulis armatis.
Cell, diani. 19, alt. Ib/i, s.sp. Lismore. Cum priori.
Cf. Pveinsch, Monog. Folyedrimn, p.500, T. iv., fig.7r/. This is
a variant of f. inlnor with alternate sides conca\ e, the other sides
generally somewhat convex. ,
Tetraedkon caudatum v. australe, n.var. (PI. Ivii., f.21, 22).
Celluky inaiqualiter pentaedricic; angulis acutis, spinulis min-
utis singulis armatis; lateribus coiicavis; utrinque infra marginem
angulis mainmillatis armatis singulis instructa;. A latere, plaiue
(ichnographia universa respecta), lanceolato-elliptica^ apicibus
acutis armatis; altero latere convexai, altero autem angulis main-
millatis armatis binis projicientibus, instructjv.
Cell. lat. max. c. spinis 21/x. Lismore. Cum priori.
BY G. I. PLAYFAIR. 535
Cf. Polyedrium pentagonum Reinsch, Algenfl. v. Frank., T.iii.,
fig. ii.c( = Tetr. caudatum Corda) which is exactly the same shape
but without the two niamniillate angles which jut out from one
side.
Tetraedron regulare v. octaedricum (Rein.) mihi.
Cellulse angulis senis octonisve praeditae. (PI. Ivii., f.23).
Cell. diam. 1 7/x. Lismore.
Cf. Polyedrium octaedricum v. spinosum Reinsch, Algenfl. v.
Frank., p. 78, T. v., fig. v., 1867. In this place, Reinsch has
united two distinct types under onename"^ {P. octaedricum Rein.,
Monog. Polyedr., p. 507, 1888). The first of these. T. v., fig. iv.,
must retain the specific name, while the other, T. v., fig. v., is
evidently a form of I'etr. regulare Kiitz. ( = P, tetra'edricum Nag.),
with from six to eight angles instead of four. Our specimens
are the same shape as Reinsch's fig. v. 6, but very much smaller;
he gives lat. 38-47/x.
Tetraedrox hastatum v. elegaxs Playf.
Cell. diam. c. proc. 30/x. Noted lately at Lismore (362), only
known previously from Parramatta. Cf. Austral. Frw. Phytopl.,
p.845, PI. Iviii., f.27.
Tetraedron acutum v. rectilineare Playf.
Cell. diam. c. spin. 34-40/x, sp. long. 1 0/x. Confirmed from
Lismore (362), only recorded previously from Enoggera; ibidem,
p.8-45, PI. Iviii., f.26.
Tetraedron conicum, n.sp. (}'l. Ivii., f.24).
Celkdjc tetraedricai; angulis conicis vix inflatis; apicibus mu-
ticis, obtuse-rotundatis; lateribus levissime conciivis.
Cell. diam. 19-25//,. Lismore (362).
The cells are tetrahedral, composed of four conical* angles
meeting in the centre. The angles can hardly be called inflated,
the sides of the cones being almost straight. The apices are
bluntly rounded, without point or spine.
" Polyedrium acuminatum spino8iun at the bottom of Plate v., I.e., is
either a slip of tlie pen or a priiiter'.s error.
536 NKW ANr) RAKR FRESHWATER ALC.X,.
Tetraedron minimum V. RECTiLixEARE Plavf. (PI. Ivii., f.25).
Cell. long. 11 J, lat. lOJ/x. Botanic Gardens, Sydney (3).
Cf. Australian Freshwater Phytoplankton, these Proceedings,
1917, p. 842, PI. 58, f.l6, 17. Known to me previously only from
Guildford. A rare form.
Tetraedron obtusum, n.sp. (PL Ivii., £.26).
Cellulas tetraedricai: lateribus rectis; angulis rostratis; rostris
crassis quadratis, brevibus; apicibus truncatis.
Cell. diam. 21/x. Lismore (362).
Cell tetrahedral, sides straight, angles produced into a siiuV^
nosed rostrum, the latter broad, quadrate, truncate.
Tetraedron trigonum Nag., forma (PI. Ivii., f.27, 28;.
Forma lateribus in medio quam levissime retusis; spinis bre-
vissimis oblique recurvis; membrana crassa.
Cell. diam. 36, crass. 20/x. Lismore (345).
Svn., Polycdrium tetragonum f3 piinctatum (Kirchn.) Lag.,
Bidrag till Sveriges Algflora, p. 62, T. i., f.28. P. trigonum e
pu7ictatum Kirchii., Alg. v. Schles., p. 104 (sec. Lagerheim, I.e.).
The type has sides simply arched, with or without a small
straight spine at the angles. In this form, the sides are slightly
flattened or retuse in the middle, and the spines are obliquely
recurved. Our specimen was not punctate. For the type, cf.
Nageli, Gatt. einz Alg., T. iv.B, figs-ln- and 3«, b only.
Fam. HYDRODICTYACE/E.
Genus P e d i a s t r u m Meyen.
Pediastrum simplkx Meyen. (PI. Ivii., f.29, 30).
Ccenob. (cell. 4) c. proc. long. 35-50, lat. 33-48; s. proc. long.
17-24, lat. 16-22; cell. long. 24-34, lat. 12-17/x. Ca?nob. (cell. 8)
diam. 50-64; cell. long. 18-24, lat. 8-1 2/x.
Parramatta Park (99, 136).
Meyen, Beob. ub einig. nied. Algenf.,( 1829) T.xHii., f.l. 8yn.,
P. simplex var. radians Lemra., Zeitschrift f. Fischerei, 1897; P.
s im])le.i: fS aiinulatum Choda^t, Alg. vertes, p.225, 1902; P. simplex
V. clathratum f. radians (Lemm.) G. J-^. West, Third Tanganyika
Exp., p.l34, 1907.
BY G. 1. PLATFAIR.
537
Meyeii gives five fig\ii'es in illustration of his P. simplex, and
these have since been arranged as three difterent variations.
His fig.l is var. radiaiiH Lemni.(1897); figs. 2, 5 are f. orata (Ehr.)
llacib., {Asterodiction ovatu/ni Ehr., 1845); while figs. 3, \ arc
f. dathrata Schroter (1883). One of his figures, liowever, must
be left to represent the type, and it falls to fig.l to do this, var.
radians Lennn., being the last in order of priority. It seems to
have been overlooked also, that Meyen himself arranged it so
in the text. He says {I.e., p. 4) "c^ie unter Fiy.l mochte als die
Norm zu betrachten seyn.'' The 8-celled ccenobium without
central cells is, therefore, the type of P. simplex, and, incident-
ally also, of the genus Pediastrum.
Forma clathrata (Schroter) Rac. (Text-fig.9).
Cienob. diam. 60-92; cell, periph. long. 26-28, lat. 12//.
Parramatta Park (99, 136).
Ptdiastrum ximjilc
Text-fig. 9.
f. cloJhrala (Schroter) Rue.
( - ()()()).
Schroter, in Jahresbericht d. Schles. Gles. fiir vaterland.
Cultur, 1883, p. 182; llaciborski, Gatunki rodz. Pediastrum, 1889,
p. 8. Syn., P. enoplou W. ifcG. S. West, Frw. Alg. Madag., 1895,
p. 81, T. 5, f.1,2; P. dathratum (Schrot.) Lemm., For.<ch. Biol.
538
NEW AND RARE FRESHWATER ALGM,
Stat. Plon, vii., 1899, p. 20. A name that would take priority
of 8chroter"s is P. duodenarium (Bail.) Rabh., Fl. Eur. Alg., iii..
p.71, 1864, but, from Raciborski's notes, it would appear to in-
clude several different forms, I.e., pp.9, 10.
Forma ovata (Ehr.) Rac. (PI Ivii., f.31).
CcEnob. (cell 4) diam. 53; cell, long.- 25, lat. 16/y-.
Parramatta Park (136).
Raciborski, Gatunki rodz. Pediastrum, 1 889, p 8. Syn., As-
terodiction ovatum Ehr., Ber. iib. d. Verb. d. Akad , 1845, p. 71:
P. ovatum (Ehr.) A.Br., Alg. unicell., 1855, p. 81; P. Sturmii
Reinsch, AlgenH. v. Frank , p. 90, T. vii., f.l; P. Schroteri Lemm.,
Forsch. Biol. 8tat. Plon, vii., 1899, p. 20, T.2, f.33; /'. dathratum
f. ovatum (Ehr.) 8chm., in Eng. Bot. Jahrb., xxxii., 1902, p.84
T.3, f.l8.
All forms of P. simplex are very rare here; I have them from
one spot only.
Pediastrum duplex v. rkticulatuiM Lag. (Text-tig. 10).
Coenob. (cell. 4) diam. 52; cell. long. 16, lat. 14/x. Ccenob. (cell,
permult.) long. 90-130, lat. 90-120: cell. long. 26-30, lat. 15-23/x.
Sydne\' AVater-supply ( 80,
81, 90, 100, 115); Botany
(50).
Lagerheim, Stockholms
Pedi., Protococc, o. Palm.,
p.56, T. ii., f. 1. Syn., /'.
dnjylex v. clathratuin Schro-
ter, SchwebeH. uns. Seen, p.
37, f.86. This form is com-
mon enough in the Sydney
Water-supply, but rare else-
where here. In the large
coenobium figured, the two
dead cells on the left show
that the early stage of the
plant was more akin to P.
JJoryaituni.
Text -fig. 1(1.
Ptdia-strnm dt(p/e,v var. r(JicuJafii/ii
Lagerh. ; ( x 500).
BY G. I. PLAYFAIR. 539
Pediastrum Boryanum (Tuip.) Menegh. (Pl.lvii., f.32).
Forma; coenobium e cellulis 4 exstructuiii. Ccenob. diam. 40/x.
Botany (15).
Var. Haynaldii (Istv.) mihi. (PI. Ivii., f.33).
Coenobium long. 180, lat. 155; cell. diam. 32//.
Botanic Gardens (3).
P. n<iynaldi% Istvanffi, Jel. Magyar. Alg., 1888, p. 252, T. ii.,
f. 38. Syn., P. angidosum v. araneosum Rac, Gatunki rodz.
Pediastrum, 1889, p. 18, T. ii., £40. This form is characteri.sed
by the fine ridging on the cell- wall. There is no difference
between Racil)orski's form and that of Istvantii, save that in the
latter the lines on the cell are granulate.
Var. AUSTRALE, n.var. (Pl.lvii., f.31).
Cellula3 perimetri cornibus pleiumque brevibus: incisuris altis,
oblongis, intus rotundatis, lateribus parallelis.
Ccenob. long. 116-150, lat. 1 00-127; diam. cell. 12-20//.
Botanic Gardens (3).
A form of P. Bori/anum common and widespread here, but
apparently not so elsewhere. It is characterised bv the shape
of the marginal incisions, which are in the form of the letter U,
oblong with rounded ends and parallel sides. Tlie horns are
generally short. The margin between the horns of the two
adjacent cells is very slightly concave.
Pediastrum biradiatum var. Braunii (VVarim.) (Jhud.
Cell. long. 24, lat. 12/x. Kyogle(219). (PL Ivii., f.35).
C/'. Chodat, Alg. vertes, p. 230. >Syn., P. Praionii' Wiirlnrdiun,
8chw. Crypt. No. 32."^ P. rotula Ehr., {no7i Nageli) in A. Braun,
Alg. unicell., T. vi., figs. 2, 5, 6. The dimensions of the peripheral
cells of Braun's fig. 5 work out at long. 22, lat. 13/x. P. biradi-
atum Meyen, is extremely rare; 1 have never come across the
type, and only one specimen of the variation. The lobules of
the cells are bifid in this form, only incised in the type.
^ Not having seen Waitniann's t3'pe, I .siniplj^ follow Chodat in the
.synonymy. Latter] lei in, however, Veg. siissw. plankt. Biiren-Insel, p. 10,
and Borge, Algentl. v. Schweden, p. 00, identify' it with P. trirormifum
Boriie.
540
NEW AND RARE FRESHWATER ALG.?:,
Pediastrum tetras var. Crux MiCHiELi (Rein.) Playf.
Cell. diam. 18//. Botany (109). (PL Ivii., f.36).
Noted also in the Sydney Water-supply, these Proceedings,
1913, p. 518, PI. 56, f.6. In the specimen here figured, the cells
were closely appressed, and the slits were observable through
w^iich the zoogonidia had escaped.
Fam. CCELASTRACE.^.
Genus Tetrastrum Chodat.
Tetrastrum elegans V, DENTATUM Playf.
Coenob. c. spin. diam. 32: corp. 8; sp. long. 12; dent. long. 2-3/z.
Lismore (258).
CJ\ Austral. Frw. Phytopl., p. 833, Pi. Ivii., f. 7. Hitherto
known only from Parramatta: this form is now confirmed from
Lismore.
Genus Ccela strum Nag.
Ccelastrum reticulatum (Dang.) Senn. (Text-fig. 1 1 ).
Cellulae matric. diam. 14/x; cell,
coenob filial. 6/x.
Parramatta Park ( 1 36).
Syn., Hariotina reticulata Dan-
geard, Mem. s. 1. algues, ii., 1889.
Generally found in plankton-gath-
erings as broken coenobia. Tn this
case, six cells were noted, partially
united by their processes, each
with a perfectly formed autocolony
Text-fia.ll.* within.
EXPLANATION OF PLATES LIV.-LVIll.
Plate liv.
Y'\g.\.~(Mdllaloria^ticxa, n.sp.: ( x (iHO).
Fig.'i. — O. simplicissimn v. 'irmntldfa, n.var. : ( x 100<3).
Fig.3. — O. simplicisftima v. roiislrirhi. n.var.; ( x HHO).
Fig. 4. — O. princeps\-A\\>Aw\-\ ])(>rli(jii of an empty filament, shoAving the
thick sheath and permanent dissepiment.s; ( k 880).
Fig. 5. — FhorinidiHin t/rande, n.sp.; ( x500).
* Coelastrum reticidatam {Daxig.) Senn, three isolated cells of a ctjeno-
bium, each containing an autocolonj^; ( x 1000).
BY G. T. PLAYFAIR. 541
Figs 6, 7.— Ph. immarium, n.sp.; (0) x 1330, (7) x 2000.
Figs. 8, 9. — Auabcena auMralica, n.sp.,( x 1000); (8) cells and heterocyst of
infertile trichome; (9) pai't of a fertile triclioiiie; the r^onstrictions
of tlie cells can be discerned only with the /.j in. obj.
Fig. 10. — A. aufifraJica var. conffricta, n.var. ; ( x 1330).
Figs. 1 J -13. — A. rariahih's xar. cyiindrncea, n.var. (x 1000); ill) cells and
heterocj'^st; (12) immature catenate gonidia; (13) matuie spore.
Fig. 14. — A. oarlUarioides var. cyas-m, n.var.; ( x 6f)0).
Figs. 15. 16. — A. /a.t-a var. hoiieih^i>i, n.var., ( x 1000); (!,')) cells and hetero-
cyst: (16) cells and spore.
Figs. 17-21. — Cyllndrospennnm sta>/iui/e var. nn-^fra/e, n.var.; (17, IS) with
conical heterocysts( x 1000); (19. 20) with ol)long ]ieterocysts( x 660);
(21) coiled trichome.
Figs. 22, 23. — ('. rerfani/ii/rtrf Plaj'f. ; (22) tricliome with thin-walled iiinna-
ture spore ( x 10(M>); (23) with mature thick- walled spore ( x 660).
Figs.24, 2o. — C. i-ectaiKjnlare var. parvicellHla,n.xM\ ( x 1000); (25) trichome
with apical cell.
Figs.26, 27.— C. GoeJzei Schm., forma: ( x KM)).
Plate Iv.
Figs. 1, 2. — Carteria ruynlosa, n.sp.; (2) endview; ( x 1000).
Fig. 3. — C. ruf/ulosa var. angidata, n.var.; ( x 1000).
Fig 4. — ('. ragidosa var. inrt^qualis, n.var.; ( x 1(X)0).
Figs.5, 6.-6'. mulfifdis (Fresen.) Dill, formed; ( x 1330).
Figs. 7, 8. — C. granulosa, n.sp.; (8) sideview; ( x 660).
Figs. 9, 10. — Chlamydomonas reniformis, n.sp.; (10) sideview; ( x 1000).
Fig. 11. — Chi. Lismorensis var. gracilis, n.var.; ( x2000).
Fig. 12. -Chi. rotida, n.sp.; ( x 1000).
Fig.l3.— C7</. alpina Wille; ( x 2000).
Y\g.U.—ChI. caudata Wille; ( x 1000).
Figs. 15-17.-67;/. timcidata, n..sp.; (15) x 1330; (16) with stratified cell-wall
( xt)60); (17) mother-cell ( x 660), autospores ( x 1000); for conveni-
ence, the ccpnobium is drawn smaller, and the autospores therefore
appear more crowded than natural; the latter are in optical section,
the spotted surface of the chloroplast being omitted; the tiagella
were just as figured.
Fig. 18. — Chi. macidata var. ohlonga, n.var.; ( x 1330),
Fig. 19. — Chi. macniata var. planktoaicu, n.var.; ( x 1000).
Fig.20.— CA/. mefastigma Stein; ( x 1000).
Figs,21, 22.-67?/. paupercnla, n.sp.: (21) x 2(100: (22) x 1800.
Fig. 23.— 6'/i/. pvsHIa, n.sp.; ( x 660).
Fig.24. — Chi. angidosa var. ohesa, n.var.; ( x 13.30).
Fig.25. — Chi. pisiformis \nr. orellafa, n.var.; mother-cell and autcxspores;
(x 1000).
542 NEW AND RARE FRESHWATER ALfiiE,
Fij^.'iG. — CJifo/'Oj/oniitm minimnm, n.sp. ; ( x 1330).
ritf;s. 27, "28. — Pfei'omona.-< am/ulom var. auMraJis Playf.. foniia; ( x 1000).
ri,i^.29. — /-*/. anfjiilo>ia var. ■■<cn1ifornu>i. n.var. ; ( x 1330).
Fi,u'..3(». — Pt. (iiKjulo.sn var. i-f.vi1Hforini-<, n.var.: ( x 1330).
Plate Ivi.
Fit's. 1-3.— .S'?///»/v/ ;/raini/o-^<i Playf.: (1) elongate Cienobium witli zooids
attaclied to a central mucous peduncle ( x 060) ;. (2, 3) individual
zooids showing one or moie dark red granules near the anterior end
( X 1000).
Fig. 4. —TexseJIarla ro/rocina Pla3'f., supposed free zooid; { x loOO).
Figs. 5, 6. — Xanfhod/^iciis Luuferbadii Scliew., (so-called); (o) face-view of
a preserved specimen with shrunken contents showing central
nucleus, but no chlamydomonadine arrangement of the chromato-
phore; (G) sideview; ( x 1000).
Figs. 7. H. — Phacotn.^ rra.^-sii>i, n.sp., ( x 1000); (S) sidevicAv.
Figs.9, \(). — fVi. (ilaher, n..sp., ( x 1000); (10) sideview.
Figs. 11, Vl.~Pk. cj/aher, formc-e; ( x 1000).
Figs. 13, 14.— PA. anstralk, n.sp., ( x 100(»); (14) sideview.
Fig. 15, — Spondi/Iomorum qnaternarinm Rhr. : ( x 1000).
Figs. 10, 17 — Sp. quale rnarhim var. roslrahiin,n.ya.\\,[ x 1000); (10) bunched
ecenobium; (17) rosette-form.
Fig. 18. — Vol I'll/ i na Steinii Pla,yi., single cell of a crcnobium, in sideview,
showing nucleus, contractile vacuoles, and stigma in the usual
chlamj'domonadine positions.
Figs. 19, 20. — Volrox aureus var. hemiyj/uericn.^, n.var. ; (19) cells from
above ( x 660); (20) three cells in sideview ( x 1000).
Figs.21, 22 — Volrox Carte.ri Stein; (21) immature zygote as described by
Carter ( x 660); (22) mature zygote ( x oOO).
Plate Ivii.
Fig.l. — EJakatothrix (lehitinom Wille, isolated cell; ( x 1330).
Fig.2. — E. gekitiiiosa, mucous matrix with newly divided cells; ( x 1000).
Figs. 3, 4. — Dactyl othecearciiatiLm,\\.^\)., two cells, of many noted, involved
in a mucous matrix ( x 1330); (3) face-view; (4) sideview.
Fig. 5. — E7'emosphc(ira viridis yslv. tetraedrica, n.var.; ( x 270).
Figj^. — Oocystis rotnla Playf., mother-cell with autospores; ( x66<)).
Fig. 7- — 0. rotnla, autospore; ( x 1100).
Fig.8. — 0. /acmtris var. nafaus (Lemm.) Playf , mother-cell with auto-
spores; ( X 660).
jritr.9. — 0, (acnstris var. natans, autospore, showing fragmented, reticu-
late chloroplast; ( x 1000).
Pig. 10. — Franceia ohhnga, n.^i).; the settv are, for convenience, figured
much shorter than natural ; they should be as long as the cell ;
( X 1330).
BY fi. I. PLAY FAIR. 543
Fig. 11. — Lafjerheimia eUiptica, n.sp. ; ( x 1330).
Fig 12. — Bernardia fetraiidrka, n.sp.; { XlOOO).
Figs.l3-1().— A'//r/<7K'?vV7/rt o^ft.^Y< W. & G. S, West; (xlOOO.
Figs. 17, IS. — Titrai'drou ;/raiii(/o.siun, n.sp.: '17) face-view; ( IS) sidevieM'
( X 1000).
Fig. 19. — Tetr. qiiadrdlnin f. minor Keiiiseli: ( X SOO).
Fig.2(>. — T^//-. qnadra/nm f. <jrara/(i, n.f. ; { X SOO).
Figs. 21, 22. — Tifr. r((iidafv/N xar. a it.-<l rah . ii.var.: (21) face-view; (22) side;
( X 1000).
Fig.23. — Tefv. rej/n/are var. ortal-dricnrn (Keinscli) niilii; (x 1000).
Fig.24.— 7'(>//-. conirum, n.sp.; ( X 1000).
Fig.2o. — Tetr. minimum var. rectiliiieare Playf.: (x lOOO),
Fig.26.— Te/r. oUnmm, n.sp.; ( x 1000).
Figs. 27, 2S. — T(tr. friyonnm Niig., forma; (27) face- view; (2S) side; ( XfifiO),
Figs. 29, 30. — Pediafffrnm simp/ex Meyen, 4-eelled forms; ( Xf)60).
Fig.31,— f*ef??. mn2)Iex f. orata (Ehr.) Pvacib.; (x660).
Fig..S2. — Pedi. Boryanum (Turp. ) Menegh., 4-celIed form; (X.3(K)).
Fig. 33. — Pedi. Dorijannm var. llayiaddii (Istv. ) milii: ( X ilOO).
Fig.34. — Pedi. Boryanum var. anMrale., n.var. ; (x(IHO).
Fig.3r>. — Pfdi. hiradiafii/ii var. /jVavr^r/ (Wartm. ) Cliodat ; (X(]r.O).
Fig.36.— P^r//. fefnts var. C'riiu- Mirluwli (Reinsch) I'layf.: ( X l(>Ol>).
Plate Iviii.
Spiroijyra nef//ecfa (Hass. ) Kiitz.
Fig. 1. — Conjugated filaments with elliptic /.j'gospores and non-inHated
sporangia! cells; (X220).
Fig.2. — Conjugated tilaments witii oval zygospores and inflated sporangial
cells; ( X 220). Figs. 1 and 2 formed part of one and the same
scalariform filament.
Figs.3-6. — Four types of zygospore in the one gathering (X 400) ; fig. .")
(xr)00).
Figs. 7, 8. — Unaltered cells of a conjugated filament, with simple delicate
chloroplast; (8) x 330; (7) chloroplast much enlarged.
Figs. 9, 10. — Long cell of a vegetative filament, with delicate, fimbriate
chloroplast and minute pyrenoids; (9) chloroplast much enlarged;
(10;x330; the fringed edge of the chloroplast is tof) delicate to be
seen under this low magnification.
Figs. 11-13, — Cells with more robust, scalloped chloroplast; (11) chloroplast
much enlarged; (12) short infiated cells probably about to conjugate
(X 330); (13) long uninflated cell of vegetative filament (x330).
V\^sA4:-\6.- -Spirofjyra ne(j/ecfa var. amylacea, n.var., vegetative filaments
and chloroplast; (14) long cell ( x330,'; (1.)) short cells ( x330); (16)
chloroplast enlarged Conjugating cells and zygospores of this
variation as in fig. 1,
544
THE MOSSES OF NORTH QUEENSLAND.
By Dr. Y. F. Brotherus and the Rev. AY. Walter Watts.
Foreword.
(By the Rev. W. W. Watts.)
The following paragraph is reprinted from my paper, " Some
Notes on the Ferns of North Queensland" (These Proceedings,
1914, Yol. xxxix., Part 4, p.7o6). " In the winter of 1913 (July-
August), I spent a month in the Cairns district in search of
Ferns and Mosses. I collected in the rich scrub-lands between
Bartle Frere and the Russell River, more especially in the
neighbourhood of Josephine Creek; around Babinda also, and
up Frenchman's Creek to beyond the Second Falls. - Babinda
and Frenchman's Creek lie at the base of Bellenden Ker, along
the coastal railway. Following, later, the main line, I visited
Stoney Creek, Kuranda, Atherton, Malanda, Yungaburra, Lake
Barrine, and, last and best of all, Ravenshoe and the Tully
Falls."
The mosses of North-east Queensland being essentially Malay-
sian, rather than Australian, in their affinities, specimens from
all my packets were submitted to the distinguished expert. Dr.
Y F. Brotherus. Despite the war, with its submarine dangers,
every parcel reached Helsingfors in safety, and, by the end of
1917, full reports, in return, came to hand. The number of new
species is smaller than I anticipated, owing, in part, to the zeal
of the late Government Botanist of Queensland, Mr. F. Manson
Bailey, C.M.G., F.L.S., but mainly to the aforesaid Malaysian
affinities of the flora. The chief feature of the following pages
is the large number of new records. No less than seventeen
genera new to Australia are listed, and some thirty known
species. One new genus, Pterohryidinni, is described, and there
are fourteen new species. In addition, many of the species
recorded are new for tropical Queensland, though not new for
Australia. The new genus was first found by me, in 1913, at
BY V. F. BROTHERUS AND W. W. WATTS. 545
Raveiishoe, North Queensland; but, curiously enough, was col-
lected, in 1916, by Mr. J. L. Boorraan, of the Sydney Botanic
Gardens, on Mt. Lindsay on the border between Queensland and
New South Wales.
It is interesting to note that a few species, the occurrence of
which in Australia was regarded as doubtful by Watts and
Whitelegge (Census Muscorum Australiensium) are now definitely
recorded, such as Leucobryurn sanctum^ Arthronormus Schimperi,
Leucophanes octoble'pharoxdes^ and CalymiJerHH molucceiise.
It remains, on behalf of Australian botanists, to express sincere
thanks to Dr. Brotherus for his continued and generous assist
ance; and also to thank the Director of the Sydney Botanic
Gardens for the facilities afforded me at the National Herbarium
for working up my material. — W.W.W.
Note.— The sign * before a generic or specific name denotes
that it is new for Australia; and the sign t that it is new for
tropical Queensland.
A. ACROCARPI,
Dicranaceae.
Trematodon Michx.
fT. LONGESCENS CM.
Barron River, n.386; Frenchman's Creek, n. 419.
[Note. —Trematodon Baileyi Broth., belongs also to the Cairns
district (Mulgrave River), but appears not to have been collected
since Bailey found it in 1889.]
Pleuridium Brid.
Pleurid. sp. ster. incerta.
Track down to Red lynch Railway Station, near Cairns. No
Pleuridium previously recorded for Queensland.
*G A R c K e A CM.
*G. PHASCOiDES (Hook.) CM.
Babinda, n. 326 e.p. Previous record, "Southern Asia."
D I T R I c H u M Timm.
D. AFFINE CM.
Ravenshoe, n. 457.
41
546 THE MOSSES OF XORTH QUEEXSLAND_,
DiCRANKLLA Schillip.
D. DiETRiCHiiE (CM.) Jaeu.
Ravenshoe, n. 447-8.
I). PYCNOGLOSSA (Brotli.) Par.
Stoney Creek: Babinda; Malanda. F\uranda: Mrs. Brotherton.
First found by Mr. F. M. Bailey on the Mulgrave River in 1 889.
Var. LONGIFOLIA Broth, et Watts, var.nov.
Street's Gnllyj Kuranda, n. 265.
H o Lo M I T Ri u M Brid.
H. MuELLEKi Hampe.
Ravenshoe, n. 573 e.p. Previous records, Rockingham Bay:
Dallachy, 1868; Bellenden Ker : S.Johnson, 1891.
H. PERiCHiETiALE (Hook.) Brid.
Ravenshoe and Malanda (several nos.).
DiCRANOLOMA Ren .
D. DiCARPUM (Hsch.) Par.
Ravenshoe, several nos.
D. SERRATUM (Bfoth.) Par.
Malanda, n. 494, 615 e.p. (Previous Queensland record, Mt.
Mistake : Bailey, 1885).
D. Wattsii Broth., n.sp.
Dioicum; ^lauco-viride, nitidiusculum; can/is usque ad 5 cm.
longus, adscendens, inferne denudatus, dein dense foliosus, sim-
plex vel parce ramosus; folia horride patula, stricta, late lanceo-
lata, breviter acuminata, 4-4 5 mm, longa et 2-2'5 mm. lata, mar-
ginibus erectis, summo apice conniventibus ibidemque argute
serratis, ubique hyalino-limbata, limbo latissimo, apicem versus
sen.sim angustiore, nervo tenui, continuo, dorso superne argute
serrato, cellulis chlorophyllosis, elongate et anguste linearibus,
apice brevioribus, alaribus numerosis, quadratis, fusco-aureis,
ssepe decoloratis: bractese perich^tii interna' e basi longe \agi-
nante convoluta subito in pilum longiusculum contractse; setae
binre ex eodem perichsetio, tenues, vix ultra 5 mm. alta3, fusci-
dulfe: theca oblonga, obliqua, subcernua, collo strumuloso; oper-
culum e basi conica longe et oblique subulatum.
BY V. F. BROTHERUS AXD W. W. WATTS. 547
Ravenshoe, n. 534rt (type).
Species D. austro-scopario (C. Muell.) affinis, sed foliis latiori-
bus, brevius acuminatis, setis binis ex eodem perichiPtio, brevi-
oribus et tenuioril)ns dignoscenda.
[Note. — Z>. aiistro-scoparinrn (CM.) was collected by Bailey,
in 1889, on tbe summit of Bellenden Ker.]
Leucoloma Brid.
L. 8ip:berianu>[ (Hornscb.) Par.
Ravenshoe (several nos.).
L. SUBINTKGRUM Brotb.
Babinda: Frenchman's Creek: Malanda: Ravenshoe (frequent).
[Originally found at Palm Camp, Bellenden Ker, by Bailey
(1889)J.
[Note. — L. clarvuerve CM.., is recorded for "Tropical Queens-
land " in Bailey's Suppl. to his Synopsis, 1888.]
C A M p Y L o p u s Brid.
C. {Rigidi) Wattsii Broth., n.sp.
Dioicus; robustiusculus, csespitosus, csespitibus densis, viridis-
simis, subopacis; caulis adscendens, usque ad 3 cm. longus, fere
ad apicem rubro-tomentosus, simplex vel furcatus; folia sicca
imbricata, humida patentia, canaliculato-concava, lanceolato-
subulata, c. 6 mm. longa et c. 08 mm. lata, superne margine et
dorso serrata, ner\T3 basi dimidiam partem folii latitudinis occu-
pante, dorso lamelloso, lamina usque ad apicem distincta, cellulis
minutis, rhombeis, baud incrassatis, basilaribus in seriebus
pluribus quadratis, alaribus teneris, fuscis, auriculas distinctissi-
mas efformantibus. Csetera ignota.
Cairns district. Frenchman's Creek, Second Falls, n. 360 ;
Ptavenshoe, on rocks al)Ove Falls, near Major's, n. 484 e.p. :
Watts, 1913.
fC. WooLLSii (CM.) Par.
Ravenshoe, n. 452, 459, 532.
EUCAMPTODON Mont.
fE. MuELLKHi Hpe. et CM.
Ravenshoe and Malanda,
548 THE MOSSES OF NORTH QUEENSLAND^
Leucobryacese.
Leucobryum Hampe.
fL. BALLiNENSE Bi'oth., A.M., vi., 1916, n.l99.
Ravenshoe, ii.534f, cZ; Malanda, ii.550 e.p. Ballina, N.S.W.,
the only previous record.
L. BRACHYPHYLLUM Hpe.
Babinda; Frenchman's Creek; Ravenshoe.
L. CANDiDUM Brid.
Frenchman's Creek, n. 397 e.p.; near Babinda, n. 445 e.p. Ap-
parently the first record for N.Q.
Ij. sanctum (Brid.) Hampe.
Kuranda ; Bartle Frere ; Babinda ; Frenchman's Creek :
Malanda. Widespread "from Nepal to Malesia" (teste Broth.).
fL. STRiCTiFOLiUM Broth.
Ravenshoe, n. 469, 672. Previously limited to N. S. Wales
and S. Queensland.
[L. vesiculosiim CM., was collected, at Whelanian Pools, Palm
Camp and Harvey's Camp, Bellenden Ker, by F. M. Bailey, in
1889; and at Trinity Bay, by Mrs. (or Miss) Cribble, in 1894.
L. viride Mitt., is recorded, in Herb. Melb., for Daintree River,
Q., but information regarding this sp. is needed. L. Wattsii
Broth., was collected by Dallachy at Rockingham Bay, Q., but
named L. sanctum var. in Hb. Melb.]
Leucophanes Brid.
L. AUSTRALE Broth.
Kuranda, n.435. Previously recorded for Freshwater Creek,
near Cairns, and "Ker Range" (teste CM.).
L. OCTOBLEPHAROIDES Brid.
Kuranda; Bartle Frere; Frenchman's Creek; Babinda; Malanda.
Previously recorded, in Hb. Melb., for Johnstone River (Berth-
aud), and Daintree River (Pentzske). Distrib. "from Nepal to
Pacific Islands."
OCTOBLEPHA HUM Hcdw.
O. ALBiDUM (L.) Hedw.
Cairns; Kuranda; Malanda; Ravenshoe.
BY V. F. BROTHERUS AND W. W. WATTS. 549
Arthrocormus Doz. et Molk.
A. ScHiMPERi D. et M.
Babinda, n. 422, 4 38. In the Melb. Herb., there is a doubtful
record of ''■Leucobryuiri' Schimperi for the Johnstone River. A.
Schimperi is found in the Indian Archipelago, the New Hebrides,
etc.
*ExoDiCTYON Card.
*E. SUBSCABRUM (Broth.) Card.
Frenchman's Creek, n. 3946. Previous record, Papua.
Fissidentaceae.
F I s s I D E N s Hedw.
IF. ARBOi^Eus Broth.
Stoney Creek; Babinda; Lake Barrine; Raven shoe.
fF. ASPLKNioiDEs (Sw.) Hedw.
Ravenshoe, n. 471, 533 e.p.
F. {Semilimhidium) cairnensis Broth, et Watts, n.sp.
Dioicus(?): gregarie crescens, pallide viridis, opacus; caulis
usque ad 5 mm. longus, cum foliis usque ad 1 9 mm. latus, inHma
basi fusco-radiculosus, dense foliosus, simplex vel furcatus; folia
sicca hoiiiomallula, humidastricta, erecto-patentia, infimaminuta,
dein subaequalia, ligulata, breviter acuminata, acuta, c. 1*3 mm
longa et c. 0"38 mm. lata, marginibus supeine minutissime serru
latis, lamina vera ad medium folii producta, basi limbo albido,
e cellulis angustissimis constructo, lamina dorsali e basi nerv:
enata, ibidemque rotundata, nervo crassiusculo, lutescente
infra summum apicem folii evanido, cellulis rotundato-hexagonis
0-007-0-OlOmm., pellucidis, papilla media instructis; seta termin
alls, solitaria, rarius binse ex eodem perichsetio, usque ad 2 mm
alta, tenuis, rubra, levissima; theca inclinata vel nutans, minuta
ovalis, pallida; operculum e basi conica recte rostratum; calyptra
ignota. Planta mascula ignota.
Near Hotel, Babinda, Cairns district, on rotten logs, n. 332a
(leg. Watts). ^
Species limbo brevi cellulisque pellucidis, papilla media in-
structis dignoscenda.
550 THE IMO.S.SES OF NORTH (^UEEK.SLAND^
IF. DlETRICHI^ CM.
Raveiisljoe, 11.-465 e.p., 528 e.p., 5*29 e.p.
¥. KERIANUS CM.
Cairns, n. 2Sia, 243; Stouey Creek, n. 252.
F. (Aloina) kurand.e Broth, et Watts, ii.sp.
Corticola, gregarie crescens, pallide viridis; caulis adscendens,
usque ad 8 nun. longus, cum foliis c. 15 mm. latus, basi fusco-
radiculosus, dense foliosus, simplex; folia usque ad 18-juga, sicca
liomomalla, humida erecto-patentia, infima minuta, dein sub-
sequalia, ligulata, breviter acuminata, obtusiuscula, apiculata, c.
1 mm. longa et c. 0*25 mm. lata, elimbata, Integra, lamina vera
ad medium folii producta, lamina dorsali e basi nervi enata,
ibidemque rotundata, nervo pallido, in apiculo folii evanido,
cellulis valde pellucidis, rotundato-hexagonis, 0*01 0-0-015 mm.,
ad basin laminae vertB paucis, elongatis, limbum inframarginalem
valde indistinctum efformantibus. Csetera ignota.
Kuranda, n. 273 (leg. Watts, 1913).
Species habitu F. Holliano Doz. et Molk., similis, sed cellulis
pellucidis, malto majoribus jam dignoscenda.
IF PALLIDUS H.f.W.
Raven shoe, n. 470; Malanda, n. 512.
IF. TKNELLUS H.f.W.
Ra.venshoe, n. 500 e.p.
*F. ziPPELiANUS Bryol. jav.
Babinda, n. 308 e.p., 343. Previous records, Sumatra, Java,
Hongkong.
[A'o^e.— The following also is recorded for N.Q.: F. micro-
lecythis CM. (Trinity Bay), but the name alone has been pub-
lished (CM., Genera Muscorum, p. 59). J
Calymperacese.
Syrrhopodon Schwgr.
*S. AMCENUS Broth.
Frenchman's Creek, n. 368 e.p., 385, 393; Babinda, n. 439.
Previous record, Papua.
BY V. V. BROTHERUS AND W. W. WATTS. 551
S (Cavi/olii) cairnensis Broth, et Watts, n.sp.
Dioicus; tenellus, cpespitosus, CiBspitibus laxiusculis, pallide
viridibus; caulis erectus, vix ultra 4 mm. lougus, basi fusco-
radiculosiis, dense foliosus, simplex vel ramosus ; folia sicca
circinato-incurva, dorso nitidiiiscula, humida erecto-pateiitia, e
basi paulum latiore ligulata, obtusa, mucroiiatula, usque ad 1-7
mm. longa, superne 0-25-0-27 mm. lata, lirabata, limbo aiigus-
tissimo, hyalino, infra apicem minute serrulatam evanido, in
parte superiore basis ciliato-serrato, nervo crassiusculo, infra
summum apicem folii evanido, inferne levi, dein parce aculeato-
papilloso, cellulis laminalibus acute et elevato-papillosis, basilari-
bus laxis, inanibus, breviter rectangularibus vel subquadratis.
Csetera ignota.
Cairns district, Babinda, on tree-trunks, n.313rt(leg. Watts,
July, 1913).
Species distinctissima, a S. fiynbriatulo Sauerb., foliis multo
brevioribus, nervo inferne levi, superne parce aculeato-papilloso
facillime dignoscenda.
*S. CROCEUS Mitt.
Frenchman's Creek, n. 418 e. p. An Indo-Malayan species.
8. FASCicuLATus Hook. et Grev.
Base of Bartle Frere ; Babinda ; Frenchman's Creek, etc.
Previous record, Johnstone River : Berthaud, Dec, 1882. A
widespread tropical species.
tS. FiMBRiATULUS Sauerb. S.Jimbriatus (J.M., non Mitt.
Frenchman's Creek, n. 408; Tully Falls, n. 462.
*S. MuELLEiii (l)oz. et Molk., Galymjyeridium) Lac.
Frenchman's Creek, n. 361, 368 e.p. An Indo-Malayan-Poly-
nesian species.
S. NoViE Valesi^ cm.
Ravenshoe, n. 461, 468, 53 1<'.. A forma inicrocarpa was col-
lected at Trinity Bay by Saver, in 1886; otherwise new for Q.
S. UNDULATULUS Broth.
Babinda, n. 313 e.p.; Frenchman's Creek, n.421 e.p.; Kuranda,
n 427. Previous record, Bellenden Ker : Mrs. Uribble (Herb.
Melb.).
552 'THE MOSSES OF NORTH QtTEEKSLAND_,
*S. UNDULATUS (Doz. et Molk.) Lindb.
Base of Bartle Frere, n.293; Babinda, n. 310; Frenchman's
Creek, n. 405, 412, 414, 420 e.p., 421. Previous record, Sunda
Islands.
[Note.— 5. i Thy ridium ) humile Mitt., is recorded in the Melb.
Herb, for Johnstone River (leg. Berthaud).]
C A L Y M P K R E S Sw.
tC MOLUCCENSE Schwgr.
Frenchman's Creek, n. 372. Hampe, in PL Preiss , recorded
it for West Australia, but this record is not found in Mitten's
Catalogue, nor in Jaegers xA.dumbratio. It is almost certainly
new for Australia. Previous range, the Moluccas.
C. NiGRESCENS Broth. et Geh.
Cairns: Allen, Sept., 1906 (Herb. J. M. Murdoch). Earlier
record, Bellenden Ker : Mrs. Gribble (Herb. Melb.).
*C. SERRATUM A.Br.
Malanda, n. 497, 530. Previous records, Java, New Hebrides,
etc.
*(/. TENERUM CM.
Fig-tree, Police Paddock, Cairns, n. 226-8, 230, 232: base of
Bartle Frere, n. 285. Previous range, Tndo-Malaysia.
[Note. — C. Motleyi Mitt., is recorded for Cape Yorke (leg.
Micholitz, 1897): and C. Keunedyo.num Hampe, (1876) was found
at Rockingham Bay by E. B. Kennedy, and at Goode Island by
Powell in 1883.]
Pottiaceoe.
W E I s I A Hedw.
W. viRiDULA (L.) Hedw.
Ravenshoe, n. 475 (f. eperistomata).
H Y 0 p H I L A Brid.
*H. MiCHOLiTZii Broth.
8toney Creek, n. 237 e.p., 261. Previous record, Papua.
B A R B u L A Hedw.
B. CALYCiNA Schwgr.
Ravenshoe, n. 464 e.p.
BY V. V. BROTHERUS AND W. W. WATT8. 553
Gpimmiaceae.
G L Y P H O M I T R I U M Bl'id.
to. COMMUTATUM (CM.) Broth.
Ravenshoe, n. 463 e.p., 488, 532 e.p.
Orthotrichaceae.
M A c R o M I T R I u M Brid .
tM. Daemelii cm.
Baveiishoe, n. 490, 536, 616 e.p.
tM DIMORPHUM CM.
Stoney Creek, n. 258; Ravenshoe, n. 485; Malaiida, ii. 491;
Kurauda, n.511, 6076.
M. HEMITRICHODES Schwgr.
Ravenshoe, several nos.
fM. PUGIONIFOLIUM CM.
Malanda, n.493, 495; Ravenshoe, n.506; Yungaburra, n.520.
Earlier records limited apparently to K.S.W.
tM. ScoTTi^ CM.
Ravenshoe, n.481, 486, 5346. Previous record, N.S.W., from
Monga and Nowra northwards to the Richmond River.
[Note. — J/, anrescens Hpe., is recorded from Mt. Archer, near
Keppel Bay (leg F.v.M.j; J/, micro stomum {lA.ook. et Grev.), for
Rockingham Bay (Dallachy, 1868); M. pusillunt. Mitt., from
Tringilburra Creek, Bellenden Ker (F. M. Bailey); J/. Sayeri
Mitt., from the Russell River (W. A. Sayer, 1886, Herb. Melb.),
and M. Whiteleygei Broth, et Geh., from BeJlenden Ker and
Bartle Frere (S. Johnson, 1891).]
Schlotheimia Brid.
S. Baileyi Broth.
Ravenshoe, several nos. The original recoi'd was Bellenden
Ker; the species spreads south well into N.S.VV., and probably
into Victoria.
Splachnaceae and Funariaceae.
These two families are quite unrepresented in my N.Q. col-
lection, but the following record may be noted : Goniomitriwm
acuininatuin H.f.W., (Clermont, Miss Gore). — W.W.W.
554 THE MOSSES OF KORTH QUEENSLAND^
Bryaceae.
B K A C H Y M K N I U M Schwgl'.
B, {Dicrauobryum) Wattsii Broth., sp.ii.
Dioicum; gracilesceiis, cfiespitosum, ca^spitibus densis, lutes-
centi-viridibus, sericeo-nitidis ; caulis erectus, vix ultra 1 cm.
longus, interne fusco-radiculosus, dense t'oliosus, simplex; folia
imbricata, ovata, breviter acuminata, aristata, marginibus basi
recurvis, integris, nervo lutescente. in aristam denticulatam ex-
cedente, cellulis superioribus elongate et anguste rhomboideis,
basilaribus et alaribus quadratis. Csetera ignota.
Ravenshoe (Millstreara), n. 489 and 527 (leg. Watts, 1913;.
Species Br. acuminato Har\ ., valde atiinis, sed statura paulum
robustiore, foliisque brevius acuminatis dignoscenda.
* A NOMOBRYUM Schimp.
*A. CYMBiFOLiUM (Lindb. ) Broth.
Ravensiioe, n.464 e.p., 5216, 532 e.p. Previous range, India
to Java and Amboina.
B K Y u M Dill, emend. Schimper.
fB. AKGENTEUM L., var. NIVEUM H.f.W.
Ravenshoe, n. 463 e.p,, 521 e.p.
fB. ERYTHKOCARPOIDES Hpe. et CM.
Kuranda, n. 429: Malanda, n. 521 e.p.
B. (Alpiniformia) kurand^e Broth, et Watts, n.sp.
Dioicum: gracilescens, caespitosum, csespitibus densiusculis,
pallide lutescenti-viridibus, nitidis: caulis vix ultra 5 mm. longus,
basi radiculosus, dense foliosus, innovationibus pluribus erectis,
vix ultra 5 nnn longis, strictis, julaceis, obtusis: folia caulina
erecto-patentia, carinato-concava, oblongo-lanceolata, breviter
acuminata, acuta, marginibus anguste recurvis, apice minutissime
serrulatis, nervo tenui, brevissime excedente, cellulis elongate
et anguste hexagono-rhomboideis, basilaribus laxioribus, alaribus
breviter rectangularibus; seta l-l'5cm. vel paulum ultra alta,
tenuissima, fuscescenti-rubra; theca nutans, minuta, cum collo
sporangii longitudinis, pyriformis, c. 2omm. longa et c. 0-95 mm.
crassa, sicca deoperculata sub ore baud constricta, macrostoma,
BY V'. F. BROTH ERIJS AND W. W. WATTS. 555
fusca; exostomii dentes lineari-laiiceolati, subulato-acuminati,
rufi, apice hyalini, dense laniellati; endostoiiiium luteum, rainu-
tissime papillosum, membraua basilaris ultra medium dentium
producta: processus lanceolati late feriestrati: cilia tcrna bene
evoluta, longe appendiculata. Csetera ignota.
Damp wall of creek Ijy Railway line, near Kuranda, n. 498
(leg! Watts, 1913;.
Species innovationibus julaceis, tlieca minuta, macrostoma
oculo nudo jam dignoscenda,
IB. PIMPAM.E CM.
Wall of creek by railway, near Kuranda, n. 679.
IB. SUBATROPURPURKUM CM.
Malanda, n. 505: Millstream, Ravenshoe, n. 521a.
IB. SUBOLIVACEUM CM.
Millstream, Ravensliue, n. 525 e.p ; Kuranda, n. 535.
B. SUBPACHYPOMA Hampe.
8toney Creek, n. 249, 255, 259 Earlier record, Rockingham
Bay (Dallachy, 1869).
\_Note. — The following records should be noted : B. Baileyi
Broth., (Freshwater Creek, near Cairns, Bailey, 1889; Bellenden
Ker, 8. Johnson); B. leptothrix CM., (Trinity Bay, Karsten,
1881).]
Rhodobryum (8chimp. ) Hampe.
jR. OMVACEUM Hpe.
Ravenshoe, n. 506.
fR. SUBCRISPATUM (CM.) Bar.
Yungaburra, on hillside, n. 501.
Mniacese.
M N I u M (Dill, exp.) L., emend. Schimper.
Mn. sp.(?).
Ravenshoe, n. 517 e.p.
Mn. sp.(?), probably sp.nov.
('?)Loc., n. 522.
556 THE M()S8F»S OF NORTH QUEENSLAND,
Rhizogoniacese.
R H I z o G o N I u M Brid.
fR.H. Geheebii cm.
Ravenshoe, n. 483. 502.
Rh. parramattense cm.
Ravenshoe, n. 523, 615 e. p.; Malanda, n. 618c. A southern
species, but already recorded (in Hb. Melb.) for Bellenden Ker.
Rh. spiniforme (L.) Bruch.
Malanda, Kuranda, and Ravenshoe, including f. minor, n.482,
492 (previously recorded from N.S.W., and S.Q.).
[Note. — Rh. brevifoliuin Broth., was collected on Bellenden
Ker by F. M. Bailey (1889), also by S. Johnson, on the same
range, and on Bartle Frere in 1891.]
M E s o c H iE T e Lindb.
M, UNDULATA Lindb.
Babinda, n. 387. Common in the south, with, also, a record
for Port Denison (Birch, 1889).
Bartramiaceae.
Philonotis Brid.
*Ph. laxissima (CM.) Bry. jav.
Babinda, n. 334; forma, Ravenshoe, n. 517, 524.
tPh. pseudOxMollis (CM.) Jaeg.
Ravenshoe, n. 489 e.p., 516: Barron River, n. 496, 534(7.
Polytrichaceae
POLYTKICHUM Dill.
P. JUNiPERiNUM Hedw.
Ravenshoe, n. 519.
B. PLEUROCARPL
Cryphseaeese.
C R Y p H yE A Mohr.
C TEN ELLA Hornsch.
Ravenshoe, n. 610 e.p., 616 e.p.
BY V. F. BROTHERUS AND W. W. WATTS. 557
C R Y p H I D I u M fMitt. ) Broth.
C MoELLERi (Hampe) Par.
Ravenshoe (Millstreani, etc.), several nos.
Cyrtopodaeese.
B E S C H E R K L L K A Dub.
B. Cyrtopus F. v.^r.: B. hrevifolia Hpe.
Ravenshoe, several nos.
Ptyehomniacese.
H A M P E E LL A CM.
H. FALLENS (Lac , Cladomiiioii) Fleisch.; Lepidopihi.m aiistrale
Broth.; HampeeUa Kurzii CM.
Tally Falls, ii.591. [Previously recorded from Harvey's Creek:
Bailey, 1889].
(Edieladiaeese.
"*M Y u R I u M Schiinp., Fleischer.
*M. RUFESCKNS (Hsch. et Reinw.) Fleisch.
Ravenshoe, n.534« (f. depauperata). Range, Indo-Malaysia.
Neekeraeeae.
T K A c H Y L o M A Brid.
T. PLANIFOLIUM (Hook.) Brid.
Ravenshoe, n. 543; Malanda, n. 650 e.p.
Endotrichella cm.
E. DlETRICHIiE CM.
Babinda, n. 440; Ravenshoe, n. 558, 5726, 616 e.p.; track to
TuUy Falls, n. 561.
E. LEPIDA CM.
Frenchman's Creek, 417 e.p.; Kuranda, 511 e.p., 607«; Ravens-
hoe, 476 e.p., 564 e.p.
Garovaglia Endl.
G. MuKLLERi (Hpe.) Mitt.
Kuranda; Babinda; Malanda; Ravenshoe, etc.
MUELLERIOBRYUM Fleisch.
.M. Whiteleggei (Broth.) Fl.
Stoney Creek ; Babinda ; Frenchman's Creek ; Malanda ;
Ravenshoe,
558 THE MOSSES OF NORTH QUEENSLAND^
*Pterobryopsis Fleisch.
P. FiLiGEHA Broth, et Watts, n.sp.
Dioica: gracilis, ij;regarie crescens, Ifete viridis, nitidiiiscula;
caiiles secundarii usque ad 3 em. loiigi, inferne simplices, nudi,
dein densissime pinnatim raniosi, filis elongatis, sat nuinerosis,
articulatis, papillosis instructi, ramis erecto-patentibus, vix ultra
6 mm. longis, complanatulis, simplicibus, obtusis; folia imbricata,
subcymbiformi-concava, dorso Ijfpvia, ovato-oblonga, breviter
acuminata, acuta, marginibus erectis, ubique serrulatis, Tiervo
tenui, subcontinuo, dorso superne serrato, cellulis aiigustissimis,
apice papillose exstante, basilaribus infimis laxioribus, alaribus
sat numerosis quadratis. Cnetera ignota.
Malanda, on trunks of trees, n. 549; Ravenshoe, on trunks of
trees, n. 605a.
P T E R o B R Y I D I u M Brotli. et Watts. n.gen.
Pt. australe B. et W., n.gen. et sp.
Dioicum; robustiusculum, Isete viride, setate lutescens, nitidum;
caulis primarius longe repens, plus minusve fusco-radiculosus:
caules secundarii conferti, rigidi, superne ssepe arcuati, usque ad
12 cm. longi, dense foliosi, plerumque obtusi, rarius tiagelli-
formiter attenuati, inferne simplices, dein plus minusve regu-
lariter pinnati, ramis patulis, teretibus arcuatulis, usque ad
2cm. longis, obtusis vel plus minusve longe, rarius flagelliformiter
attenuatis, simplicibus, rarius longioribus, parce ramulosis; folia
caulina infima, minuta, dein sensim accrescentia, cochleariformi-
concava, sicca imbricata, Isevia, late ovalia, apiculo lato, obtuso,
recurvulo terminata, nervo simplici basi validiusculo, dein raptim
tenuiore, ultra medium folii evanido, cellulis inter se porosis,
linearibus, chlorophyllosis, basilaribus infimis fusco-aureis, alari-
bus numerosis, subquadratis; bractese perichaetii erectse, concavse,
internse oblongaj, raptim breviter acuminatse, c. 5mm. longse et c.
1*5 ram. latse, integrse, nervo tenui, vix ad medium bracteie pro-
ducto, cellulis linearibus, inter se valde porosis, basilaribus laxi-
oribus; seta c. 1 mm. alta, stricta, fusca; theca immersa, breviter
ovalis, sicca deoperculata siib ore paulum constricta, c. 2 mm.
longa et c. 090 mm. crassa, fusca; peristomium duplex, infra
BY V. F. BROTHERUS AXD W. W. WATTS. 559
orifieinm oriundum, rufaii), la^ve, cum praeperistomio; exostomii
dentes angiiste lanceolati, linea dorsali recta, articulationibus
lamelliformibus: corona hasilarisrudimentaria: processus angusti;
sublineares. Csetera ignota.
North Queensland: Malanda, n. 619a; Ravenshoe in scrub,
n.551e.p.,r)5o, 556, 557 e.p., (Watts, 1913). Queensland-N.8 W.:
Mt. Lindsay (J L. Booiman, n.l522 in Herb. S3^dn.).
Genus novum insigne, habitu foliorumque structura speciebus
nonnullis Pterohryopsidis simillimum, sed theca immersa nee
non peristomio dignoscendum.
P A P I L L A R I A (CM. ) CM.
P. AMBLYACIS (CM.) Jaeg.
Lake Barrine, n. 636.
P. FLEXiCAULis (Tayl.) Jaeg.
Ravenshoe, n. 556 e.p., 610 e.p., 654a; Malanda, n. 619 e.p., 647.
P. KERMADECENSis (CM.) Jaeg.
Babinda; Malanda; Ravenshoe; several nos.
fP. NiTiDiuscuLA Broth.
Malanda, n. 618 e.p., 6196. Previous records, N.S.W.
M E T E o R I u M Doz. ct Molk.
M. Baileyi (Broth., Papillaria) Broth., Bryales, p. 818.
Ravenshoe; Malanda; Yungaburra; several nos.
*M MiQUELiANUM (CM.) Fleisch.
Malanda; Ravenshoe; several nos.
Akrobryopsis Fleisch.
A. SIGMATOPHVLLA (CM.).
Base of Bartle Frere; Babinda; Frenchman's Creek; Malanda;
Lake Barrine; Ravenshoe: many nos.
■''"F L O K I B U N D A K I A CM.
*Fl. floribuxda (Doz. et Molk., Mfteorinm) Fleisch.
Malanda, n. 4946, 619^; Yungaburra, n. 668 e.p.; Ravenshoe,
n.579.
■*Fl. pseudofloribunda Fleisch.
Malanda, n.651 e.p., 655; Ravenshoe, n.582, 624 e.p., 656.
560 THE MOSSES OF NORTH QUEENSLAND^
Fl. robustula Broth, et Watts, n sp.
Dioica; robustiuscula, palHde lutescenti-viridis, opaca: caulis
secundarius breviusculus, dense pinnatim ramosus, ramis patulis,
usque ad 2 cm. longis, apicem versus decreseentibus, laxiuscule
foliosis, simplicibus, obtusis; folia ramea horride patula, e basi
cordatoovata lanceolato-subulata, comalia in pilum tiexuosum,
denticulatum attenuata, marginibus basi uno latere inflexis,
apice crenulatis, nervo tenui, ultra medium folii evanido, cellulis
elongatis, angustis, seriatim papillosis. Csetera ignota.
N. Queensland : Malanda, on tree-trunks, n.618rt.
Species statura robustiuscula foliisque horride patulis oculo
nudo jam dignoscenda.
Bakbella (CM.) Fleisch.
B. ENERVis (Thw. et Mitt.) Fl.; Neckera trichophoroides Ilpe.
Ravenshoe, n. 558, 624, 628 e.p.
*M ETEORioPSis Fleisch.
*M. RECLINATA (CM.) Fl.
Malanda, 604, etc.; Yungaburra, n.626a; Ravenshoe, 541, etc.
*T RACK Y PUS Reinw. et Hornsch.
*T. NoViE CALEDONIiE CM.
Malanda, n. 633.
Orthokrhynchium Reichdt.
O. CYMBIFOLIOIDES CM.
Ravenshoe, n. 600, 605 e p., 611 e.p.
Calyptothecium Mitt.
C acutum (Mitt.) Broth.
Babinda, n. 317; Ravenshoe, n. 551, etc.
C HUMiLE (Mitt.) Broth.
Ravenshoe, n. 557 e.p.; Yungaburra, 612,etc.; Malanda, n. 61%.
0. RECURVULUM (CM., Trachylomo) Broth.
Babinda, n. 312 e.p.
Neckeropsis Reichdt., emend. Fleisch.
N. Lepineana (Mont.) Broth.
Malanda; Ravenshoe; Yungaburra: several nos.
BY V. F. BROTHERUS AXD W. W. WATTS. 561
*HiMANTOCLADiUM (Mitt., Neckero) Fleisch.
*?[. LORIFORME (Bryol. jav.) Fl.
Base of Bartle Frere, n. 294; Babinda, n. 437.
*H OMALIODENDRON Fleisch.
*H. EXIGUUM (Bryol jav., Homalia) Fl.
Ravenshoe, n. 546, 576e.
*H. FLABELLATUM (Dicks, 8m., Hookevia) Fl.
Ravenshoe, n. 547-8, 564 e. p., G09.
*PiNNATELLA (CM.) Fleisch.
*P. INTRALIMBATA Fl.
Ravenshoe, n. 600a, 611, 632, 667 e.p.
Lembophyllaeese.
C A M p T o c H ^ T E Reichdt.
C. BRLSBANICA (CM.).
Ravenshoe, ii 542, 595, 648 e.p.; Lake Bairine, n. 636.
C VAGA (Hornsch.) Broth.
Babinda; Frenchman's Creek: JVIalanda : several nos.
Entodontaeese.
*C A M P Y L O D O N T I u M Doz. et Molk.
*C. PLAVESCENS (Hook., Ptevogonium) Bryol. jav.
Ravenshoe, n. 538, 581 e.p.
Fabroniacese.
*A U S T IN I A CM.
*A. LUZONENSis Broth.
Ravenshoe, n. 667 e.p.
Hookeriaeese.
DiSTICHOPHYLLUM Doz. et Molk.
D. Baileyanum CM.
Kuranda; Ravenshoe; Yungaburra : several nos.
Cyclodictyon Mitt.
C LEPIDUM (Mitt.); Hookevia karsteniana Broth. Geh.
Yungaburra, n. 576a, 674.
42
562 THE MOSSES OF NORTH QUEENSLAND^
Callicostella (CM.) Jaeg.
■*C. KiERNBACHII Broth.
Kuranda, n. 266, 272; Babinda, 332 bis.
Ch^etomitrium Doz. et Molk.
Ch. entodontoides Broth, et Watts, n.sp.
Dioicum; gracile, ca)spitosum, csespitibus laxis, mollibus, de-
pressis, lutescenti-viiidibus, nitidiusculis; caulis lepens, fusco-
radiculosus, dense et complanate foliosus, dense pinnatim ramosus,
ramis vix ultra 1 cm. longis, complanatis, obtusis; folia erecto-
patentia, concaviuscula, ovato-oblonga, rotundato-obtusa, dorso
Isevia, marginibus erectis vel basi anguste recurvis, superne
argute denticulatis, nervis binis, brevibus, indistinctis, cellulis
angustissimis, linearibus, infima basi brevioribus et latioribus,
aureis, omnibus l?evissimis. Caetera ignota.
North Queensland: Frenchman's Creek, Cairns District, n.352.
Species distinctissima, habitu entodontoideo foliorumque forma
facillime dignoscenda.
Ch. Geheebii Broth.
Babinda, n. 336^.
Ch. nematosum Broth.
Babinda: Frenchman's Creek; Ravenshoe : several nos.
Lepidopilu7n austral e. See Hampeella pallens.
Hypopterygiaeese.
H ypoPTERYGiUM Brid.
H. Muelleri Hpe. et CM.
Lake Barrine, n. 643.
Helieophyllaeese.
P 0 W K L L I A Mitt.
P. AUSTRALis (Hpe., Helicophyllum) Broth.
Frenchman's Creek, n. 401. Found earlier by Dallachy at
Rockingham Bay.
Rhaeopilaeese.
Rhacopilum Palis.
Rh convolutaceum cm.
Babinda; Frenchman's Creek; Malanda; Ravenshoe.
BY V. F. BROTHERUS AND W. W. AVATTS. 563
Leskeaeese.
*H ERPETiNEURON (CM.) Cardot.
*H. ToccOiE (Sull. et Lesq., Anomodon) Card.
Ravenshoe, n. o98 e.p.
T H u I D I u M BryoL eur.
ITh. lilliputanum Broth.
Ravenshoe, n. 648, 665.
Th. plumulosiforme (Hpe.) Jaeg.
Stoney Creek; Frenchman's Creek; Kuranda; Ravenshoe.
ITh. suberectum (Hpe.) Jaeg.
iStoney Creek; Malanda; Ravenshoe.
\^Note. — The following species have also been recorded for
N.Q.: Til. rameihtosum Mitt , ( Bellenden Ker: Karsten); Th. spar-
simi (Bellenden Ker : Karsten), and perhaps others. — W. W.W.J
Hypnaeese.
StereodontecE.
E C T K O P O T H E C I a M Mitt.
E. SERRIFOLIUM Broth. et Watts, n.sp.
Dioicum (1) : gracile, csespitosum, c?espitibus densiusculis^
rigidis, depressis, Isete viridibus, nitidiusculis; caul is elongatus,
repens, fusco-radiculosus, dense subpinnatim raniosus, ramis vix
ultra 5 mm. longis, strictis, dense et complanate foliosis, simplici-
bus, obtusis; folia erecto-patentia, caulina superiora plus minusve
distincte homomalla, concava, caulina ovato-lanceolata, plus
minusve longe acuminata, marginibus inferne indistincte re-
curvis, superne erectis, argute serratis, nervis binis, brevibus,
cellulis anguste linearibus, apice distinctissime papillose exstante,
marginalibus in serie unica brevioribus, ramea breviter et late
acuminata. Csetera ignota.
Cairns District : Millstream Falls, Ravenshoe, n. 528 (leg. Dr.
Joynt).
Species E. Zollingeri (CM.) Jaeg., affinis, sed foliis argute
serratis jam dignoscenda.
E. UMBICULATUM (CM., Hyjmuni) Par.
Stoney Creek; Kuranda; Babinda; Ravenshoe, etc.
564 I'HK MOSSES 01' NORTH QUEENSLAND^
*E. VERRUCOSUM (Hpe.j Jaeg.
Raveiislioe, n. 539 e. p., 606, 653 e. p., 661.
►Stereodon Mitt.
S. MOSSMANNIANUS (CM,, Hypnum) Broth,
Millstream, Ravenshoe, ii. 521 e.p,
fS. SUBCHRYSOGASTER Brotll,
Rocky Hill, Kuranda, n. 680 e.p. Previous loc, Blackall
Range: leg, WhitteroD.
A C A N T H O C L A D I U M Mitt.
A. EXTENUATUM (Brid.) Mitt.
Frenchman's Creek; Malanda; Ravenshoe; Tully Falls.
A. sp.?
B-avenshoe, n. 605 e.p.
Flagiotheciece.
I S 0 P T E R Y G I U M Mitt.
I. AusTRo-PusiLLUM (CM.) Jaeg.
iStoney Creek; Kuranda; base of Bartle Frere; Babinda.
I. CANDiDUM (CM.) Jaeg,
Kuranda; Mai and a.
I. SUBLATIFOLIUM Broth.
Base of Bartle Frere, n. 299 e,p., 305.
Taxithelium Sprue.
T. KERIANUM (Broth., Trichosieleum) Broth., Bryales.
Base of Bartle Frere, n. 285 e.p., 298; Babinda, n. 320 e.p.;
Tully Falls, n. 591 e.p., 597.
fT. Nov^ Valesi^ (Broth., Isopterygium) Broth.
Babinda; base of Bartle Frere: Malanda; Yungaburra; Tully
Falls, etc.
*T. PAPILLATUM (Harv.) Broth., Bryales.
Babinda, n.424; Frenchman's Creek, n. 362a.
BY V. F. BROTHERtTS AND W. W. WATTS. 565
T. (Poli/stigma) Wattsii Broth., n.sp.
Dioicum; robustiusculum, csespitosum, csespitibus densis, de.
pressis, viridibus, nitidis : caulis elongatus, repens, per totam
longitudinem fusco-radiculosus, dense pinnatim ramosus, ramis
patulis, valde complanatis, dense foliosis, obtusis, rarius f^agellis
microphylliiiis instructis, cum foliis c. 1 7 mm. latis, vix ultra
5mm. longis, siinplicibus vel longioribus, pinnatim ramulosis;
folia ramea lateralia patentia, valde concava, e basi paulum
angustiore oblonga, sensim lanceolato-acuminata, marginibus
erectis, apice minute serrulatis, nervis binis brevibus vel nullis,
cellulis angustissimis, seriatim minutissirae papillosis. basilaribus
infimis vesiculosis, hyalinis. Caitera ignota.
Babinda, North Queensland, on living tree-trunks, n. 441.
Species valde peculiaris, cum nulla alia commutanda.
Vesiculauia (CM.) CM.
tV. RiVALis Broth.
Stoney Creek, n. 256 e.p. A second locality. The type was
collected by Watts on the Richmond River (Whian Creek).
Sematophyllacese.
M E I O T H E C I U M Mitt.
M. Brotheri Watts; M. tenerum B.P., non Mitt.
Cairns (Police paddock, on tig tree), n. 227, 228, 230; Street's
Gully, Kuranda, n. 270«, 607.
fM. Wattsii (^Broth., Ptcrogoniella) Broth., Bryales.
Kuranda, n. 607^, 642.
R H a p h I d o s t e g I u m Schimp.
Rh. aciculum (CM.).
Stoney Creek; Kuranda; Ravenshoe, etc.
\iu. OVALE Broth.
Babinda; Frenchman's Creek; Ravenshoe.
Rh. pseudo-homomallum (CM.).
Kuranda; base of Bartle Frere; Babinda; Frenchman's Creek;
Ravenshoe.
566 THE MOSSES OF NORTH QUEENSLANd^
*Rh. saproxylophilum (CM.) Jaeg.
Frenchman's Creek, n. 362; f. rohnsta (n. 415).
Sematophyllum (Mitt.) Jaeg.
8. ERYTHKOPODiuM (Hpe.) Jaeg.
Malanda, n. 625 e.p. Previous record, Rockingham Bay :
Dallachy.
S. sp.'? (ster.): Ravenshoe, n. 588.
S. sp.] (ster.): Tally Falls, n. 591 e.p.
*T R I c H o s T E L E u M (Mitt.) Jaeg,
T. ( Papillidiu77i) elegantulum Broth, et Watts, n.sp.
Autoicum; tenellum, csespitosum, ca^spitibus densis, pallide
viridibus, retate lutescenti — vel fuscescenti- — viridibus, nitidi-
usculis; caulis breviusculus, per totam longitudinem f usco-radicu-
losus, dense ramosus, ramis adscendentibus, complanatis, brevi-
bus, simplicibus vel longioribus plus minusve ramulosis; folia
erecto-patentia, concava, e basi anguste oblongo-elliptica sensim
lanceolata vel lanceolato-subulata, raargiriibus late recurvis,
superne argute serratis, enervia, cellulis anguste ellipticis, superi-
oribus papilla media altiuscula instructis, infimis aureis, alaribus
magnis vesiculosis, hyalitiis vel aureis; bractea^ pericheetii internee
erect?e, e basi vaginante in acumen loriforme, argute serratum
sensim attenuatte ; seta vix ultra 8 mm. alta, tenuis, rubra,
ubique mammillis humilibus latis, luteis obtecta; theca suberecta,
majuscula, ovalis, fusco-rubra, Isevis; operculum ignotum.
Cairns District (Frenchman's Creek), n. 382.
Species distinctissima, foliis angustis, nee non theca majuscula
oculo nudo jam dignoscenda.
T. HAMATUM (Doz. et Molk.) Jaeg.
*Var. SEMiMAMMiLLOSUM (CM.) Par. ""
Babinda, n. 3 19 e.p., 337«; Frenchman's Creek, n. 3626, 383.
The var. previously recorded for Papua and the New Hebrides.
RhegmatodontaeesB.
*M A C R O H Y M E N I U M CM.
*M RUFUM (Reinvv. et Hornsch.) CM.
Malanda, n. 623; Ravenshoe, n. 657, 660, 662.
BY V. F. BROTTTERI^'^ AXD TV. W. WATTS. 567
Braehytheciaceae.
Rhynchostegiella (Bryol. eur. ) Limpr.
Rh. convolutifolia (Hpe.) Broth.
Base of Bartle Frere, n. 297 e.p.
R H Y N c H o s T E G I u M Bryol. eur.
Rh. nano-pennatum (Broth.) Par.
Base of Jiartle Frere, n. 285 e.p., 296-7; Babinda, n. 320 (bis);
Frenchman's Creek, n. 416.
Rh. tenuifolium (Hedw.) Jaeg.
Ravenshoe, n. 425 e.p., 476 e.p.; Yungaburra, n. 601.
Hy pnodendraeese .
H Y p N o 1) E N D R o N (CM. ) Lindb.
H. SPININERVIUM (Hook.) Jaeg.
Kuranda; Babinda; Yungaburra; Ravenshoe.
Mniodendkon Lindb.
Mn. comatulum Geh. in Herb. Brotii.
l)ioicum; gracilescens, viride, setate fuscescens, nitidiusculum;
caules secundarii approxiniati, usque ad 4 cm. longi, a foliis
squama^formiV)us, remotis, patentissimis, reflexis, tomento den-
sissimo fusco-purpureo occultis obtecti, superne verticillatim
ramosi, ramis 1-1 5 cm. longis, gracilibus, arcuatulis, undique
dense foliosis, simplicibus; folia caulina e basi late cordata, tri-
angularia, longe aristata, marginibus erectis, basi minute denti-
culatis, superne argute et insequaliter serratis, nervo longe ex-
cedente, cellulis angustissimis, alaribus sat numerosis, ovali-
hexagonis, fusco-aureis; folia ramea patentia, ovato-lanceolata,
breviter acuminata, marginibus basi integris, superne argute et
iiia3qualiter serratis, nervo apice evanido, superne dorso serrato;
seta usque ad 2 cm. alta, flexuosula, tenuis, fuscidula; theca
horizontalis, oblongo cylindrica, c. 3 mm. longa ; operculum e
basi convexo-conica longe rostratum.
Charmellan's Creek, on Gordon's track from Ravenshoe to
TuUy Falls, n. 676. Mt Armit : Pentzke in Herb. Melb.(type),
568
MESOZOIC INSECTS OF QUEENSLAND.
No. 4. Hemiptkra Hkteroptera : The Family Dtnstaxiid^'E.
With a Note on the Origin of the Hkteroptera.
By R. J. Tillyard, M.A., D.Sc, F.L.S., F.E.S., Linnean
Macleay Fellow of the Society in Zoology.
(Plate lix. ; and Text-figures 17-22).
In 1916, I described (3) the l)eautifull\' preserved wing of
Dunstanio, pulchra Till., from the Upper Trias of Ipswich,
Queensland, and placed it as the sole representative of a new
family D\i7istaiiiid(E within the Order Lepidoptera. This deci-
sion had the concurrence of Dr. A. Jefferis Turner, of Brisbane,
and was based mainly upon the presence of eight longitudinal
veins in the fossil, this being the number found in the hind wings
of Frenate Lepidoptera.
While the description of this fossil was going to press, I hap-
pened to be on a visit to Brisbane. A day or two before I left,
Mr. Dunstan showed me four other specimens bearing the label
*^ Du7ista7iia '' in pencil; these had just been discovered at
Ipswich. Two of them were broad wings resembling the type,
and two were much longer and narrower wings. Thinking that
these latter must be the forewings of the same insect as that
whose hindwdng I had assumed the type to be, I added the note
on p. 32 of the paper quoted above. However, when I received
from Mr. Dunstan, later on, the complete collection of Ipswich
Insects of which these wings formed a part, and had time to
work at them in detail, I soon found that the two narrow wings
did not belong to Dutistania at all; so that there remained onl}"
the two broad wings for study in connection with the original
type. Neither of these is in anything like as good a state of
preservation as the type is, and one of them is very poorly pre-
served indeed.
BY R. J. TILLYARD. 569
Meanwhile the publication of the original description had
created considerable interest amongst entomologists, and discus-
sion soon became rife as to its true affinities. In August, 1916,
Mr. E. Meyrick, F.R.S., contributed a paper on the subject* (2),
in which he argued strongly against the admission of Diinstania
to the Order Lepidoptera, and suggested that its affinities might
rather be with the Homoptera. He reasoned as follows :— "In
order to appreciate the position, I looked up the record of
Palfpontina oolitica Butl., from the Jurassic, hitherto the oldest
known Lepidopteron, ; T must express my opinion that
there is little doubt it belongs to the Ilepialidce Now it
is clearly proved on structural grounds that the Micropterygina
(the small group to which the Hepialidre belong, otherwise
termed Jugatse) were the primitive form of the Lepidoptera, and
the nature of Palceontina is in accord with this conclusion and
confirmatory of it. Parenthetically, I take occasion to notice
that Goss, Tillyard, and others are troubled as to what the early
Lepidoptera can have fed upon, when Howering plants were not
yet in existence, and honey was not available; it is a problem
easy of solution, since the more primitive forms (including the
Hepialidce) have in general no proboscis or feeding apparatus,
and, therefore, fed on nothing in the perfect state."
" With the evidence recounted above, Dmistaiiia, if regarded
as Lepidopterous, is violently discordant As the hind wing
possesses neither frenulum nor prominent basal angle of costa,
it must be assumed that the forewing was furnished with a
jugum; all existing Lepidoptera exhibit one or other of these
structures. But all jugate Lepidoptera have the hindwing
similar in neuration to the forewing, with at least 1 1 veins,
whereas this wing seems to be of the modern 8- veined type,
though conspicuously different from any known form, and in fact
quite as highly specialised as any now existing. Finally, there
is the apparently corneous margin round the costa and termen,
which is altogether abnornial, no other Lepidoptera showing a
* See also " Nature," No. 2488, Vol. 99, Sept. 28tli, 1916, p. 75, where
this paper is reviewed.
44
570 MESOZOIC INSECTS OF QUEENSLAND, iv.,
trace of it, and in my estimation a far more important systematic
character than the two which the author lays stress on."
"It appears to me, then, that this specimen can only be
regarded as Lepidopterous on the theory that it is a member of
an entirely unknown line of development of that Order, which
had attained a high degree of specialisation at a period far
anterior to the earliest certainly Lepidopterous insect known,
itself a lowly organised form; this cannot be termed impossibffe,
but it involves a very great improbability. The alternative view
that the resemblances are accidental and the insect not Lepi-
dopterous seems to me, on the whole, less improbable."
" I cannot venture to express any positive opinion on its pos-
sible relation to other Orders, but I suggest that theie are some
points of resemblance to the Hemiptera-Homoptera, some species
of which have a semi-Lepidopterous facies. The corneous margin
of the wing, the central cell (usually, it is true, much larger), and
approximate number of veins rising from it, and even the curious
dark banding of the membrane alongside the veins, can all be
paralleled in this group, which, moreover, is already known to
have been in existence since the Carboniferous period."*
Mr. Meyrick's criticism led me to study intensely the various
types of Lepidopterous wing-venation extant, about which 1
knew very little at the time that I first described the fossil. I
very soon became convinced that Dunstania was not a Lepidop-
teron; but that conviction was not based, for the most part, on
the argument set forth by Mr. Meyrick, which I regard as un-
sound in several important particulars.
Firstly, as regards Palceontina. The latest authority on the
family to which this fossil belongs is Handlirsch(l), who gives a
masterly treatment of this and allied fossils, with photographs
and restorations of a number of types, in some of which the
hindwing is complete enough to allow of definite proof that the
Falceoniinidce were most certainly not Hepialidm, as Meyrick
avers, nor even Jugatse of any kind, but definitely Frenatse allied
* This is surely an error. No true Homoptera were known below the
Lias, until I described Mesojassus from the Ipswich Trias, and, later on,
two new genera from the Permian of Newcastle, N.S.W.
IBY n. J. TiLLYARt). 571
to the Limacodidce. If Mr. Meyrick " turned up the record of
Palceontina oolitica,'' how is it that he ignored Handlirsch's
epochal work, which does not fit in with his own opinions 1
I accept without any doubt whatever Handlirsch's proof that
the PalcEontinidcE were Frenate moths, though I take leave to
doubt that they were as closely allied to the Limacodidce as he
imagined. The evidence of the hindwings, which are small, with
only eight or nine veins, is conclusive on the point. Conse-
quently, there is no absurdity in supposing that Frenate types
might have existed also in the Lias, or in the Upper Trias of
Australia. That being so, all this part of Mr. Meyrick's argu-
ment rests upon an unsound basis.
There is, therefore, no a priori reason why a Frenate moth, of
a type^a?' removed from those we know at the present day, might
not also have been in existence in the Upper Trias at Ipswich,
though I am quite willing to admit that it might not be a very
probable supposition.
(Secondly, as regards the "apparently corneous margin" round
the costa and termen. In my description, I simply called this
" a wide margin without pits," and added that " the external
border, or termen^ shows signs of a delicate transverse ribbing "
which " extends also somewhat indistinctly round the apex on
to the costa." I purposely refrained from suggesting that it was
a corneous margin, as it seemed to me to be equally likely that
it had been formed by the crushing down of the deep bases of
insertion of a series of stiff and closely packed marginal hairs or
bristles, such as one sees in many Diptera, for instance. There
was no inherent improbability that such a margin might not bo
found on a strongly-built and hairy-winged, primitive Lepi-
dopteron.
If, however, the margin be really corneous, then I admit that
the character is of great importance, and definitely rules Dun-
stania out of the Lepidoptera. In the actual fossil, the appear-
ance of this margin is certainly not such as to suggest a positive
solution to this question, one way or another; but Mr. Meyrick,
of course, had only my drawing to go upon.
Thirdly, I should like to reply to Mr. Meyrick's parenthetical
572 MESOzoic Insects of Queensland, iv.,
statement tliat the early ancestors of the Lepidoptera took no
food in the iraaginal condition. This argument defeats itself.
For, if these insects took no food as imagines, then 'surely their
descendants must have even more aborted mouth-parts than
their ancestors, and thus they should all be either Hepialidce or
Saturniidcp. I Surely the ancestors of those Lepidoptera that are,
to-day, of the haustellate type were originally of the weak man-
dibulate type still preserved in Micropteryx and allies, and were,
therefore, pollen-feeders and lappers-up of dew and rain-drops,
as the great majority of the Planipennia, Mecoptera, Trichoptera
and Diptera are to day !
To my mind, the great argument against Diinstania being a
Lepidopteron is simply this, that the venation, quite apart from
the mere number of the veins, does not show any of the typical
arrangements of that Order, whether Frenatae or Jugatpe. As
soon as I had sufficiently mastered the wing-venation uf the
Order, I became convinced of this, and definitely rejected my
original placing of the fossil. It is part of the object of this
paper to relate how the opinions of different entomologists, look-
ing at the problem from different points of view, favoured the
claims of various Orders to receive this fossil; but none, includ-
ing myself, who had the type-specimen itself to study, succeeded
in finding the true solution, which was at last given by the
study of the new material.
Starting from Mr. Meyrick's suggestion that Dunstania might
be Homopterous, I find that a number of correspondents enter-
tained this idea, the strength of which lies, of course, in the in-
terpretation of the margin of the wing as corneous. However,
Mr. J. Edwards, of Cheltenham, England, an acknowledged
expert upon this Order, wrote to Mr. Meyrick, and also to me,
strongly opposing the idea, on the ground that he could see
nothing in the fossil to indicate such a relationship. As the
Homoptera are well represented in the Ipswich Trias by forms
differing entirely from Dunstania, and as I was unable to estab-
lish a single venational character common to the two types, I
soon abandoned any idea of relationships in this direction.
In correspondence, other entomologists discussed with me the
tiV R. .t. TlLLYARt). 573
probable affinities of Dunstania, but their opinions were not
published. Jn particular, I desire to mention Mr. Herbert
Campion's share in the clearing up of the mystery. It was
through his unfailing.courtesy that I was kept in touch with
the trend of opinion in England on the fossil, and from him I
obtained valuable suggestions as to clues that were worth follow-
ing out. One of the most interesting of these, originating with
Mr. Durrant of the British Museum, and stated in publication
by Dr. Bather, F.R.8., in the course of a short review of my
paper in " Nature,"* was, that the venation of Duiistnida might
be compared with that of certain groups of Diptera, such as the
Fsycliodidce. At about the same time. Dr. C. J. Gahan, of the
British Museum, had before him a small collection of insects
from New Zealand, and suggested to Mr. Campion that the fossil
should be compaied Avith the Anthomyiid Exsul sing^day'is
Huttou, which it closely resembles in the shape and general ap-
pearance of the wing.
Acting on these suggestions, I put myself into communication
with several New Zealand entomologists, in the endeavour to
obtain material of Exsul. This, however, was not easy to get,
as the fly is exceedingly rare, and regarded as a great prize.
After a considerable time, I ol)tained two specimens, one from
Mr. David Miller, Government Entomologist of New Zealand,
and an expert Dipterologist, and another from Mr. G. V. Hudson,
of Wellington. To both of these gentlemen I offer my cordial
thanks for their generosity in sending me such valuable material.
While I could not admit any affinity between Dunstania and
the Psychodid(e^ I was impelled to study more closely its possible
affinity with Exsul., in spitn of some cardinal differences in the
venational scheme, which would be hard to explain on any
hypothesis. Believing that I had here, at last, a good solution
of the problem, and wishing to give due credit to those to whom
the suggestion was originally due, 1 wrote to "Nature,"! stating
ray adherence to the probable Dipterous nature of Diuistaida.
However, it was necessary to carry out a very detailed studv of
* " Nature," No. 2441, Vol. 97, Aug. 10th, 1916, p. 489.
t " Nature," No. 2488. Vol. 90, July 8th, 1917, p..373.
574 MESOZOIC INSECTS OF QUEENSLAND, iv.,
the fossil before anything could be published. Here I was
helped and guided by a regular correspondence with Mr. Miller.
He, like other entomologists, found in the supposed corneous
border of the wing a stumbling-block to the acceptance of any
theory of Dipterous affinity for the fossil. He also pointed out
the remarkable difference between the structure of the costal
vein in Dunstania and in Exsul, and finally declared his belief
in the Homopterous nature of the fossil.
The only other possibility that occurred to me was, that Dun-
siania might be the wing of a large Oligoneurous Mayfly. The
shape and venation of the wing made this likely; but I was
unable to follow this line of research far, as I soon became con-
vinced that the evident toughness and strength of build of the
Dunstania wing could not possibly have belonged to the wings
of any representative of the Order Plectoptera.
Here, then, after having reviewed possible affinities with four
Orders, Lepidoptera, Diptera, Homoptera and Plectoptera, I
abandoned the study of this puzzling fossil for a time, in the
hope that a respite from the work might rid my mind of any
preconceived notions, that were bound by now to be present,
after so long a time spent in thinking over the problem, and
misfht allow some new lierht to come in.
In the meanwhile, I had received from Mr. Dunstan the com-
plete series of fossils found at Ipswich since my firbt paper had
been published, and had been arranging these for study. For
some months, I had put the new material of Dunstania by, as it
was evidently in very poor preservation compared with the type,
and I did not expect to get any new evidence from it. It was,
however, from this new material that the solution of the problem
came; and that solution was of such a nature that it could have
scarcely been suggested as a possibility, on the evidence of the
type alone.
The number of the type-specimen was 2a; the two wings of
Dunstania received after the type had been described were
numbered 107a and 147 respectively. In the present paper, I
propose to give a complete description of the new material,
together with a re-examination of the type in the light of the
BY R. J. TILLYARD. 575
new evidence, and an attempt at a complete restoration of the
wing.
Specimen No. 147 is in a very poor state of preservation, and
represents practically the same area of the wing as is found in
the type. It clearly belongs to a somewhat larger insect than
the type; and, as the venation differs from that of the latter in
some important respects, it will form the type of a new genus
within the family Duiistaniidce.
Specimen No.l07« is in much better preservation than No.
147, but it is not so well preserved as the type. The apical part
of the wing is missing, but the whole of what was taken to be
the basal part in the type is fairly well preserved. On the other
side of this there is preserved a large flattened area, with only
traces of disconnected venation. On making a careful drawing
of this, and trying to establish its connection with the rest of
the wing, on which the venation is well preserved, it became at
once apparent that this new fossil represented almost the whole
{excluding the base and apex only) of a huge Heteropterous
hemelytron, and that the supposed almost complete wings pre-
served in the type-specimen and in No. 147 were, in reality,
nothing more than the distal membranous half of the hemelytron,
which is separated anteriorly from the basal half, or coriwm, by
a strongly developed node upon the costa.
With this new and unexpected light upon the problem that
had been exercising me for so long, the mystery of the affinities
of DuiisUmia became cleared up in a moment. There cannot
now be any doubt that these fossils are Heteropterous; the only
question that remains for discussion is as to what, if any, recent
or fossil families of Heteroptera show any affinity with them.
Before the fossils, including the type, can be correctly de-
scribed in terms suitable to their new position, it will be neces-
sary to examine them very carefully, with a view to trying to
establish the homologies of their unique venation upon the
Comstock-Needham system. This is a difficult task. I propose,
therefore, to use the new names under which I shall describe the
two new fossils at the end of this paper, in order to facilitate
the discussion that must take place before those descriptions can
be properly given.
576
MESOZOIC INSECTS OF QUEENSLAND,
Specimen No. 107a is to be made tlie type of a new genus and
species, Dunstaniopsis triassica, n.g. et sp., while specimen No.
147 will also be the type of a new genus and species, Paradun-
stania affinis, n.g, et sp. All these are evidently members of
the same family, Bunstaniidff, the venational differences being,
at the most, only of generic value.
Tn discussing the venation of the Bunsfaniidce, we shall make
use of the most complete specimen, Duiistaniopsin triassica, n.g.
et sp., supplemented by the type of the family, Dnii^tanui
pidchra Till. It will be necessary to compare the venation on
the corium of the former with the tracheation to be found on the
same area of the hemelytta of recent Heteroptera. Unfortu-
nately, few reliable figures of the latter are available. I have
selected as the most suitable for our purpose the excellent figure
of the tracheation of the nymphal fore wing of Syromastes {idixnWy
Coreidcp) given by Handlirsch (1, Atlas, PI. vii., fig. 3). It is
not necessary to figure the corresponding venation in the imago,
as it onl}' differs from that of the nymph in the greater differ-
entiation Ijetween corium and membrane^ the dividing line
becoming a complete whole, and effectually concealing the con-
nections between the veins on the corium and those on the mem-
brane. Consequently, such an archaic type as Dunstaniopsis is
more closely comparable with the tracheation of the nymphal
wing of Syromastes than with the venation of the imaginal wing.
Text-fig. 17.
Tracheation of forewing of iiympli of Syromastes sp. ; ( x 8). After Hand-
lirsch. For lettering, see p. 592,
Text-tig. 1 7 shows the tracheation of the nymphal wing in
Syromastes. The wing consists of a hard coriaceous basal half,
BY R. J. TILLYARD.
577
called the corfwm (co), and a softer membranous distal half, called
the meAnhrane (ni). These are separated by the dividing line,
running somewhat obliquely across the wing from the node (n)
on the costa to the antinode (n) on the dorsum. Posterior to
the corium is a narrow projecting anal area, known as the cl(wiis
(cl). Around the membrane is a complete coriaceous harder (e.b.),
such as we have already noticed very prominentlv in the tvpe-
specimen of Dnnst<iiiia.
Text-fig. 18.
Hemelytron of Dunstaniopsis triamica, n.g, et sp.; { x2*S). Upper
Triassie, Ipswich, Q. For lettering, see p. 592.
Turning next to Dunstaniopsis (Text-fig. 1 8) we see that the
node is more prominent, but that the dividing line is, on the
whole, not so clearlv marked, and its course is far more irregular.
This line also runs more transversely across the wing in Dun-
staniopsis than in Syromastes-, but, in many recent Heteroptera,
especially in the family Pentatomidce, it takes a similar trans-
verse course, though much straighter than m Dutistaniopsis. A
further point of difference is that no true claval area can be seen
in Dunstaniopsis. But, as the extreme base of the wing is not
preserved, there may well have been a short elavus present on
the lost part.
Turning next to the venation, we notice that, in Syromastes,
the corium is crossed by four longitudinal veins, not counting
the analis, which borders the elavus anteriorly. By reference
44
578 MESOZOIC INSECTS OF QUEENSLAND, iv.,
to the precedent traeheation, these veins are shown to be the
subcosta, radius, media, and cubitus respectively. As in most
Heteroptera, the subcostal trachea lies close up to the costal
border; in the imago, the subcostal vein becomes more or less
fused with this border. The radius and media are fused for
some distance from their bases, and then diverge gradually at an
acute angle. The cubitus is a separate vein, lying about half-
way between the media and analis.
Having made the above comparison with Syromastes, it now
becomes an easy task to name the radius, media, and cubitus in
the corium of Dunstaniopsis. This is shown in Text-fig. 18.
Owing, however, to the bad state of preservation, in this fossil,
of the dividing line between corium and membrane, we cannot
proceed to name the numerous veins in the latter area with any
certainty. Here, however, we may fall back upon the type-
specimen of Du7istania, in which there is just enough of the
corium preserved to enable us to make the necessary connections.
In my original description of Dunstania, I paid little atten-
tion to this part of the wing, believing it to represent only a
small flattened-out portion of the thorax of the insect. I now
give, in Text-fig. 19, a careful drawing of the region of the node
and dividing line in this fossil. From the figure of Dunstani-
opsis, it will be seen that Pv and M, after diverging from one
another in the middle of the corium, again begin to converge,
and are then unfortunately lost in a break in the rock-surface.
In Dunstania, however, we are able to pick up these two veins
as they approach the dividing line. R is seen to be just dividing
into Rj and Rs, while M converges towards the latter as a stout
vein from below. Sc is fused with the costa, as in Dunstaniopsis.
Between Sc and R there are a number of oblique ribs, half fused
into the corium; these can also be seen in Dunstaniopsis, further
basad along the same area.
Just before the level of the node, Rs turns upwards, and
partially fuses with R^. From the combined stem thus formed,
three branches pass out into the membrane, viz., Rj, R.^, and R3.5.
Of these, Rj is a short vein lying close alongside the costal
jnargin, and supporting it from below; it ends up at about one-
BY R. J. TILLVAIID.
579
third of the length of the membrane, upon the costal margin.
The second branch, R,, is a longer, gracefully curved vein,
running below Rj and parallel to the costal margin for most of
[f*+lS(J ^^^ length, and finally
curving up to meet the
margin at about two-
thirds of the length of
the membrane. The last
branch, Rs.s, gives off R.,
close to its base, and im-
mediately after divides
into R4 and R.,. Ro passes
in a gentle curve beneath
Ro, and ends up just
above the apex of the
wing. Rj and Rg meet
again (in Dimstania OT\\y)
beyond the middle of the
membrane, and from there
on continue straight to
the termen as a single
vein R4+5. In Dunstan-
Rii^ is a sincrle unbroken vein from its origin to its
Text-fig. 19.-
lOpSlS, iV4^5
termination.
Returning to the corium of Dunstania (Text-fig. 19 and Plate
lix., fig. 15), we can pick up M converging towards R as it ap-
proaches the membrane. On the dividing line, it is connected
with R by a short, stout cross-vein, forming portion of the
dividing line itself. From the swollen root of M upon the
dividing line, two branches pass out into the membrane. Of
these, the upper, Mj, arches gently upwards, so as to approach
close to R3.5, and is connected with R5, just distad from its origin,
by a short cross-vein. Thence onward, Mj runs with a slight
double-curving to the termen. The lower branch, M2+3, almost
" Region of node and dividing line in hemelytron of DunManla pnlchra
Till., showing the small anterior portion of the coriuin preserved in the
fossil; ( X 7'«3). Upper Triassic, Ipswich, Q.
580 MESOZOIC INSKCTS OF QUEENSLAND, iv.,
immediately divides into two; of these, the upper, Mg, runs to
tlie termen on a similar course to that of Mj, but well below it;
while the lower branch, M3, diverges from M2 for some distance,
until it gives off a backwardly descending vein, whose destina-
tion cannot l)e followed out in Dunstania\ at this point, M3 is
bent at a very oblique angle, and passes on probably to near the
tornus of the wing.
From the base of M3, in Vutisfania, a very weakly indicated
vein, M4, can be made out, passing downwards, about midway
between Mg and the dividing line; it can be seen to be con-
verging towards the cross-vein mentioned in the preceding para-
graph; but the fracture of the rock hides its further course from
our view.
In order to follow out the further structure of the w'ing, pos-
teriad to the region of M, we must now turn again to Duiistati-
iopsis (Text-fig. 18), in which mast of this part of tlie wing is
fairly well preserved. In this genus, we can easily pick up the
cubitus, running with a gentle curve upon the corium, well below
M. In Syroinastes^ we see that M and Cu are connected by a
short cross-vein, not far from the dividing line. In Dunstan-
iopsis^ Cu gives off an oblique anterior branch, which approaches
closely to two oblique posterior branches given off' by M at about
the same level; these two branches of M unite close to the
dividing line, at a point where the branch from Cu lies very close
below them. Here there is a break in the rock-surface; but we
can pick up, on the other side of the break, a single weak vein
curving into M4. This is evidently the prolongation of the
united branches from M and Cu.
Turning now to the main stem of Cu, we can follow it, in
Dunstaniopsis^ up to the dividing line between corium and
membrane, where it gives off an oblique anterior branch, Cuj^;
this branch meets M4, the weak vein already mentioned as de-
scending from the base of Mo. At the angle formed by these
two, which is almost a right angle in Dunstaniopsis, a third vein
comes in from the membrane; following this distad, we see that
it arises from M3, which, in this genus, is a straight vein running
to near the toruus of the wing.
nv R. j. TILLYARt). 58l
We can now complete this part of the wing in Dunstania.
The descending vein from M3 must be the homologue of the
corresponding vein in Dunstaniopsis ; the only difference is in
the angle of departure from M3. On this difference, as well as
upon its more numerous and slightly more divergent veins, we
see at once that D unsiania had a broader and shorter membrane
than Dunstaniopsis.
Following the main stem of Cu, which I call Cun,, across the
dividing line in Dunstaniopsis, we see that it continues in a
gentle curve, until it ends up on the dorsum of the wing, not far
from the tornus. Below Cu^,, another branch of Cu, Cu„, arises
on the dividing line, and passes into the membrane as a curved
vein running to the dorsum below Cuj.
The dividing line hereabouts is not well preserved, but its
ending up on the dorsum can be just made out. The division
between corium and membrane is clearly noticeable in the fossil
on account of the difference between the colour of the rock and
the fossil itself. The former is a dark grey; the latter a pale
brown, 'i'he vvhole of the corium is coloured pale brown, and is
Hat and smooth. The membrane, on the other hand, is vari-
coloured, showing pale veins bordered by narrow bands of dark
brown pigment, and, between these, areas of the darker rock-
colour on those parts where no veins are present. Also, the
membrane is lightly pitted all over, whereas the corium is quite
smooth in this genus.
Sufficient will now have been said to make it abundantly clear
that we have, in Dunstania and its allied genera, a primitive
type of Heteropteron, of large size, and very probably of amphi-
biotic habits. From recent Heteroptera, such as the Lyyceidce
and PyrrJiocorid(e^ in which some of the venation is still pre-
served upon the corium, the DunstaniidcB differ in the incom-
pleteness and tortuosity of the dividing line between corium and
membrane: and they show us how the anterior portion of this
dividing line has been evolved, apparently in a most complicated
manner, from a series of cross-veins and branches of main-veins
linking up at different angles. Tiie process by which the straight
division between corium and membrane, found in most recent
582 MESOZOIC INSECTS Of QUEENSLAND, iv.,
Heteroptera, has been evolved from the stage seen in the Dun
st'aniidce, is simply one of reduction and alignment of parts.
When it is completed, as in recent Peiitatomidce, the clue to the
venation of the membrane would be quite lost, were it not for
the preservation of the tracheae in the nymphal wing. I have
not been able to find a single existing type of Heteropteron in
which this venation is at all closely comparable with that of
Dunstania. In the aquatic forms, especially, there does not
seem to be any clue at all. Judging chiefly by the shape of the
wing, and the condition of the venation of the corium, I am
inclined to regard the Coreidce, Lygceidce, and Pyrrhoeoridce as
the nearest relatives of the Dunstaniidtfi at the present day.
These three families, and no doubt also the Peniatomidce, may
well lie almost in a direct line of descent from the Dunstaniidce.
We should have to suppose, in restoring the latter, that the
development of the scutellum had advanced very little beyond a
normal size; and consequently, that the clavus, which, in the
position of rest, borders the scutellum, was also very short. As
no anal vein or clavus can be seen in the preserved portion of
the hemelytron of Bunstaniopsis, this was almost certainly the
case.
In Plate lix., fig. 15, I offer a restoration of the complete
hemelytron of Dunstania pulchr a TiW., based upon the preserved
portions of the type and of Bunstaniopsis triassica. The poorly
preserved Paradunstania ajfinis has not been brought into the
discussion at all, and its only use in the restoration is to indicate
the course of the wing-border below the tornus.
It now remains for me to revise my original definition of the
family Bunstaniidce in the light of our new knowledge, and to
give definitions of the type-genus, and of the two new genera
here proposed, and descriptions of the new species.
Order HEMIPTERA.
Suborder Heteroptera.
Family DUNSTANIID^.
Large bugs, with hemelytra more than 30 mm. long. Hemely-
tron suboval, about half as wide as long, the corium either
smooth or lightly pitted (hairy), the membrane pitted (hairyj.
BY ft. J. TILLY ARD. 583
Veins present on the coriiim are R, M, and Cii; no clavus visible
in the preserved portion, but the base of the wing is incomplete
posteriorly in all known specimens. 8c fused with costal margin.
Costa broken between corium and membrane by a strongly de-
veloped node. Dividing line between corium and membrane
crosses the wing from node to antinode in a very irregular
manner, being formed anteriorly by short cross-veins and por-
tions of main veins, posteriorly by a sinuous line not made up
of veins. On the membrane there are either eight or nine
longitudinal veins, of which either three or four arise from R,
three from M, and two from Cu; these veins runs distad in a
slightly diverging manner, Rg ending up at or near the apex, and
Cui at or near the torn us. From M3 a downward and back-
wardly running branch passes to the dividing line, where it
meets the intersection of M4 and Cui^. Few cross-veins present
on membrane. Veins of the membrane pale, generally bordered
on either side by a band of dark brown pigment. A fairly broad
border without pits, and probably coriaceous, runs completely
round the membrane.
Horizon, Upper Trias of Ipswich, Queensland.
The genera comprised in this family are Danstania Till.,
Dunstaniopsis, n.g., and Fai^adunatania, n.g.
Key to the known genera of Dunstaniidc'.
f'S] 1 connected with the lowest branch of R by a short cross-
I vein, not far distad from the dividing line 1 .
I No cross-vein present at this level between Mi and lowest
I Ijrancli of R PavoAlunstania, n.g.
I ''Four branches of R crossing the membrane, viz., Rn to R5 ; R4
and R5 unite a little beyond half-way, forming a closed cell
Dunstania Till .
Tluee branches of R crossing the membrane, viz, Ro, R.t, and
R4+5; the last a single straight vein connected with M, near
its origin Dunstaniopsis, n. g.
Genus Dunstania Tillyard.
(Plate lix., fig.15, and Text-fig.l9).
Membrane and a very small portion only of the corium pre-
served. Preserved portion of corium distinctly pitted all over.
Membrane about as broad as long, the apex prominent but grace-
584 MESOZOIC INSECTS OF QUEENSLAND, iv.,
fully rounded. Node exceediDgly prominent. Rj a weak vein
fused with costal margin from below node up to about one-tbird
the length of the membrane. Kg ending well before the apex,
R3 at tlie apex itself. Between K and M, at the base of the
membrane, a closed cell is formed, bounded by the dividing line
basally, by R..^ anteriorly, by Mj posteriorly, and by a short
cross-vein connecting these two veins distally. Distally from
this cell a second closed cell is formed between R^ and R5, these
two veins at first diverging, and then converging to meet again
at a point beyond the middle of the membrane. M, and M.^
united distally for a short distance upon the membrane. The
backwardly projecting branch from M3 comes off almost at right
angles from the main stem. (Rest of hemelytron missing).
Genotype, Dunstania pulchra Till., (3, p. 32, and Plate iii.,
fig. 6).
Genus D u N s t a x i o p s i s, n.g. (Text-fig. 18).
Hemelytron completely preserved except for the extreme l)ase
of the corium (including the clavus, if present) and the apical
portion of the membrane. A longer wing than that of Dun-
stania^ and narrower in proportion. Differs from Dunstania as
follows: — Corium smooth, membrane lightly pitted all over.
Coriaceous border narrower and less defined; node somewhat less
prominent. R4-1-5 a single straight vein, and hence no closed
cell formed between R4 and R5. The basal cell completed by
the presence of the short cross vein betw^een M^ and lowest
branch of R is present, but is much shorter than in Dunstania.
The backwardly projecting branch from M3 to the junction of
Mj and Cuj^^ comes off from the main stem at an angle of about
30^, and runs almost parallel to the posterior border below it.
On the corium, M curves upward distally, converging towards
R, and giving off posteriorly two branches which unite further
distad and then meet an anterior branch given off from Cu; this
latter vein divides at the dividing line into three branches, CU)^,
Cujb, and Cun, of which the first runs to the intersection of M4
and the backward branch fi (jm M3. Cu^, ends up at or near the
tornus.
Genotype, Duiiatniiioijsis triassica^ n.sp.
fiY il. J. TiLLYARl). 585
DuNSTANioPSis TRiAssiCA, n.sp. (Text-fig. 1 8).
Greatest length of fragment, 31 "5 mm.; breadth across the
dividing line, loo mm Esiiniated total length of hemelytron,
40 mm.
'J'he specimen is only moderately preserved, and would appear
to have been much torn distally before it became fossilised, since
a part of a frond of Thinnfeldia, lying upon the same rock-
surface, projects well into the gap where the missing distal por-
tion ought to Ije, and is embraced on one side by the projecting
torn costa, without appearing to overlie the wing in any way.
Also the beautiful pattern of brown pigmentation bordering the
veins of the membrane, though it can be seen to be present, is
not by any means so well preserved as in Dutistmda. The
corium appears to have been hard, and is fairly well preserved;
the veins lying upon it are not so distinct as those upon the
membrane.
The venational characters being considered as of either family
or generic importance, they will be found in full in the de-
T V p e. Specimen No. 107rt, in the Queensland Geological
Survey Collection.
Genus P A R A D u N s T A X I A, n.g. (Text-fig. 20).
Portion of membrane only preserved, and in very poor condi-
tion. Apparently a hemelytron of somewhat larger size than
that of Duustaiiia. No cross-veins basally between Mj and
lowest branch of R. Cross-veins developed not far from dividing
line between Rj and Ro, atjd also between R;; and K4 .-5. This
last vein receives a curved branch-vein from above, at about
the same point as where, in Diiiistania, R^ and P., unite distally;
but a break in the rock-surface prevents us from determining
whether this branch is really R4 or not. At about the same
level, Mj receives a curving branch from above; this branch ap-
pears to come from R44.5, but its origin is not determinable with
certainty. Rest of venation similar to that of Dunstania.
Genotype, Faradunsiania afjittis, n.sp.
586
MES0Z(JIC INSECTS OP QUEENSLAND, iv.
Paradunstania affinis, 11. sp. (Text fig. 20).
Greatest length of fragment, 17 mm. ; greatest breadth,
14-5 mm. A very poorly preserved specimen, with very little
trace of the brown pig-
mentation bordering the
veins. A large and deep
triangular break has cut
out a considerable por-
tion of the courses of all
the branches of R; the
rest of the veins present
can be traced out by the
use of careful lighting.
This specimen appears
to represent a hem ely-
tron intermediate in Text-fig. 20.'^'
size between that of Duni<fania pidchva Till., and the much
larger hemelytron of Dunstaniopsis triassica, n.g. et sp. In
shape, also, it was probably intermediate between these two.
Type, Specimen No. 147, in the Queensland Geological Survey
Collection.
JVote on the Origin of the Heteroptera.
In dealing with the phylogeny of the Order Hemiptera, Hand-
lirsch(l, pp. 1244-1 249) rightly insists upon the clear-cut dicho-
tomy between the two Suborders Heteroptera and Homoptera.
Neither of these two Suborders can be derived from the other:
for, on the one hand, the Homoptera have preserved the more
primitive wing-form and venation, while, on the other, the
Heteroptera have preserved the more archaic form of the head
and antenniB. As the direct ancestors of both these recent Sub-
orders, Handlirsch indicates some known Hemipterous fossils of
the Upper Permian and Lias of the Northern Hemisphere, which
he places in a distinct Order, Paleeohemiptera, on the ground
that it is not possible to demonstrate that they are definitely
either Heteropterous or Homopterous. The Palseohemipteia he
Paradiimtaula njini^, n.g. et sp. ; ( x2'8). Upper Triassic, Ipswich, Q.
BY R. .). TILLYARD. 587
would, in turn, derive from the Lower Permian fossil genus
Eugereon, wliich he places in a third Order, Protohemiptera.
Had it not been for the preservation of its mouth-parts, which
form a primitive elongated sucking beak, Euyereon would never
have been considered to have any relationship whatever with
the Hemiptera. It was a huge insect, with primitive densely
veined wings that were held out away from the body; the pro-
thorax had lateral expansions suggestive of rudimentary wings,
as in the case of some of the Carboniferous fossils. Apart from
the dense venation, it is not possible to establish any close re-
lationship between the venational plan of Euyereon and that of
primitive Hemiptera; although it might be admitted that such
a connection may possibly have actually taken place in the
course of evolution, between Eugereon and the most densely
veined types of Fulgoridcn^ if the venation of the former had
undergone a great deal of reduction and fusion of parts. That
being so, it seems to me that the Protohemiptera are best re-
garded as a separate Order lying right off the main line of descent
of the Hemiptera proper; the connection between the two being
only establishable through a lost Carboniferous ancestor common
to both.
The Palseohemiptera, on the other hand, are so obviously
Hemiptera, that there seems to be no point at all in erecting a
separate Order lo contain them. Most of the Liassic fossils
known are, in fact, definitely placed within tiie Homoptera by
Handlirsch; and it is quite evident that they rightly belong
there. Only the genus Dysmorpho}>tila Brodie, is retained within
the Palseohemiptera; and, as this is probably a fragment, we
may disregard it. Turning to the Permian fossils, we find two
beautifully preserved wings, Scytinojytera kokeni Handl., and
Prosbole hirsuta Koken, which are placed together in the Order
Palpeohemiptera. The former, I would regard definitely as an
Homopteron; the latter most certainly comlnnes the wing-char-
acters of both Heteroptera and Homoptera, and could not well
be placed in either Suborder without disregarding its evident
affinity to the other. I propose, therefore, to sink Handlirsch's
Order Palseohemiptera to the status of a Suborder within the
588 Mesozoic inskcts oP Queensland, iv.,
Order Hemiptera, this Suborder representing the original main
stem of the Order, while the Heteroptera and Homoptera repre-
sent two distinct branches evolved from it.
Let us now compare Proshole (Text-tig.21) witli the restored
wing of Dunsfania, with a view to the establishment of a more
complete phylogeny of the Heteroptei-a.
n
Text-fig.21.
Proshoh hirsuta Koken, hemelytron, after Handlirsch; ( x2"l).
Upper Permian, Kama River, Russia; n, node.
We notice, at once, that Dunstania is much more definitely
Heteropterous, in that it shows the distinction between corium
and membrane much more plainly than Proshole does. More-
over, the corium of Proshole is strongly pitted, while the mem-
brane is smooth. In Dunstania, both are strongly pitted, while,
in Dunstaniopsis, the corium is smooth and the membrane
pitted. If, then, these genera had a common ancestor, it must
have had a wing pitted all over, such as we find still preserved
in many Homoptera. In most recent forms, the pits are swollen
into tubercles, and no longer carry hairs; but there can be little
doubt, from the nature of the pits in Dunstania, that, originally
at any rate, they were the bases of insertion of macrotrichia.
As we have already seen in the Protomecoptera, the macrotrichia
were originally carried upon a dense meshwork of veinlets all
over the wing. If, then, the wings of the Protohemiptera were
also hairy, we are led to suggest that the true Hemiptera
BY R. J. TILLYAKD. 589
diverged from them by a reduction of tlie original meshwork, in
the course of which the pits, or bases of insertion for the maci-o-
trichia, became seated upon the membrane of tlie wing; just as,
in the Lepidoptera, the scales have appeared in a similar manner.
The original pitted wing of the true Hemiptera may be con-
sidered to have given origin directly to the various wing-types
still extant amongst tlie older families of recent Homoptera; the
main developments in this latter Suborder are not to be found in
the wing at all, but in the specialisation of the head and antennae,
and in the development of the power of leaping. In this con-
nection, we should bear in mind that a number of Homoptera,
especially in the Cicadidat, still show the dividing line between
corium and membrane.
We have now to consider the origin of the Heteroptera. For
this, we cannot take Prosbole itself as a starting point; but we
must go back a little way beyond this type, and assume a closely
similar ancestral form with a wing pitted all over. Keeping the
main scheine of venation unaltered, we may now see the origin
of the Heteroptera in a form in which tlie tendency of evolution
in the basal half of the wing was towards suppression of the
venation and hardening of tlie wing-membrane (with or without
loss of the pits); while, in the distal half of the wing, the cross-
veins of dense Fulgorid-like venation still preserved in Prosbole
become obsolete, and the remaining venation becomes greatly
altered, in correlation with the differentiation of the dividing-
line between corium and membrane.
I think that a very little consideration, aided l)y a comparison
of the figures of Prosbole smd Dunsiania here given, must con-
vince us that Dunstania represents a very typical immediate
derivative from the hypothetical ancestor oi Prosbole, from which
we started in the preceding paragraph, along the lines there
indicated as leading directly to the Heteropterous type. That
is to say, Dunstania, though not directly derivable from Pros-
bole, is nevertheless closely related to it; but, while Prosbole
itself cannot be accepted as having advanced along the line of
the true Heteroptera, and must, therefore, be kept in a separate
Suborder PaUeohemiptera, Dunntdnia, on the other hand, has
590 MESOZOIC INSECTS OF QUEENSLAND, iv.,
advanced far enougli to be considered a definite Heteropteron
of a primitive type.
Let lis now consider the very interesting question of the
evolution of the dividing line between corium and membrane in
the Suborder Heteroptera. Dunstania shows us this h'ne in an
exceedingly primitive condition. From it, we see that its
anterior portion was originally composed of either branches of
main veins, or cross-veins. But, as the main veins are all pro-
ceeding more or less distad, it is, therefore, inevitable that these
various parts should meet at difterent angles, as we see so de
finitely shown in Dunstania.
The parts of the dividing line may be shown as follows, start-
ing from the node or costal end (see Plate lix., fig. 15, and Text-
figs. 18-1 9):—
A. Anterior Division, formed from cross-veins and branches
of main veins :
1. The node, formed from the fused C and Sc.
2. Radial portion, formed from short parts of Rj and R.^,
partially fused together.
3. A short cross-vein connecting the radial portion with the
median portion.
4. Median portion, consisting of a swollen root on M, from
which the three branches of M are given off.
B. Posterior Division, consisting of a sinuous line crossing the
wing more or less transver.sely, and not carrying either cross-
veins or parts of main veins :
5. Medio-cubital portion, from the swollen root of M to Cu;
the curvature of this part is concave to the corium.
6. Cubito-anal portion, from Cu to the antinode; the curva-
ture of this part is slightly convex to the corium.
In the evolution of such a type as a recent Lygseid or Penta-
tomid Bug, all these diverse parts have to become aligned and
fused into one harmonious whole. Probably a careful study of
existing types amongst the Heteroptera would reveal many traces
of the method by which this change has been brought about.
The other point of outstanding interest in the evolution of the
Heteropterous hemelytron is the development of the clavus,
•
BY R. J. TILLYARD.
591
which is correlated with the growth of the scutelluni. We are
bound to assume that, in the original Hemiptera, the scutellum
was of more normal size than it now is; and, consequenth% the
wings v/ere not so completely folded over one another distally as
they now are. In fact, we have to start from the normal type
of scutellum and the roof-like position of holding the wings, still
to be found in many Homoptera of the present dav. From this,
a gradual enlargement of the scutellum, together with a close
folding-down of the wings upon one another, leads us to the
condition found in recent Heteroptera. Here the dividino- line
marks the limit of the portions of the wing that are folded upon
one another; while the anal area, or clavus, borders the two pos-
terior sides of the large triangular scutellum. Thus we have to
correlate the length of the clavus with the size of the scutellum,
the tendency being towards enlargement as evolution progresses.
►Since, in Du7istaniopsis, there is no trace of a clavus in the pre-
served part of the wing, and the same is
true of Prosbole, we have to conclude that
both these forms had only a small or moder-
ate-sized scutellum, bordered by a corres-
pondingly short clavus, or even, perhaps,
by scarcely any claval area at all.
As soon as the overlapping of the two
distal parts of the wing, or membranes, has
been brought about, it will be clear that
these two parts will in future act (while
the insect is at rest) as one only; whereas
the coria of the two wings remain separate.
There will, therefore, be little tendency
towards the thickening of the membrane Text-fag. 2i..
at all, and the " heteroptery" might be expected to advance
more quickly than ever. The line of evolution would then
culminate in forms having a thick corium, from which all traces
* Diagram of a Shield-bug, family Pentatomidce, to show the five parts
of the shield, viz., 1, the pronotum, pn; 2, the scutellum, .sc; 'A and 4, the
right and left coria, co; and 5, the two completely' overlapping membranes,
7nh, The clavus, cl, borders the scutellum on either side.
592 MESOZOIC INSECTS OF QUKKXSLAND, iv.
of venation had been eliminated, and a tliin membrane, in which
the veins were arranged more or less parallel to one another and
to the wing-border. Probably the highest point reached by this
line of evolution to-day is to be seen in the dominant family
Pentatomidrt', in which the shield-shaped or " cut-into-five " ap-
pearance of the insect becomes perfected, and is often enhanced
by bizarre sculpture and brilliant colouration. Text-fig. 22 shows
the outline of such an insect, with the five separate areas of the
shield-design named.
BIBLIOGRAPHY.
1. Handlirsch, A., 190S. — Die Fossilen lusekten, etc. Leipzig. {Pahvon-
tinklce, pp. 018-627, and Atlas, Plates xlix.-l. ; Phjdogeny of Hemip-
tera, pp. 1244-1249; Piotoliemiptera, p. 387, and Atlas, Plate xxxvii.,
figs. 21-23, Plate xxxviii., figs. 1-4; Pala?ohemipteia, p. 390, and
Atlas, Plate xxxvii., figs. 24-27.
2. Meyrick, E., 1916. — "Note on Some Fossil Insects."' Knt. Mo. Mag.,
3rd Ser,, No.20, (No.627), Aug. 1916, pp. 180-182.
3. TiLLYARD, R. J., 1916. — " Mesozoic and Tertiaiy Insects of Queensland
and New South Wales." Queensland Cleol. Survey, Publicatioii
No. 253, pp.31 -33, and Plate 3, fig. 6.
EXPLANATION OF PLATE LIX.
Fig. la. — Restoration of right hemelytron (forewing) of Daastania pulchra
Till., based upon the preserved portions of this fossil and of the
allied Dmistaniopsis triassica, n.g. et sp. ; ( x ,■>),
Letterimi of Text-Jlynrefi.
A, analis — C, costa — c.h., coriaceous border — d, clavus — co, coriuni —
Cu, cubitus; Cuia, Cuib, Cu.j, its branches on the membrane^-wi or w?/>,
membrane — M, media; Mj to M4, its branches on the membrane — n, node
— n , antinode — pn, pronotum — R, radius; R^ to Rg, its brandies on the
membrane — Rs, radial sector — sc, scutellum — Se. subcosta.
593
THE GEOLOGY AND PETROLOGY OF THE GllEAT
SERPENTINE BELT OF NEW SOUTH AY ALES.
Part viii. The Extension of the Great Serpentine Belt
FROM THE NUNDLE DISTRICT TO THE CoAST.
By W. N. Benson, B.A., D.Sc, F.G.S., Professor of Geology
AND Mineralogy in the University of Otago, N.Z., late
LiNNEAN MaCLEAY FeLLOW OF THE SOCIETY IN GeOLOGY,
(With one Text-figure).
Tlie previous parts published liave described, in greater or less
detail, the whole of the country along the Serpenthie Belt, ex-
tending from Warialda to I fanging Rock, a distance of nearly
150 miles. Some facts, however, have been collected with regard
to the further extension of the Belt, which may be wortljy of
record before this series of papers is concluded.
Somewhat to the west of the Serpentine-line, and about twelve
miles south of Nundle, commences, on Wombramurra Creek, one
of the largest masses of limestone in the State, which mass con-
tinues for some miles across the Main Divide, at Crawney Pass,
into the head of the Isis River. This has, as yet, been little
studied. It was briefly described by Phillips, in 1875(1). Mr.
Etheridge described a new coral therefrom, in 1898(2); and Mr.
Dun, two years later, determined a large collection of fossils ob-
tained by Mr. Cullen(3). It was visited by the writer in 1910,
and has recently been studied by Mr. Came, and is to be de-
scribed in his forthcoming account of the limestones of New
South Wales (4). Yery little is known of its stratigraphical re-
lationships to the other Devonian rocks; it seems to occur among
a series of banded claystones like those of the upper portion of
the Taraworth Series, or even of the Barraba Series, though, as
45
594 GREAT sp:hpentink belt of new south wales, viii.,
pointed out in previous papers, the lithology of the Tarn worth
Series is not very constant, an(] tlie chertvfacies is more marked
MVJOO
in the Tamworth-Nundle region than elsewhere. The following
fossils were found to be present in this limestone, the letters M.,
L., and N. indicating whether the forms are also present in the
BY W. N. BENSON. 595
Moore Creek, Loomberah, or Nemingha horizons respectively.
Favosites gothlatidica ...
F. basaltica var. '^yiooiibiensis
F. salebrosa
F. multitabulata
A Stromatoporoid
Diphyphyllum porteri ...
D. sp.nov.
* Sanidophyllum davidis
IVyplastna, sp.nov.
* Spongophyllani yiyanteum
*AcHnocystis cornubovis
Cyathophyllum sp.
Cystiphyllwn auslralasiciim
* Microplasma parallehtm
Heliolites po7'osa
*Sy7'ingopo7-a auloporoides
Litophylhiin konincki ...
Alveolites sp. ...
Endophyllum schlueteri.
Atrypa sp.
Euomphalus sp.
The association of tlie species marked with an asterisk, and
the abundance of Heliolites are features very characteristic of
the Moore Creek limestone, with which the great development
of pure grey or white limestone also accords. The absence of
the great abundance of Stromatoporoids, or Pentameroid shells,
or of the signs of shallow water, distinguishes this from the
Loomberah Limestone; and the want of association with frag-
mental igneous rocks, or with banded cherts, separates it from
the Nemingha Limestone. It seems permissible, therefore, to
correlate it with the Moore Creek Limestone.
East from here, across the Main Divide, the (Serpentine Belt
extends across the headwaters of the Manning River, through
extremely rugged and broken country. It is the deeply dissected
remnant of a plateau, over which extends a widespread series of
more or less alkaline basic rocks, which were described by Mr.
M.,
N.
N.
M.,
N.
M?,
L.,
N.
M.,
L.,
N.
M.,
L.,
N.
M.,
N.
(rare).
M.I
, L.], N.I
M.
M.
L.I
L.I
M.
M.,
L.,
N. (rare),
M.
M.,
L.,
N.
M.?
, N,
.?
M.T
, N.
?
M.?
596 GREAT SERPENTINE BELT OF NEW SOUTH WALES, viii.,
Andrews, at the head of the Hunter and Manning Rivers, as
being of two types, "one, a holocrystallinerock with large augite
crystals so abundantly scattered throughout its mass as to obtain
for it locally the name of 'plum-pudding stone.' Other types
found there are dense, fine-grained, vesicular olivine-basa]ts"(5).
The writer has shown that, among these, are to be found
various types of basalt, basanite, dolerite, essexite, teschenite,
and theralite, with sometimes an abundance of nepheline (6, 7).
Indeed, there is an assemblage of basic rocks quite analogous to
those of the Bohemian Mittelgebirge, which Becke described as
a typical instance of an assemblage of Atlantic rocks. The
shapely cone of Wombramurra Peak, a few miles east of the
Pass, was not visited by the writer, but is very analogous to the
conical Gragin Peak near Delungra, Wariakla, which Mr. Carne
believes to be a centre of intrusion, and in which Mr. Card found
olivine-dolerite. Mount Jellore, near Mittagong, 80 miles S.W.
of Sydney, may possibly be of a like nature.
Serpentine occurs on this line at the Barry, at the head of the
Barnard River, twelve miles distant from Nundle, and was here
observed by Mr. Andrews, W'ho remarked on the association with
it of "diorites and basic rock-types," probably dolerites, and
spilites(8). Mr. Longrigan, of the Barry, informed the writer
of the occurrence of limestone east(?) of the serpentine on his
property: and Mr. Stonier records the finding of Favosites in
limestone on the Pigna Barney Ptiver, two miles above its junc-
tion with the Manning (9). Serpentine occurs again at Glenrock,
some twenty-five miles south-east of Nundle (8), and local pros-
pectors say that it is also present at the old " Polly Fogal "
Diggings, which are not marked on any map, but are stated to
be sixty miles east from Scone, and forty south-east of Nundle.
An analysis of chromite obtained from here has been recorded (10).
This must be near the Curracaback River, which is crossed by
the serpentine (8).
North-east from this line, serpentine has been recorded from
Nowendoc(8), doul)tless among rocks of the Eastern Series, and
will thus probably be on or near a line extending from the belt
of serpentine recorded by Stonier (11), which extends from near
fJY W. N. BENSOX. 597
Duiigowan Creek, at a point about six miles south-east of Dun-
go wan township, across to the Mulla Creek, east of Moonbi. At
the south end of this, the writer found antigoritic serpentine,
and various intrusive doleritic rocks. A further nortliern con-
tinuation of this beyond the Moonbi granite is, perhaps, to be
seen in the serpentine which occurs {fide D. A. Porter) near the
liead of Moore Creek, from which a nickelifeious opal was ob-
tained (12).
Further to the south-east, Mr. Siissmilch and the writer(13j
have observed the occurrence of serpentine at Mt. George,
on the Manning River, and have remarked on the presence
of the same rock at Glen Lewis, five miles to the n'orth of
this spot, and at Bow Bow near Tinonee, south of the Manning
River, twelve miles to the south-east of the last-mentioned oc-
currence. This last record, we owe to Mr. Card, who received
specimens therefrom. Between this group of intrusions of ser-
pentine and Gloucester, to the south-west, there is a great de-
velopment of Middle and Upper Devonian and Lower Carboni-
ferous rocks, extremely like those in the areas to the north,
described in the preceding parts of this series. The strike is
generally N .W.-S.E., with a S.W. dip. In the Middle Devonian
beds thei-e is an immense development of spilite with pillow-
structure, rising to form Kangat Mountain. The geology is
further complicated by the presence of an infaulted outlier of
Permo-Carboniferous rocks, perhaps analogous to that recorded
from the Nundle district(14). What exactly is the relationship
of this district to those further to the north, has yet to be dis-
covered. It lies rather too far to the east to be in the direct
continuation of the main Serpentine-line, unless (as is quite pos-
sible) the direction of that line has been bent to the E.iS.E. in
the Manning River valley. In the absence of such a deflection,
it may be suggested that the George Town serpentine is a con-
tinuation of the Mulla-Nowendoc line of intrusions, and that
the southerly continuation of the main Serpentine-line is indi-
cated by the serpentine and chromite, which occurs near Barring-
ton (15). and the serpentine recorded by Mr. Andrews, on the
Myall River(8). If this be so, however, the Carboniferous and
598 GREAT SERPENTINE BELT OF NKW SOUTH WALES, viii.,
Devonian rocks of this region are so remarkably little altered,
for rocks east of the main Serpentine-line, as to indicate some-
what different conditions from those which have prevailed in
the districts north of Nundle. In either case, it is probable
that the Serpentine-line will eventually be traced from Nundle
down to the coast, thus adding another hundred miles to the
hundred and fifty already mapped, and thus forming one of the
most continuous structural lines in the State.
A few words may "be given with respect to the serpentines of
Port Macquarie, which were described by Mr. Carne(16). In
this region, the predominant strike is to the N .N.E. It has been
suggested that the serpentines here lie on the same line as those
of the Great Serpentine Belt, which has swung round from the
S.S.E. to the S.E., and eventually into the N.N.E direction,
and thence swings further round to include the serpentines of
the Clarence River, recorded by Professor David and others(17).
The Serpentine-line, on this hypothesis, forms a discontinuous
ring about the strongly compressed Permo-Carboniferous rocks
and the great granitic masses of north-eastern New South
Wales(18). The writer has pointed out, however, that the pre-
sence of a N.W. to N.N.W. strike so close to the coast as the
Gloucester District renders this suggestion improbable, and in-
dicates rather that the Port Macquarie N.N.E. line of strike is
more likely to be on a virgation passing off from the main N.N.W.
direction. Several such virgatious have been noted between
Binsara and Nundle, notably at Mundowev on the Namoi
Pviver(19), though they have not been traced into a greater
divergence from the main direction than a N.-S. line of strike.
Such an hypothesis, however, involves a much simpler distribution
of folding forces than that necessary to explain the discontinu-
ous ring of intrusions, though the latter would not be entirely
without analogies. A comparison, though an extremely strained
one, might be made with the discontinuous line of serpentine-
intrusions around the Central Granite of the Austrian Tyrol, but
it is very doubtful whether such a comparison would indicate
any real analogy, so diverse are the other features in the two
areas ( see 20) .
BY W. N. HKNSON. 599
BIBLIOGRAPHY.
1. Phillips — Mines and MintMal StatiRtics of New South Wales, IS?-'),
pp. 149-151.
•2. Etheridoe — "On the Occurrence of the Genus Endophyllum in the
Lower Pah«ozoic Rocks of N. S. Wales," Ree. Geol. Snrv. N. S.
Wales, Vol. vi., 1898, pp. 43-46.
3. Dun— Ann. Report Dept. of Mines N. S. Wales, 1900, p. 195.
4. Carne and Jones — " The Limestone Deposits of N. 8. Wales.'' Min-
eral Resources, No. 24.
5. Andrews — "The Tertiary History of New England." Rec. Geol.
Surv. N. S. Wales, Vol. vii., 1903, p. 63.
6. Benson — " Preliminary Note on the Nepheline-bearing Rocks of the
Liverpool and Mount Royal Ranges." Journ. Proc, Roy. Soc.
N. S. Wales, 1911, pp. 176-186.
7. " The Geology and Petrology of the Great Serpentine Belt of
New South Wales." These Proceedings, 1913, pp.593, 701-3.
S. Andrews — " The Geology of the New England Plateau, with special
reference to the Granites. " Parts ii. and iii. Rec. Geol. Suiv.
N. S. Wales, Vol. viii., pp. 108-152, especially 118, 146-7.
9. Stonier— Ann. Report Dept. of Mines N. S. Wales, 1894, p. 150.
10. Analyst's Report. Ann, Report Dept. of Mines N. S. Wales, 1915.
11. Stonier — "The Geology of Swamp Oak and Niangla."' Rec. Geol.
Surv. N. S. Wales, Vol. iii., 1892.
12. Porter — " Note on the Occurrence of a Niekeliferous Opal near Tam-
worth, N.S.W." Journ. Proc. Roy. Soc. N. S. Wales, 1897(1898),
pp. xxviii.-xxxix.
13. Benson and Sussimilch — "The General Geology of the Gloucester
District." Journ. Proc. Roy. Soc. N. S. Wales, 1916, pp. xxxi.-
XXXV.
14. Benson—" The Geology of the Nundle District.'" These Proceedings,
1913, pp.586-7.
15. Analyst's Report. Ann. Report Dept. of Mines, N. S. Wales, 1906.
16. Carne — "Geology of the Coast between Port Macquarie and Cape
Hawke." Rec. Geol. Surv. N. S. Wales, Vol. v,, pp. 53-64.
17. David— Annual Report Dept. of Mines N. S. Wales, 1891, p.208.
18. Handbooks for the Meeting of the British Association, 1914. Austra-
lian Federal Handbook, p.308; N. S. Wales State Handbook, p.619.
19. Benson — "The Geology of the Western Slopes of New England,''
These Proceedings, 1917, p. 223.
20. Becke and Loewl — Exkursionen im westlichen und mittleren Abschnitt
der Hohen Tauern. Livret-Guide s. Congr^s Internationale Gt'o-
logique, Vienna, 1903. Map.
600
THE STEMS OF CLIMBING PLANTS.
By John Shirley, D Sc, and C. A. Lambert.
(Plates Ix.-lxvi.).
In botanical excursions with the Field Naturalists' Club or
Royal Society, it has been a constant source of interest that
climbing plants of the Orders Bignoniacese and Menispermacese
could be partly determined by studying the cut ends of their
stems. This determination extended only to the Order, and not
to the genus or species. In Bignoniacere, the determining factor
was the arrangement of the bast in four masses, or in multiples
of four, each mass usualh' presenting rectangular outlines in
transverse section. In Menispermacese, the rays of wood and
bast have usually a stellate arrangement, the vascular bundles
being separated by broad, multiseriate medullary rays, which, in
transverse sections of the stem, appear club-shaped from dilation
of the last-formed cells.
To test whether similar peculiarities marked the structure of
climbing plants of other Orders, some tifty-three stems were
sectioned, and their slides photographed by my colleague, Mr.
C. A. Lambert.
List oj' Species examined.
AcANTHACEiE. — Thunhevgia yrandijiora Roxb., T. laurifolla
Lindl.
Ampelide^. — Vifis hypoylauca F.v.M., V. opaca F.v.M., V.
sterculifolia F.v.M.
APOCYNACEiE. — Beaiiinontia yratidifiora Wall., Alelodrims acu-
tijiorns F.v.M., yVachelos^jei'mum jasmitioidea Lindl.
AsCLKPEDiACEiE. — Cr'i/])tostegia yra7idiflora R.Br.
BiGNONiACKiE. — Adeitocalymma nitidnm Mart., Bic^iiottiaJJori-
bunda H.B.& K., B picta Lindl., B. Tweediana Lindl., H. venusta
Ker, Tecoma Hillii F.v.M., T . jasminoides Lindl.
BY JOHN SHIRLEY AND C. A. LAMBERT. 601
CoMBRETACEi^. — Qiiisqualis itidica Roxb.
CoMPOSiTiE. — Senecio tanwides DC.
(JoNVOLVULACE.E. —Ipomcea Horsfallice Hook.
CucuRBiTACEiE. — Bryonia laciniosa Linn.
Leguminos^. — Ahrus precatoi-his Linn., Bauhinia scandens
Burni.,( = B. corymhosa Roxb.), Derris scandens Benth., Lathyrus
odoratus Linn., Milletia australis F.v.M., M. megasperma F .\ .M..,
Wistaria chinensis DC.
LoGANiACEiE. — Buddleia niadagascariensis Vahl.
MALPiGHiACEiE. — Banisteria chrysophylla Linn., Riptage Ma-
dahlota Gsertn., Stigmaphyllon jatrophcefolium Juss.
MENiSPERMACEiE. — Carrouia mnUisepalea F.v.M., Legnephora
Moorii Miers, Stepha7iia hernandicefolia Walp.
^ YCT AGINEJE. — Bougainvillea lateritia Hort., B. Sanderiana
V. variegata Hort,, B. spcctahilis Juss.
PASSiFLOREiE. — Fassiflora suberosa Linn.
PiPERACEiE. — Piper nigrum Linn.
y O'LY GO^ A.CY.IE. — Antigonon leptopus Hook.
RuBiACEiE. — C(elosj)ermuin paiiimdatum F.v.M.
SoLANACEiE. — Solauum Wendlandii Hook. f.
URTiCACEiE. — Ficus pumila Linn.
VERBENACEiE. — Faradaya splendida F.v.M., Petrcea volubilis
Linn.
AROiDEiE. — Monstera acicininata C. Koch, Pothos aureus
Linden.
Flagellarie^. — Flagellaria indica Linn.
L1LIACE.E. — Asparagus racemosus Willd., Geitonoplesium cymo-
sum A. Cunn., Ehipogonum albnm R.Br., Sniilax australis R.Br.
ORCHiDACEiE.— Galeola cassythoides Reichb.
The study had not proceeded far. when it was seen that any
grouping in Natural Orders was impossible, as similar character-
istics were common to plants of many different families, especi-
ally among Dicotyledons. It was necessary, therefore, to create
classes, among which the stems could be distributed, and the,
following are submitted.
46
:C
I g R A R YI30I
602 THE STEMS OF CLIMBING PLANTS,
Suljclass i, DICOTYLEDONES.
1. Normales. — The stem shows a single cambium-ring, with
the wood and bast of each bundle lying along the same radius,
the exterior outlime of the bast approximating to a circle.
Type, CirJospermum paniculatum F.v.M.
2. Chiastoxylon. — A single zone of cambium, with anomalous
distribution of tissues in young stems into four rays, each of alter-
nating wood and bast, the bast reaching (a) partly to the pith,
or (h) stopping short of it. T^^pe 2 (<x), Bignonia Tiveediana
Lindl., 2(6), B. jncta Lindl.
3. Astroxylon. — Normal cambium, with definite and continu-
ous separation of fibro-vascular bundles by means of pluriseriate
medullary rays, that present, in transverse section, a stellate
arrangement. Type, Carroiiia multisepaJea F.v.M.
4. Endophloia.— Besides the normal bast, a second develop-
ment of bast occurs at the inner margin of the ring of wood,
forming bicollateral bundles. Type, Bryonia lo.ciniosa Linn.
5. Exoeyela. — Besides the normal cambium-ring, new cam-
bium-zones appear successively in centrifugal order. Type,
Wistaria chinensis DC.
6. Phloioeyela. — New zones of bast produced in each period
of vegetation in centripetal order. Type, Vitis sterculifolia
F.v.M.
7. Polycyela. — The oldest vascular bundles lie in the pith;
outside the pith, a normal zone of wood and bast is formed, or
alternating rings of wood and bast may be formed. Type, Boii-
gainvillea spectabilis Hort.
Subclass ii. MONOCOTYLEDONES.
1. Vulgares. — Possessing the usual rind, hard ground-paren-
chyma, and scattered, closed bundles of the ordinary mono-
cotyledon.
2. AbnOPmales- —Differing from the typical monocotyledonous
stem in one or other of the above peculiarities.
Subclass i. DICOTYLEDONES.
L Normales. — To this section belong Banisteria chrysophylla
L., Ccelosjjermum 2^<^^i'iculatum F.v.M., Cryptostegia grandijiora
R.Br., Faradaya splendida F.v.M., Hiptage Madablota Gsertn.,
BY JOHN SHIRLEY AND C. A. LAMBERT. 603
Milletia australis F.v.M., M. megasperma F.v.M., Se^iecio tamoides
DC, Solanum Wendlandii Hook, f., and 2'ecoiiia jasminoides
Lindl , etc., etc.
(a.) CCELOSPERMUM PANICULATUM F.V.M. (Plate Ix., fig.l). —
Although this stem is normal in having a single ring of cambium,
and in regard to the formation of secondary wood, the pith pre-
sents peculiarities. It is made up of two elements; one, the
ordinary pith-cell, containing starch: and the other, found mainly
in the centre of the pith, a thick-walled, sclerotic cell. The
latter form strands which do not extend to the circumference of
the pith. They are akin to the gum-resin sacs, described by
Karsten,* in the allied genera, Cinchona and Ladenbergia.
(b.) Milletia australis F.v.M. (Plate Ix., fig.2). — Although
normal in the development of its cambium-ring, this stem shows
some special features. The pith is of very large cells, and, in
transverse section, these seem to be radially arranged, forming
conical masses. In the rays, the cells are elongated, and have
not the usual hexagonal outline. The cambium-ring is very
well defined, and, outside the bast, is a sclerenchymatous ring, 6
to 8 cells in depth, with intervals in which are inserted a few,
much larger, thinner- walled cells, evidently a transfusion-tissue.
Sacs containing single crystals abound in the cortex, and others
with much larger single crystals are scattered through the pith.
Milletia megasperma F.v.M. — The arrangement of the pith
is normal. The wood- vessels form radial rows, with large masses
of murally arranged wood-fibres between them. There is a thick
sclerenchymatous ring outside the bast, with transfusion-tissue
at intervals; but, in this species, the thin-walled cells are oval,
and elongated along radii passing through the transfusion-tissue.
A second, much narrower, sclerenchymatous ring separates the
periderm from the cork. The medullary rays differ little, in a
transverse section from the wood-parenchyma.
(c.) Solanum Wendlandii Hook, f., (Plate Ixi., fig. 3). —The
central pith is of large cells. The ring of wood is very regular,
the wood-fibres showing very even ranks, the cells increasing in
size towards their inner boundary, and gradually merging in
'" Karsten, Die medic. Chiiia-rindeu Neu-Grenadas, Ges. Beitr., p. 382.
604 THK STEMS OF CLIMBING PLANTS,
those of the pith. Outside the bast is a ring of sclerenchymatous
cells, that apparently acts as an endoderniis also, The bark
shows all the usual tissues excellently — phelloderm, phellugen,
cork, and epidermis with cutin. Scattered, sclerenchymatous
elements are found also in the circumferential parts of the pith.
•2. Chiastoxylon.
(a.) BiGNONiA TwEEDiANA Lindl., (Plate Ixi., fig 4).— In trans-
verse section, the pith is roughly square in outline, and a rect-
angular ray of bast strikes the centre of each side of the square
at right angles or nearly so. From the inner end of each bast
ray, a small cone of wood (protoxylem) penetrates the pith to
about one-sixth of its diameter. The pith-cells are filled with
starch-grains. The wood is continuous round the pith, but forms
a very narrow connecting band at the base of each bast-ray, near
the protoxylem-masses already mentioned. The vessels occur in
V-shaped bands, which are most definite near the bast. The
wood-parenchyma forms irregular, radiate rows. The bast-rays
project into the cortex at their circumferential limit, and pre-
serve their shape as rectangular masses. Their bast consists of
alternating strips of hard and soft bast, the latter of more rows
of elements. The bast rays are connected by a bast-ring sur-
rounding the four masses of wood, and showing alternate strips
of hard and soft bast, as in the rays. There is an incomplete
circle of sclerenchyma within the phellogen, beyond which are
6-7 tiers of cork-cells bounded by dead bark.
[b.) BiGNONiA PiCTA Lindl., (Plate Ixii., fig.oj. — The bastforms
four square masses of five thin layers of hard bast, alternating
with five broader layers of soft bast. Where these masses occur,
the wood-tissues have about half their usual depth, and press on
the central pith, giving it almost a square outline. The cambium
is thick and definite. It is depressed at each of the bast-patches,
so that the cambium-ring has a crenulated outline. On a circle
outside the bast are a number of roughly elliptical masses of
thick-walled sclerenchyma. A second narrow and almost com-
plete ring of sclerenchyma, bounding the cork-cambium, is
constructed of brick-shaped cells, whose cavities are almost
obliterated.
BY JOHN SHIRI,EY AND C. A. LAiMBERT. 605
3. Astpoxylon. — Type, Carronia multisepalea F.v.M., (Plate
Ixii., fig.6). — The stele consists of a central pith, composed of
very fine polygonal cells in which a few thick-walled ducts are
inserted at irregular intervals. Of wood and bast, there are
about thirty fibro-vascular bundles, each showing six to eight
large vessels, increasing in size, as usual, from the centre out-
wards. Bordering the pith in each, is a cluster of mainly spiral
vessels. Each medullary ray is well defined, and shows five to
eight rows of brick-shaped cells, parting at the circumferential
end to send, right and left, divisions round the separated bundles.
The bast-masses are oval and well defined. They are flanked
outwardly by older bast-tissues, now changed into sclerenchyma,
to be finally cast off with the outer bark. The different layers
of the bark are well defined, and the phelloderm assumes a wavy
outline, parallel to the outward curves of the various bundles.
Lighter in colour than the rest of the stele, the medullary rays
present a stellate pattern on a freshly cut, transverse section of
the stem.
4. Endophloia. — Bryonia laciniosa Linn. .(Plate lxiii.,fig.7).—
The development of the stem is apparently normal, but the
vascular bundles are bicollateral, having bast on both inner and
outer faces. The vessels of the xylem are unusually large for a
young stem in its second year of development. A large pad of
cork is seen on the side of the stem that has flattened by pressure
against the supporting plant. Very little pith is to be noted,
and the cells of the medullary rays are peculiarly large and broad.
There are signs of the production of interfascicular bast from
the sides of vascular bundles.
5. Exoeyela. — Wistaria chinensis DC, (Plate Ixiii., fig.S). —
The medullary rays are exceedingly well defined, usually of 2-6
rows of radially elongated cells. At their outer extremities, the
cells, lying in the bast, are of larger size. The cambium-ring is
also clearly defined, and outside it are 6-8 ranks of thin-walled
bast-cells. The older bast-cells, pushed out towards the circum-
ference, have most of their cavities obliterated by internal thick-
enings. The phellogen-ring is also well marked, and 3-5 layers
of dead cells lie outside the cork. Cortex and pith show sacs
606 THE STEMS OF CLIMBING PLANTS,
containing single, lozenge-shaped, octahedral crystals. The
remains of a former cambium- ring are seen in the wood about
one-third of the distance from pith to cortex. A third cambium-
ring is seen forming in the bast, from which additional rings of
xylem and phloem will be created.
6. Phloioeyela. — ViTis sterculifolia F.v.M., (Plate Ixiv., fig.
9). —The pith appears homogeneous. The medullary rays, 2-8
rows of radially arranged, brick-like cells, contrast clearly with
the murally arranged wood-fibres. The vessels are large, and
evidently capable of division. The medullary rays are as well
defined between the bast-masses as between the wedges of wood.
They increase in size towards the circumference of the transverse
section, and take a clavate outline. The bast is composed of
8-10 rows of thin-walled cells, with a terminal, almost circular
mass of sieve-tubes. Outside the stele, the remains of former
bast-masses, now sclerotic, form a strengthening layer in the
cortex. A few sacs containing raphides may be noted here and
there; others contain single, larger, flattened crystals, and others,
again, store starch-grains.
7. Polyeyela.
(a.) BOUGAINVILLEA SPECTABILIS JuSS., (PI. Ixiv., flg.lO). — The
centre of the transverse section of the stem contains the largest
vascular bundles, scattered through the pith. The xylem and
phloem portions of each bundle are clearly defined, but the
cambium is usually marked by a scar, where it has torn under
the knife of the microtome. Outside these older and scattered
bundles, is a clearly defined ring of two years' growth of younger
bundles, normally dicotyledonous in general appearance, but
having alternate circles of wood, bast, wood, bast. J^etween any
two bundles of the outer ring are well defined medullary rays;
but, at the interior end of each ray, the cells composing it become
roughly polygonal, and merge into the pith-cells. When a new
cambium appears, outside the last ring of bundles, the cells of
the outer ends of the rays unite round the outer extremity of
each bundle, so as to isolate the ring previously formed b}^ a
definite wavy circle of parenchymatous tissue. The bast arising
BV JOHN SHIRLEY AND C. A. LAMBERT. 607
from this latest-formed cam])ium consists of 6-8 rows of lii-ick-
like, tbin-walled cells, outside of wliich is a single-rowed circle
of sclerencb\'ma.
(b.) BouGAiNViLLEA LATERITIA Hort., (Plate Ixv., fig. 1 1 ), differs
little from Z>. spectabilis and B. tSatideriana, except that numer-
ous sacs containing clinorhombic crystals — raphides — are found
in the periderm and pith. Where a bud is formed on the stem,
the tissues entering the bud from the stele are full of these
needle-crystal sacs.
(c.) BouGAiNViLLEA Sanderiana Hort., (Plate Ixv., fig.l2).—
This section ( x 90) shows the central and first-formed vascular
bundles isolated in the pith, as in the case of its allies; and four
annual zones of alternating wood, bast, and encircling medullary
ray-tissue. This last might be taken for cambium, but is quite
distinct in the character of the cells composing it, from that of
the last-formed cambium-ring. Periderm, cork, and the dead
cells of the outer bark are well shown.
(d.) Piper nigrum Linn., (PI. Ixvi., fig. 13). — The general struc-
ture of this stem is similar to that of Bougainvillea. The first-
formed set of bundles, seven in number, develops cambium within
each bundle, but not between them. These bundles are further
isolated by the growth of the pith. Circumferential ly, other
cambium-tissues form, making a complete wavy ring, and give
rise to a normally developed set of vascular bundles of the usual
dicotyledonous arrangement. Lying in the pith, outside each
of the original isolated bundles, is a resin-duct; and a larger one
occupies the centre of the stem. Bounding the outer, normal
ring of vascular bundles on the inner side, is a complete wavy
ring of sclerenchyma, of about six irregular rows of thickly
strengthened, polygonal cells. An indefinite ring of sclerenchyma,
its cells elongated circumferentiall}', surrounds the outer ring of
wood and bast.
Stephania HERNANDiiEFOLiA Walp. — In a first year's shoot,
there is a remarkable similarity between the cells of the pith,
medullary rays, and periderm. As in Piper nigrum, the stele is
bounded by a scolloped line, outside of which new cambiums
arise, and new ringS of wood and bast are developed.
608 THE STEMS OF CLIMBING PLANTS,
ii. MONOCOTYLEDONES.
1. Vulgares, — To this section belong Asparagus racemosus
Willd., Flayellaria indica Linn., Geitonoplesium cymosum A.
Cunn., Rhipogonum album R.Br., and Sviilax australis R.Br , tfec.
2. Abnormales. — Galeola cassythoides Reichb. f., (PL Ixvi.,
fig. 14). — In this leafless, climbing orchid, the bundles are numer-
ous, and contain six large vessels, with a few smaller marking
the protoxylem. In each vascular bundle there is a limited
quantity of soft bast, and a few sieve-tubes. The ground-paren-
chyma is formed of larger cells than usual, and these are massed
round the bundles, forming bundle-sheaths. In transverse sec-
tions of the stem, a roughly radiate arrangement of the cells of
the ground-parenchyma is evident, in some sections showing four
rays, in others with more and irregular rays. Numerous sacs
containing raphides are seen in the ground-tissue. The contents
of cells lying immediately within the bark show a brown dis-
coloration; these are evidently attacked by a parasitic fungus,
whose hyphte can be seen within the brown cells.
Conclusion. — The whole of the abnormal stem-structures in
climbing plants have, for their object, the free flow of elaborated
sap in the bast-tissues. In Chiastoxylon, the stem so presses on
the support as not to put pressure on the bast-masses ; in
Astroxylon, the broad medullary rays ensure the nutrition of
the stele; in Hiptage, Bryonia, and Solanum Wendlandii, a thick
pad of elastic cork is developed on that side of the stem subject
to pressure, and so on.
Reference letter.^. — h., Bast — c, Cambium — c.c, Cork-cambium — ck.,
Cork — cr., A crj^stal— ec. , Protoxylem— //>. , Fibro-vascular bundle — //.,
Hair — m.r., Medullary ray— ^., Pith^pA., Phelloderm — r.d., Resin -duct
— ry., (Fig. 14), Three of the four rays traversing the ground-parenchj^ma
— .§., Sclerotic cells — sc, Sclerenchyma — 4.y.6. (Fig. 14), Secondary vascular
bundles — t.t., Transfusion-tissue — c, Vessels of wood — i:.h. (Fig. 14), Vas-
cular bundle with apparent bundle-sheath — w.. Wood — x (Fig. 14), Traces
of fungal hypha;.
BV JOHN SHIRLEY AND C. A. LAMBERT. 609
EXPLANATION OF PLATKS LX.-LXVI.
Plate Ix.
Fig. I . — T. S. C'l-lospenn iim pan/ritJafiim F. v. M . ; ( x 24). ( 1 . Normales].
Fio-.-2.— T.S. Mllletia au-^fra/!.^ F.v.M. ; ( x 17). [1 .Normales].
Plate Ixi.
Fig.o. — T.S. Solaitiim Weiidlandii Hook. f. ; ( x 90). [1. Normales].
Fig. 4. — T.S. B/';/iioiiia Tireediana Lindl. ; ( x 17). [2 Chiastoxylon].
Plate Ixii.
Fig.o. — T.S. /J i(/iio)i id picf a h\nd\.; (x23). [2. Chiastoxylon].
Fig.O. — T.S. Carronia mnlti.'<e.palea F.v.M.; ( x lo). [H. Astroxylon].
Plate Ixiii.
Fig. 7. — r.S. Jir//oiu'a /acinio>!a hinn.; c/j. , cork-la^eis on side facing sup-
port: ( x 23). [4. EndophloiaJ.
Fig.S. — T.S. Wixtaria rhinensis DC; ( x 40). [.l.Exocycla].
Plate Ixiv.
Fig.U.— T.S. 17//^' stirred [folia F.v.M. ; ( x 40). [6. Phloiocycla].
Fig. 10. — T.S. l)On<ia'nirillea Kpertahilis .Tuss. ; ck., pad of cork opposite
supporting branch; ( x 28). [7. Polycycla].
Plate Ixv.
Fig. 1 1. — T.S. JJou;/ainril/ea laterkia Hort. ; ( x 90). [7. Polycycla].
Fig. 12. — T.S. /j. Sanderianax. ran'ef/ata Hort.; (x 90). [7. Polycycla].
Plate Ixvi.
Fig. 18. -T.S. Piper ui(/rum Linn. ; ( x 40). [7. Polycycla].
Fig. 14. — T-S. UaJeola cassythoide-^ Reichb. ; (xl9). [Monocotylei)0>' :
2. Abnormales].
610
OEDINARY MONTHLY MEETING.
October SOth, 1918.
Professor H. G. Chapman, M.D., B.S., President, in the Chair.
The President announced that the Council was prepared to
receive appHcations for four Linnean Macleay Fellowships,
tenable for one year from April 1st, 1819, from qualified Candi-
dates. Applications should be lodged with the Secretary, who
would afford all necessary information to intending Candidates,
not later tlian 30th November, 1918.
The Donations and Exchanges received since the previous
Monthly Meeting (25th September, 1918), amounting to 3 Yols.,
50 Parts or Nos., 11 Bulletins, 1 Report, 5 Pamphlets, and 1 Map,
received. from 43 Societies, etc., were laid upon the table
NOTES AND EXHIBITS.
Mr. Fred Turner exliibited a specimen of a very rare plant,
IWichi Ilium corymhosum Gaud., var. parvijiora Benth., from near
Barringun, the only specimen he had seen in New South AA^ales.
Mr. Froggatt showed specimens of an Indian wood-borer
[BostrychuH cequalis Jud.) introduced in the wood of boxes con-
taining helmets; and likely to cause trouble unless looked after.
Mr. E. Cheel exhibited specimens of two distinct forms of
plants commonly known as "Cobblers' Pegs'' {Frig e7'on linifolius
Willd.) showing the following characters :— (a) Plants with
lacinated, basal leaves, and narrow, linear, upper leaves; and with
comparatively small Howers; the commonest form, (h) Plants
with very large, basal leaves, more or less serrated or toothed;
and with much larger fiower-heads than in (a). The two forms
had been noted for the past five years in gardens, and on vacant
land at Wynyard Square and other places in the city and sub-
NOTES AND EXHIBITS. 611
urban districts. He suggested tliat they were two distinct
species, or it may be that one or othei- of tiie forms is heterozy-
gous, Botli are easily distinguished from the "Canadian Flea-
bane" {E. canadensis Linn.) found at Cronulla, but comparatively
rare in this State; as well as from E. honariensis, recently
brought under notice by Mr. A. A. Hamilton, which is fairly
common in New South Wales. — Also a series of seeds of "Soy
Beans" {Glycine hispida), which originated from a form known
in the trade as " Ebony Soy." The mother-plant " Ebony Soy,"
is a black-seeded form, which had been cultivated for four years,
and had bred true. In the tifth year, one of the plants gave
seeds of a greenish-yellow colour and distinctly green seeds in
the same pods. This was distinguished under the name "Aus-
tralia." When seeds of "Australia" were sown, the resultant
Fj plants gave seeds of the following colours : — (1 ) Plants with
light brown pods, and seeds similar to those of the parent. (2)
Plants with dark brown pods, and seeds similar to those of the
parent. (3) Plants with dark pods, and black seeds. (4) Plants
with ginger-coloured pods, and brown seeds, (a) Plants with
brown pods, and dark brown seeds. When samples of the above
were sown, the resultant plants (F.^) gave the following results.
No.l gave plants yielding greenish-yellow, black, brown, and
green seeds similar to those of "Australia." The seeds of No. 2
were similar to those of No.l. No. 3 produced plants yielding
some brown seeds, and others with black seeds, the pods and
seeds in both instances varying in colour and in the shape of the
seeds of different plants. No. 4 gave plants yielding all brown
seeds. No. 5 produced results similar to No. 4. Some distinctly
green seeds were separated from No. 1 , and sown separately, and
these yielded distinctly green-coloured seeds. The behaviour of
these plants is very similar to that noted in certain " French
Beans " (PAft.s'eo^ws vidgaris), recorded in these Proceedings 1914,
xxxix., p.l60; and 1916, xli., p.l92.
Dr. A. J. Turner exhibited a collection of J^epidoptera made
on the journey across Canada, on his way back to Australia
from Enijfland.
612 NOTES AND EXHIBITS.
Dr. Tillyard showed a collection of Canadian Lacewings and
Caddisfiies, presented to him by Dr. A. J. Turner. Also a large
Dragonfly {Petahira gigantea Leach), picked up on the pavement
outside the Society's gate on the 25th inst., in an exhausted, but
otherwise uninjured condition — a stray from the Blue Mountains
under the influence of westerly winds.
Professor H. G. Chapman, on behalf of Miss M. Lilley and
himself, exhibited some samples of yeast, on which observations
had been made on the rate of multiplication of the yeast-cells,
on the amount of sugar present, and on the production of alcohol
and carbon dioxide. The figures obtained showed that most of
the growth of the yeast occurred before the decomposition of
crlucose into alcohol and carbonic acid fjas.
613
THE RELATION BETWEEN THE FAT-CONTENT AND
THE ELECTRICAL CONDUCTIVITY OF MILK.
By H. S. Halcro Wardlaw, D.Sc, Linxean Macleay Fellow
OF THE Society in Physiology.
(From the Physioloyical Laboratory of tJie University of Sydney.)
Milk consists of water containing substances in solution, in
the colloidal state, and in suspension. The electrical conduc-
tivity of milk is due to the presence of ionised salts among the
substances in solution. The colloidal and the suspended matter,
the protein and the fat, do not contribute appreciably to the
transport of electricity. These substances, in fact, depress the
conductivity, for they displace a certain amount of conducting
material.
The eftect of protein on the conductivity of milk has been
studied bv Jackson and Rothera (1913). They removed the
proteins from separated milk by dialysis, and found that each
gram of protein in 100 cc. of the separated milk depressed the
electrical conductivity by 2 •76%.
The effect of fat on the conductivity is shown by the figures
given by Taylor (1913). He found, as an average of five ex-
periments, that the removal of 5% of fat from milk caused an
increase of electrical conductivity of 11 4%. This increase of
conductivity is more than twice as great as that which would be
accounted for by the increased concentration of electrolyte due
to the removal of the fat. The excess is ascribed by Taylor to
the removal of the mechanical obstruction which the fat-globules
may offer to the movement of the ions.
The object of the present work was to investigate more closely
the relation between the fat-content of cows' milk and its elec-
trical conductivity.
48
614 FAT-CONTENT AND ELECTRICAL CONDUCTIVITY OF MILK,
Tt has been found that, in a given sample of milk, the depres-
sion of conductivity is directly proportional to the fat-content;
but tliat, in different samples of milk, the removal of equal
amounts of fat does not lead to equal increments of conductivity.
Various other properties of the milk have been examined in the
endeavour to account for this difference between samples, but no
definite correlation has been found.
1. Itelati07i between the fat-content of a sample of milk and its
electrical cond/activity.— Mixtures of the same sample of milk
with different percentages of fat were prepared by spinning the
.samples in a centrifuge to remove as much of the fat as possible,
and mixing various proportions of the spun and the whole milk.
The conductivities of these mixtures of known fat-content were
determined.
The percentages of fat in the spun and whole milk were de-
termined by the Rose-Gottlieb method; the percentages of fat
in the various mixtures were obtained by calculation. The
results have errors of not more than 2%.
The electrical conductivities were determined at 25''C., by the
Kohlrausch method; a Wolff's Wheatstone-bridge was used, with
a telephone as a null instrument The determinations were
made in a conductivity- vessel provided with vertical electrodes.
The measurements obtained were concordant to 0*5%. The
details of these methods are given in a previous paper (Wardlaw,
1917).
As the effect of the fat on the conductivity is probably related
more closely to the volume occupied than to the weight in a
given quantity of milk, some preliminary determinations were
made of the density of the fat in the samples examined; from
these figures, the volumes can be calculated.
The density of the milk-fat was calculated from the densities
of the spun and the whole milk. The figures obtained are given
in the accompanying Table.
BY H. S. HALCRO WARDLAW.
615
Table i.
DtTmity of fat remoi'ed from cows' milk hy spinning.
Sample.
Density
of milk.
Percentage of fat.
Den.sity of
fat!
Spun.
Whole.
Spun ,
Whole.
5
10312
1 '0349
3-ol
0-19
0-933
7
10307
I '0347
8-98
O'lG
0-938
S
I '030S
r0352
3-78
017
0-923
9
ro30(;
1 '0849
4 03
0-13
0-933
10
1 '0812
1 '0854
3-80
0-18
0-9'29
11
1 -0310
J -085 1
3-78
0'17
0-927
12
1 -0808
1 -0847
4-0o
0-17
0'922
18
I -0308
I 0302
3-70
016
0-921
21
r0317
1 -0351
3ol
0-13
0 934
Mean
0-929
The determinations of density were made as described in a
previous paper (Wardlaw, loc. cit.), water at 25°C. being taken
as standard. The results have an error of al^out 1 in 10,000.
If the two errors are maximal and fall in opposite directions,
the calculated density of the fat will have a possible error of
about 2% The actual values obtained, as shown above, have
an extreme range of rather less than this, from 0-921 to 0-938,
and the variation from the mean is less than 1%.
The above results show that the densities of the fat removed
by spinning from the samples of milk examined varied between
0-921 and 0-938; 0-93 has been taken as the mean value in the
subsequent calculations. This value agrees well with that ob-
tained by previous investigators (Fleischmann, 1885).
The following Table shows the percentages of fat and electrical
conductivities (K) of various mixtures in whole and spun milk in
the case of different samples, and the ratios of the decrements of
conductivity to the corresponding increments of fat-content.
If the decrement of conductivity is directly proportional to
the increment of fat, these ratios will be constant for each
sample of milk.
ni6 FAT-CONTENT AND ELECTRICAL CONDUCTIVITY OF MILK,
Table ii.
Relation hefirefii fleet, -leal conductivity and fat-content of indiridual
.samples ofcoics' milk.
Sample.
Percent.
iii fat
Increment
K X 10"^
Decrement
D.KxlO-"
bj' vol.
of fat.
of K X 10--
D.Fat.
1
87.-)
2-60
4-72
2-9
11
8-42
2-27
47«
2-5
11
3 12
1-97
479
2 2
11
2-8-)
1-70
4-83
1-8
115
2-60
r4o
4-85
1-6
\'\
2-38
1-24
4-87
1-4
11
2-19
ro4
4-88
1-3
1 -25
ri.")
0
5 01
^»
2
0-29
()
5-43
0
1-57
1-28
5 -30
1-3
ro
205
1-76
5-25
1-8
ro
2-45
2-16
5-23
20
0-95
2 -76
2-47
5-19
2-4
roo
2-98
2-69
5-18
2-5
10
3-59
3-30
515
2-8
0-85
4-01
3-72
5 06
37
ro5
6
0-23
0
5-38
0
1-78
1-55
5-26
1-2
0-8
2-42
2-19
5 20
1-8
0-85
2-80
2-57
5-18
2 0
0-8
3-38
3 15
5-15
2-3
075
7
017
0
5-40
0
1-65
1-48
5-28
1-2
0-8
2-18
2-01
5-26
1-4
07
3 02
2-85
5-20
2 0
07
3 -53
3-35
5-18
2 '2
0-65
4*28
4-11
5-10
3-0
07
The values of K in this Table have a possible error 20 to 50
times that of the corresponding measurements of conductivity.
The above figures shoM^ that the ratio of decrement of conduc-
tivity to increment of fat is constant to within the limit of error
of the determinations for the samples of milk examined.
From these results, it will be seen that, for a given sample of
milk, the depression of conductivity is direcitly proportional to
the percentage of fat.
BY H. S. HALCRO WARDLaW.
617
2. Relation betiueen fat-cotUent and conductivity of different
samples. — The above figures also indicate, however, that the
depression of conductivity due to a given increase of fat-content
is not the same for different specimens of milk. To confirm this
observation, a number of determinations have been made of the
percentage-decrease of conductivity of spun milk due to the
addition of 1% of fat. These figures are given in the last column
of the following Table.
Table iii.
PerceiUaye-depi'essioii of etecfricaf conduct ir if i/ due to f/ie addihon of 1 c.c.
of fat to different samples of cowk'' milk.
Percent.
1
Sample
of fat
by vol.
D.Fat.
1
D.KxlO-'.
DKxlO-^
1)K% for
1 c.c. fat.
• 1
3- 75
4-72
1-15
2 -(50
5-01
2-9
2-25
2
4-01
5-06
0-29
3-72
5-43
37
1-8
8
3-87
4-84
0-19
3-68
5-10
2-0
1-3
4
4-15
5-07
0-18
3-97
5-37
3 0
1-4
5
3-78
5 10
0-23
3-55
5-41
3-1
1-65
6
3-38
5-15
0-23
3-15
5-38
2-3
1-35
7
4-28
5-10
0-17
4-11
5-40
3-0
1-35
S
4 -OK
5-11
0-18
3 '88
5-42
31
1-5
9
4-33
5-07
0-16
4-17
5-37
3-0
1-35
10
4-09
5-08
♦
0 19
3-89
544
3 •<)
17
11
4-01
5-09
0-18
3-83
5-37
2-8
1 -35
12
4-36
5-05
0-18
4-17
5-3(5
3-1
1-4
18
3-98
5-07
0-17
3-81
5-39
3-2
1'55
14
3-92
5 -OS
0-20
372
5-41
3-3
1-65
15
3-84
4-99
0-16
3-68
5-35
3 (J
1-8
618 FATL-CONTENT AND ELKCTKICAL CONDUCTIVITY OF MILK,
Table iii. — (conllnmd).
Sample.
Percent,
(jf fat.
by vol.
D.Fal.
U.K X 10 ".
D.Kx 10 -.
D.Xr for
1 c.c. fat.
Ki
3*98
o'Oo
0-U
3-84
.V33
2-8
1 •3.-)
17
4 -39
O-02
0-17
4-11
o'32
3 0
1-4
I-"*
370
-vol
<ri7
3 03
.5-42
41
2- 1.3
19
3-90
5-21
0-1-")
3-81
4-93
2-8
1-4
2(J
3-97
5-81
0-12
3 •8.1
0-47
3-4
r.3
21
3H.i
-3-37
0'14
3ol
0.71
3-4
17
'I'l
3 -09
5-43
0-is
3-41
572
2*9
r.j
•23
3-8H
5-49
Olo
3-71
5-83
3-4
r(i
24
3-70
o.4(5
0-14
3 -0(3
5-83
37
1-8
'2o
3-80
0-48
o-ir»
3 •().')
.V85
37
1 7.3
2(3
4-10
5 '51
0-13
3-97
0-89
3-8
1 -(33
27
3-45
0-88
(rlO
3-29
(J-21
3-3
\ry
2S
3-9(5
5-34
O-lo
3-81
o-(37
3-3
1 -33
29
3-7r>
o'oO
017
3 -08
5-81
3-1
1-3
3(J
3-72
0-48
0-10
3-56
o78
30
1-45
31
3-o5
o*o.>
o-k;
3-37
5-89
3-4
17
32
3-67
4-47
O-K)
3-51
5-81
3-4
1 '(33
Mean
1 -38
The figures in the above Table show that 1 c.c, of fat in 100
c.c. of milk depresses the conductivity 2-3 to 1*3% of the value
for spun milk in the samples examined, the mean depression
being 1"58%. The eftectof fat on the conductivity in these cases
is thus 30 to 120% greater than can be accounted for by the
amount of conducting material displaced.
BY H. S. HALCRO WARDLAW. 619
It was thought that this variable efi'ect upon the conductivity
might be due in some way to the method of removal of the fat
from the milk. The milk might be concentrated slightly by
evaporation during the process of spinning in the centrifuge.
The conductivity of the spun milk would then be greater than
could be accounted for by the removal of the fat. Or, substances
other than fat might be removed by spinning. In this way also
the electrolytes of the spun milk might become more concen-
trated, and the effect of the removal of the fat would be ex-
aggerated.
To test the first of the above suppositions, the conductivity
of samples of milk which had been spun covered to prevent
evaporation was compared with the conductivity of samples
spun uncovered in the ordinary way. The results obtained are
lifiven below.
Electrical conductivity of milk after spinning in covered and
uncovered tubes.
Conductivity.
Sample.
Covered.
Uncovered .
3
5-08
5-08
-t
5-37
5*35
o
5-39
5-40
These results show that there is no appreciable increase of
conductivity due to the concentration of the milk by evaporation
during spinning.
To test the second hypothesis, the amounts of total solid
matter as well as of fat in the milk were determined. The
following Table gives the weights of fat and of total solids in
100 c.c. of milk before spinning, and in the volume obtained
after spinning (100 c.c less volume of fat).
620 FAT-CONTENT AND ELECTRICAL CONDUCTIVITY OF MILK,
Table iv.
]Vci;/hfs of fat and of tolal solid matter remored from 100 c.c. of milk
hy spmnhuj in a cenlrifiifje.
Fat in
100 c.c.
Solids in 100 c.c.
Wt. from 100 c.c.
Solids.
Sainplo.
—
Whole.
Spun.
Whole.
Spun.
Fat.
Solids.
Fat.
8
3-78
017
!
1 12 70
8 91
3-61
3-79
1-05
9
4 03
015
1 12 76
8-54
3-88
4-22
109
10
3-80
018
12-72
9-07
3-62
3-65
1-01
11
3-73
017
12-66
8-94
3 56
3-72
1-05
12
4 05
017
12-95
8-82
3-88
413
106
13
3-70
0-16
12-60
8-95
3-54
3-65
103
14
3-65
019
12-58
8-92
3-46
366
106
15
3-57
015
12-88
8-95
3-42
3-83
1 12
16
3-70
013
12-72
8-92
3-57
3-80
I 06
17
4-08
016
13-00
8-96
3-82
4-04
1-06
18
3-44
016
12-39
8-99
3-28
3-40
104
20
3-80
0-11
, 12-85
9 00
3-69
3-85
1-04
21
3-39
013
1 12-40
9-01
3-26
3-39
1-04
22
3-44
017
12-45
9-09
3-27
3-36
1-03
23
3-70
014
12-72
8-97
3-56
3-75
1-05
24
3-54
0 13
12-56
9-00
3-41
3-56
1-04
25
3-64
014
12 57
9-05
3-50
3-52
101
26
3-92
0 12
12-99
9 00
3-80
3-99
105
27
3-31
015
12-01
8-85
3 16
316
TOO
28
3-79
014
12-80
8-86
3-65
3-94
1-08
29
3-59
016
12-59
9 08
3-43
351
1-02
30
3 55
015
12-51
9-00
3 40
3-51
103
.•^l
3-40
015
12 35
8-96
3-25
3-39
104
32
3 51
0 15
12-57
i
9 06
3-36
3-51
1 05
The above figures show that a certain amount of material
other than fat is removed from milk by spinning. Tliis is the
material adsorbed by, and adherent to, the fat-globules, and as
would be expected from the rather indefinite nature of its asso-
ciation witli the fat, its amount is rather variable, ranging from
12% to 1% of the weight of fat removed. If this material be
assumed to have the average density of the solids other than fat
of milk, 1-6, then the volume occupied by it will be only 0-6 to
7% of the total volume removed. This increase of volume is
quite insufficient to account for the excess of the efibct of the
fat on the conductivity over the volume-effect (30-120%).
3. Effect of degree of siihdivision of fat.— ^t was thought that
some light might be thrown on the variable effect of fat on the
BY ti. S. HALCRO WARDLAW. 62l
conductivity of milk by examining the relation l)et\veen this
effect and the number of fat-globules in a given volume of fat.
The effect of non-conducting suspended matter on the con-
ductivity of electrolytes has been studied by Oker-Blom (11)00).
He determined the conductivity of suspensions of sand in jellies
made up with salt solutions, and came to the conclusion that the
size of the particles (between 2 and under 05 mm.) made no
difference to their effect on the conductivity, but that the sand
depressed the conductivity less when uniformly distributed
through the jelly than when collected into one layer. These
conclusions are to a certain extent contradictory, however, as
the difference between the first and second cases is mainlv
one of aggregation. Further, the range of size of particles ex-
amined in the first case was not very great.
The sizes of the fat-particles even in one sample of milk vary
enormously. But each sample contains a certain average size and
number of particles in a given volume, and well-defined diff'er-
ences exist between the sizes and numbers for samples of different
origin, ^trippings, for instance, contain larger fat-globules than
first milk, and the milk of Jersey cows contains larger particles
than the poorer milk of Shorthorn cows.
The number of fat-globules in the milk was counted Vjy the
method of Babcock (1886) as modified by Shaw and Eckles
(1909). One volume of milk is diluted to 50 with water. The
mixture is drawn up into capillary tubes, the internal diameters
of the tubes are measured, and the numbers of globules in a
known length are counted. From the figures obtained, the
numbers of globules in a known volume of the undiluted milk
may be calculated. The measurements are made with an ocular
micrometer. The workers quoted made the optic measui-ements
with the capillaries immersed in glycerine. It was found in the
present work, however, that owing to the differences between
the refractive index of glycerine and that of glass, the values of
the diameter of the capillaries measured in this wa}- were too
high. The present measurements, therefore, were made with
the tubes immersed in a solution of chloral hydrate in glycerine
(7:1). This solution has a refractive index of 1'508, which is
622 FAT-CONTENT AND ELECTRICAL CONDUCTIVITY OF MILK,
very close to that of the glass used. The measurements were
checked by weighing the amount of mercury contained in a
known length of the tube. The accompanying figures show that
the optical measurenients of diameter agree well with those
calculated froni the weij^hts.
Tube,
Wei-ht of Hg.
Length.
Diameter
from weight. optically.
1
2
4
r3o m-.
I 07.5 ,,
0'9.3.5 ,,
1-22 ,,
13 ■57 mm.
17-3.5 ,,
16-15 ,,
22-2 „
0-0U7 mTii.
0-074.5 ..
0-74 „
0-071 „
0-01M> mm,
0073.5 ..
0 072 ,,
0 072 „
These figures show that the results obtained by the two
methods are concordant to within their limits of error.
In the following Table are shown the ratios of percentage of
fat to the percentage-decrease of conductivity, and the numbers
of fat-globules in 1 cu.mm for different samples of cows' milk.
Table v.
De;/ree of subdivision of milk-fat and its effect on the electrical conductivity.
Sample.
Fat in 100 c.c.
D.K%for
1% fat.
Globules
cu.mm, milk
per
X 10-6
Globules per
cu.mm. fat x IQ-*,
19
1-36
1 92
4-85
20
1-36
194
4-86
21
1-71
1-96
5-40
22
1-44
1-72
4-65
23
l-o3
1-79
4-51
24
1-73
1-98
5-20
25
1-69
2 02
5-17
26
1-58
1-89
4 47
27
1-57
2-27
6-38
28
1 49
2 10
5 15
29
I 4.5
2 02
5-22
30
1-42
1-95
5 09
31
1-65
1-77
4-84
32
1-62
212
5-61
As will be seen from the above figures, no simple relation ap-
BY H. S. SALCKO WARDLAW. 623
pears to exist between the (lepressioii of conductivity due to 1 c.c.
of fat in 100 c c. of inilk and the average number of globules
into which this amount of fat is divided, or the average size of
the globules. The degree of subdivision of the fat thus does not
seem to influence its effect on the conductivity in excess of the
effect due to the volume displaced.
4. The relation between the increase of inscosUij due to the fat
of milk and its effect on the electrical conductivity. —The electrical
conductivity of a solution of electrolytes depends on the velocity
and concentration of its ions. One factor which determines the
velocity of the ions is the resistance which the liquid of the
solution opposes to their movement. In the case of aqueous
solutions of salts only, this resistance is measured by the vis-
cosity. In the case of systems which are not homogeneous, how-
ever, the viscosity of the system as a whole is not a measure of
the resistance offered to the passage of ions. The addition of
gelatine to a salt solution, for example, may increase the vis-
cosity enormously, while the resistance to the movement of ions,
as measured by the conductivity, is hardly afiected. The system
is no longer homogeneous, but consists of two phases.
Milk is such a heterogeneous system, and contains at least
three phases. No simple relation is to be expected, therefore,
between the viscosity of milk as a whole and its electrical con-
ductivity. It was thought, however, that these two properties
might be connected in some regular manner which would explain
the disparity between the volume occupied by the fat of milk
and its efiect on the conductivity.
In the following Table are shown the effect of the removal of
measured amounts of fat on the conductivity and viscosity of
milk. The viscosities were measured in an Ostwald viscosimeter
at 25''C. The values given are those compared with water as
unity. The results have a maximum relative error of 1%.
624 FAT-CONTENT AND ELECTRICAL CONDUCTIVITY OF MILK,
Table vi.
Effect offal on condnrjlrihj a\id rhros^lly of milk.
.Sample.
Decrement
Increment
Viscosity.
DK%
DV%
of tat.
of K.
Whole.
Spun.
DFat%
DFat%
28
3 -92
3-3
\m
r.V)
1-38
ro7
29
3-«9
3'1
174
1 TV.)
1 •3r>
ro3
30
3-83
3 0
1-84
1 MU
1 -32
ro3
31
3-49
3-4
1-86
1 -59
ro3
105.
32
3-61
3-4
1-81
r(3i
I'ol
103
The above figures show that in these samples of inilk a de-
crease of 1 gm. of fat ill 100 c.c. diminished tlie viscosity from
3 to 7%. The corresponding increases of conductivity, however,
are from 30 to 50% above the values due to the volume occupied
by the fat. The diminution of viscosity is thus quite insufficient
to account for the discrepancy between the volume occupied by
the fat and its effect on the conductivity. It will be seen, too,
that the increase of conductivity is not proportional to the de-
crease of viscosity.
These results also indicate that, in the samples of milk ex-
amined, the effect exerted on the conductivity by 1% of fat by
volume is as variable as the effect exerted on the conductivity,
and does not show that constancy which Taylor {loc. cit.) found
in the samples examined by him.
5. The behaviour of fat-globules in an electric field. — If the fat-
globules of milk carry an electric charge, they may retard the
movement of ions in their vicinity, and in this way be responsi-
ble for the variable effect that fat exerts on the conductivity in
addition to the effect due to the volume displaced. The attempt
was made to observe whether the fat-globules were charged by
placing milk in a shallow cell less than 1 mm. deep, provided
with platinum electrodes (("hick and Martin, 1912). A current
was passed l)etween the electrodes, and the fat-globules were
watched under the microscope. No movement was observed
with a fall of potential below about 3 volts per cm. With higher
voltages, bubbles oi gas were formed at the electrodes, and
caused movement of the globules mechanically. These experi-
ments do not show conclusively that the fat-globules of milk
BY H. S. FIALCRO AVARDLAW. 625
are not charged electrically. They indicate, however, that any
charge carried must be small; much smaller, for example, than
that carried by the particles of a suspension of coagulated pro-
tein, which show verv evident movement in an electric field
under conditions similar to tliose described.
Summary.
(1). Removal of fat from milk increases the electrical con-
ductivity.
('2). In a giv^en sample of milk, the increase of conductivity is
directly proportional to the volume of fat removed.
(3). The increase of conductivity due to the removal of a given
amount of fat is not the same, however, in different samples of
milk. The average increase of conductivity due to the removal
of 1% by volume of fat is 1*5%.
In conclusion, I wish to express my indebtedness to Sir
Thomas Anderson Stuart, in whose laboratory this work was
done.
REFERENCES.
Babcock. — Ann, Rep. New York Agrie. Expt. Station, 1885, 298, 1886.
Chick and Martin. — Journ. of Physiol., 45, 261, 1912.
Fleischmaxn. — Journ. f. Landwirts., 33, 251, 1885. Quoted in Somnier-
feld's Handbuch der Milohkunde, 269, 1909. Wiesbaden : Berg-
mann.
Jackson and Rothera. — Biocheni. Journ., 8, 1, 1914,
Oker-Blom.— Arch. f. d. ges. Physiol., 79, 510, 1900.
Shaw and Eckels. — U.S. Uept. Agric, Bur. Anim, Industrj', Bui. Ill,
1909.
Taylor.— Journ. Proc. Roy. Soc. N. S. Wales, 47, 174, 1913.
Wardlaw. — These Proceedings, 1917, xlii., 815.
626
THE PANORPOID COMPLEX.
Part 2. The AViNn Trichiation and its Relationship to the
General Scheme of Venation.
By R. J. TiLLYAHD. M.A.. D 8c., F.L.S., F.E.8., Linnean
Macleay Fellow of the Society in Zoology.
(Plates Ixvii.-lxix., and Text-figures 17-^14).
If we cut off a small portion of the wing of any archaic Holo-
metabolous Insect, such as, for instance, Archichaiiliodes guttiferufi
Walk., (Text-fig. 22) of the Order Megaloptera, and make a
cleared mount of it, ^ve shall find that, when it is examined
under a high power, both veins and membrane of the wing are
covered with hairs. These hairs will be seen to be developed
equally abundantly upon the upper and under sides of the wing.
This covering of hairs may be spoken of collectively as the
Winy-trichiatimi. Though it is to be found in all Orders of the
Holometabola, it does not occur in all Orders of Insects. It isj
for instance, entirely absent from the wings of the Odonata.
Generally speaking, there may be found upon the wing of any
Holometabolous Insect two types of hairs, which I have already
named microtrichia and macrotrichia respectively (5). These may
be defined as follows: —
Microtrichia (Gr. [xiKpo's, small; and Opi^, r^txos, a hair) are
minute hairs, generally much curved or hooked, which are de-
veloped in connection with every unspecialised hypoderm-cell of
both upper and under surface of the wing. Hence the}^ occur
indiscriminately and exceedingly abundantly upon veins and
membrane alike. Theii- bases of insertion appear as simple
circular pits, in which, under a very high power, a central per-
foration leading into a very small lumen in the interior of the
hair can be made out. They vary in length from under l/u to
r»v II.
TILLYARD.
0-27
a.
d.
about 30/x, lint in most cases lio between 10/x and 20/x. Similar
hairs are frequently present upon the softer parts of the bod}-,
as, for instance, on the abdominal sutures and pleurto. (Text-
fig. 17, a, cZ).
Macrofrichia (Gr. fiaKpo^, large; and Opi^) are much larger
and stronger hairs, much less abundant, and oidy developed from
large specialised hypoderm-cells (trichogen cells) in connection
with special nerve-endings. They are thus of the nature of
i^ptisillfp, and homologous with similar hairs on othei* parts of the
body. Their bases of insertion are in the
form of raised circular rings, enclosing an
opening in the cuticle, which is continued as
a very definite lumen within the hair, almost
to its very tip. These hairs are almost uni-
versally present upon the main veins and
their branches, less frequently upon the
membrane of the wing, and only in very
special cases upon the true cross-veins. In
several Orders they become evolved, in cer-
tain cases, by a process of flattening, into
scales; but this development is only spasmodic
except in the Order I.epidoptera, where scales
are found universally. (Text-fig. 17,6, c, e,/).
Those macrotrichia found upon the veins
are usually somewhat larger and stronger than those found upon
the membrane, and both series may vary c(msiderably in size
upon the same wing. In different insects, the limits of variation
of these hairs may be placed at from 40/x to over 200/x. Text-
fig. 17 shows two sets of fairly normal hairs, one from Panorpa,
and one from Rhyphus.
It is the purpose of this Part to study the evolution of the
Wing-Trichiation in the various Orders of the Panorpoid Com-
plex. As this is partly dependent upon the state of evolution of
* Hairs from the wings of Panorpa confitsa VVestw., (a-c), and Rhyphus
brecis WaAk., {d-f); a, d, microtrichia; b, e, macrotrichia from wing-mem-
brane; f, /, ditto, from main veins; ( x 375).
a (U)
h ^ I
c. e.
Textfiir.l';
f.
628 THE PANORPOID COMPLEX, ii.,
the general plan of the wing-venation, it will be necessary, first
of all, to define the composition of this venation somewhat more
strictly than is usual.
According to the well known, and b}^ now generally accepted,
theory of Comstock and Needham (14), the venation of an insect's
wing has been original^ determined by the courses of the pre-
cedent tracheae, which supply the growing wing-rudiment of the
larva or nymph. From the six main trachete that enter the
growing wing, tliere are thus developed six maiv i^eiiis, known
as the costa, subcosta, radius, media, cubitus and analis, respect-
ively. Some of these main veins may be branched; such primary
brandies are termed sectors.
In the case of a main vein which branches and rebranches
several times, it is clear that the final branchlets may be botli
very small and very numerous. T propose to term such branch-
lets rciuhts. They are to be distinguished from cross-veins (see
below) by the fact that they are always preceded by definite
tracheae.
In contradistinction to a veinlet, I propose to restrict the term
cr^ss-veiit to a short connecting vein which does not form part of
the branching system of any main vein, and is developed inde-
pendently of the precedent tracheation.
It should be clearly understood that there is no reason why
such true cross-veins should not be developed upon any portion
of a wing, and at any thnc, provided the need for them arises.
There is thus, in reality, no difference between true cross-veins
and the so-called false cross-veins, which appear occasionally in
newly-expanded areas of the vyings of highly specialised genera;
as, for instance, in the enlarged humeral area of the hindwing of
Lasiocampa. The only difference between these newly formed
cross-veins and those found in older wings is one of time; where
the geological record is available to us, as in the case of the
Psychoj)sida', the rise of the true cross- veins from a similar origin
is clearly indicated.
In order to appreciate the difference between veinlets and
cross-veins more clearly, I have figured the tracheation and oor-
BY R. J. TILLYARD.
629
responding venation of a portion of the costal area of the fore-
wing of PsycJiopsis degans Guerin, (Text-fig. 18), in which this
difference is very clearly shown. It will be seen that the branches
passing into the costal area from the main subcostal vein (Sc)
are all of the nature of opinleU {ci'l), since they are all preceded
by trachefe. These veinlets are, however, supported by true
eross-vpins {x), which have arisen independently of the trachea-
tion. Also, the narrow areas between tSc, R, and Rs are
strengthened by true crogs-i'plns (a*).
In the known fossils of the Psychopsidcv and allies, we find the
costal veinlets present as in recent forms. I^ut there are no
connecting cross-veins. Hence it is evident that these latter are
a later development, called into existence by the need for
strengthening the enlarged costal area of the wing.
-^^<^^
Text-fig. 18.
Portion of enlarged costal area of fore wing of P.^ychopsis efer/an-'^ Guer.,
( x27), with the corresponding tracbeation (to the left) of the pupal
wing, ( X 60). C, costa; cf, trachet^ preceding the costal veinlets;
crl, costal veinlets; R, radius; Rs, radial sector; Sc, subcosta.
Referrins: aoain to Text fio. 18, we notice a further difference
between the veinlets and the cross-veins. The former, like the
main veins of which they are branchlets, carry a regulai*
series of macrotrichia, ^\'hereas the latter have no macrotrichia
at all. This distinction holds throughout the whole of the
Holometabola, with only two exceptions, viz., the Raphidioidea
49
630 THE PANORPOID COMPLEX, ii.,
and the higher faniihes of tlie Planipennia, in which the macro-
trichia appear upon the cross-veins as well as on the veinlets.
Having now clearly explained the difference between veinlets
and cross-veins, let us now go back to the most primitive fossil
insects, and study the venational scheme that they present to us.
In the oldest known Orders, the Palteodictyoptera and their
allies of the Upper Carboniferous, we find the same scheme of
main veins as in recent insects. The spaces between these main
veins and their branches are, however, filled up with an irregular
meshwork of cellules, quite unlike anything to be seen in the
Holometabola of to-day, though very probably homologous with
the still-existing dense meshwork of cellules to be found in certain
Orders of Hemimetabola, such as the Odonata. To this oriyinal
meshwork of cdhdes, as developed in the Palseodictyoptera and
their allies, I propose to give the name archedictyon. A portion
of a typical archedictyon is shown in Text-fig. 19.
Text-fig. 10.
Portion of wing of the fossil Hypermegethes shuckerti Handl., (Order Palaso-
dictyoptera), to show the archedictj^on; (nat. size).
It is here necessary to disabuse our minds at once of the idea,
if it exists, that the dense venation found in certain Planipennia
is an archedictyon. Practically all known fossils of this Order
(Triassic, Liassic, and Jurassic) have either no or few cross-
veins, and the spaces between their numerous main veins and
branches are devoid of any meshwork. Further, the oldest types
still extant, such as the Fst/chopsidte, If/io)ddce, etc., have no
meshwork present; and it is demonstrable that the apparent
BY R. J. TILL YARD. 63]
f'U'(ni-
meshwork present in the newer families, such as the Mijrmfh
iid(P. and Asmla^^hidce, is to be traced back to a simpler arrange-
ment of numerous parallel sectors supported b}' cross-veins at
intervals. It is by the zigzagging of these sectors, and the con-
sequent production of innnerous polygonal cellules, that a mesh-
work appearance has been produced.
The essential characters of the original archedictyon, as we
find it preserved in the Paljeodictyoptera, are its irregularity, and
the delicacy of the cemiles (I use this tei'm as distinct from vein-
lets or cross-veins) that form it. Whether these first arose in
connection with precedent fine trachete in the wing-rudiment, or
entirely independently of them, or perhaps partly in one way
and partly in the other, we have now no sure means of proving.
We can only say that the irregularity of their courses, and the
fact that, in one fossil at least, macrotrichia hav^e been found to
exist upon them, strongly suggest a tracheal basis.
It is now necessary to show the inter-relationship between the
archedictyon proper, the veinlets and cross-veins of the wing, and
the trichiation. Fortunately all these are preserved together in
the fossil Order Protomecoptera, from the Trias of Ipswich,
Queensland.
Plate Ixvii., figs. 9-10, show portions of the wing of Archipanoiya
magnijica Till., the only knosvn representative of this Order.
Here we may see the archedictyon still complete, but evidently
in a stage preparatory to becoming completely merged into the
wing-membrane. The venules of the archedictyon are not de-
finitely marked out, as in such fossils as HyjJennegethes (Text-fig.
19), but appear rather as simple ridges of the cuticle, not showing
any definite venular structure. In many places, but chiefiv close
to the main veins, and at the angles of the mesh work, there can
be seen rounded or slightly oval tubercles, of a diameter con-
siderably smaller than those seen upon the veins themselves, and,
with few exceptions, less clearly preserved. These are evidently
the bases of insertion of macrotrichia; but, partly owing to the
weak development of the mesh work that carries them, and parti)'
because they apparently lay more fiatly along the wing-surface,
632 THE PANORPOID COMPLEX, ii.,
their bases are not preserved as well as those on the veins, and
very rarely show the two concentric circles which characterise
the base of a stiff and more upright bristle. In ray original de-
scription (5. PI. viii., fig. 6), I gave a diagrammatic figure of a
small portion of the archedictyon of this fossil, much enlarged.
I now consider this figure to be somewhat misleading, in that it
made the macrotrichia of the mesh%vork appear as large and as
clearly marked as those of the veins, whereas they are always
smaller, and in most parts of the wing more difficult to make out.
It will be readily seen from the photomicrographs that it is not
an easy matter to give a correct drawing of this meshwork, and
I think it better simply to give the photographs in this paper.
So w^ell is this fossil preserved, that I have been able to find,
in several parts of the wing, by the use of careful lighting,
definite indications of the fine pitting due bo the presence of
microtrichia. Their bases of insertion are of about the same
diameter as that of the average particle in the grain of the rock,
viz., from 2 to 3/a. From a comparison with the microtrichia of
Panorpa, we may therefore conclude that the length of these
microtrichia was 40/x or more; i.e., considerably larger than an}^
to be seen in existing Holometabola.
Measurements of the beautifully preserved bases of insertion
of the macrotrichia upon the main veins of this fossil show that
the outer diameter of most of them lies between 30 and 40/x.
This is much in excess of the size to be found in the wings of
existing Orders, which seldom exceeds 10/x. Allowing some
expansion due to the pressure upon the wing during fossilisation,
it would still appear that this insect i30ssessed macrotrichia very
much stouter and longer than any now known to exist; their
probable length was between 300 and 400/^. The macrotrichia
upon the archedictyon were evidently smaller, and probably did
not exceed 200/x, their bases of insertion averaging only 20/v..
The cross-veins in the fossil are clearly seen to be developed as
strong struts between the main veins, and are quite independent
of the archedictyon. With the exception of <me or two macro-
trichia that appeal' to have strayed on to them from the latter,
BY R. J. TILLYARD. 633
they are found to be quite free from these hairs. Hence they
show no evidence whatever of liaving been developed ])y straight-
ening out of any portion of the archedictyon. If they had been
so developed, it is clear that they nuist have carried macrotrichia,
which they would receive from the archedictyon itself.
It will be seen, from the fossil Ao'chipanorpa, that the macro-
trichia occurred originally only upon the main veins and their
branches, including the true veinlets, and upon the archedictyon.
Cross-veins and membrane of the wing are alike free from them.
When, however, the archedictyon is abscjrbed into the membrane
(as it is on the point of being, in this fossil), it is evident that
the macrotrichia that originally occurred upon the archedictyon
nuist then become seated upon the membrane. This will explain
their appearance upon the membrane in many Orders of Holo-
metabola. But, if this explanation be correct, then the arrange-
ment of the macrotrichia might be expected to show some signs
of the original design of the archedictyon. If we draw the posi-
tions of a number of macrotrichia in any of the older Holometa-
bola, we shall see that this is the case. For this, it will only be
necessary to refer again to Text-fig. 2 2, in which it will be seen
that the macrotrichia below the main vein there figured tend to
lie along oblique lines similar to the courses of the venules running
out from the main veins in the fossil wing shown in Text-fig. 19;
and, like them, are connected by shorter cross-branches, indicated
by the presence of macrotrichia lying in intermediate positions.
(See also Text-figs. 21, 26, 30, 34).
Further evidence as to how the macrotrichia first appeared
upon the membrane of the wing may be obtained by a careful
study of the pterostigmatic region in the Planipennia. Text-fig.
20 shows this region in Micromm^ tasmanicn Walk. ' The series
of costal veinlets, that fills the costal space in all Planipennia,
originally extended into the pterostigmatic region. Here there
is a gradual thickening of the membrane taking place, with a
correspondingly gradual absorption of the veinlets; all stages of
this process can still be seen going on by studying various species
in the families Hemerohiidoi, Chrysojndce, and Manfispidce. As
634 THE PAKORPOID COMPLEX, ii,,
these veinlets cany loacrotricliia, it will be obvious that, as they
disappear, the iiiacrotrichia will become seated upon the mem-
brane.
Text-tig. 20.
Pteiostigmatic area of forewing of Micromua tasiiuini(x Walk,, showing
aphantoneuric condition of the veinlets; { x 15). C, costa; R, radius;
Sc, subcosta.
Thus we see that the evidence both of the fossil Order Proto-
mecoptera and of the recent Planipennia agrees in pointing to
the same conclusion: — Macrotrichia icere oriyinaUy carried only
upon the main veins and their branches^ and upon the archedictyon;
they ajypear upon the membrane by absorption of the veinlets or
venules^ carrying thein^ into the membrane of the iviny.
From this we come to the further conclusion that: — The
2)resence of macrotrichia upon the inembrane of ivings of oai open-
veined type (i.e., one ichich has no close meshivork of numerous
veins) is to be regarded as evidence of the descent of sack forms
from more deitsely veined forms.
The process by which the archedictyon, or any series of vein-
lets, becomes merged into the membrane, may be termed aphan-
toneurisnt, and veins which are in process of disappearing in
this manner may be called aphantoneuric. It is often possible
to reconstruct the aphantoneuric meshwork, merely by plotting
the positions of the macrotrichia upon the membrane, and join-
ing them up into a polygonal meshwork. Text-fig. 21 shows a
reconstruction of a small portion of the aphantoneuric meshwork
of the Hawk-moth Cceqnosa triangularis Don.
Having now explained how the Wing-trichiation is connected
with the venation in general, we may proceed to trace out the
various lines of evolution within the Orders of the Panorpoid
Complex.
BY R. J. TILLYARD. 635
Order PROTOMECOPTERA. (Plate Ixvii., fig,s.9-]0).
We have already described above the condition of the trichi-
atioii in the wing of the Triassic fossil ArrhipaH.orpa, the only
known representative of this Ordej".
It will be seen that
this wing differs from all .'.* -'•* ;•' ;-' *:•.''•'
known Holometabola at ''. . : - ; .':..;,.* .-*■.* .'•; ■'.
present existing in hav V'" •. v- ' l',;* •_';. •'■'
ing the archedictyon ; '...•' V. '.\- \' ■;'■':'. ■'/.
still present, upon all •,• •'. ;.;'•. •;.' ••.■••.."'.• '.'■!■
parts of the wing-mem- •/:. •'.■ .. ..-. .v.-'; "'' .•/•/.
l)rane, in an aphanto- .' v;. '.•:v.*';-. '-': ':'/'' \:
neuric condition, and .*.'. V'v '. • J; ;, ■•; •'•.,
carrying macrotrichia ' •',•; • '■'.' .'•:- ':;-.'r'^r-' -'/'.'
upon it, mostly at the *;; vv '•/.'' \:' •'.'■ •••*.
junctions of two or more ;•' V-. ■;*":'..■ .' .•'. ;; -^
venules. '":^ '.'''•'' V .." ••: ■:'.. '.V
Further, it should be '," '.• •.' •'}' /.' "'■','
noted that both macro- V! .'.•."'■- '*•"■.• ::•'.*' '•■■
trichia and microti'ichia
are considerably largei- Text-fig. 21.*
in this fossil than in other Holometabola. The tendency towards
reduction in size can be seen to be operating independently in all
Orders; it culminates, in the case of tlie microtrichia, in their
complete elimination from the wings of the higher families of the
Planipennia and Lepidoptera; and, in the case of the macrotri-
chia, in a great reduction in their number and shortening in tlieir
length, in the case of the Diptera and certain Mecoptera.
Order MEGALOPTERA. (Text-fig. 22),
The most archaic members of this family exhibit a near ap-
proach, in arrangement of the trichiation, to that seen in the
Protomecoptera, We have selected the genus Archichau/lodes as
* Arrangement of the bases of insertion of the scales on a small portion
of the wing of the Hawk- Moth C'crqaosa triangidaris Don., to show the
pattern of the lost archedictyon; { x 50).
636
THE PANORPOID COMPLEX, 11,
ail illustraticju (Text-fig. 22); it belongs to the most archaic family
Corijdali.d<t\ of the Suborder Sialoidea.
The arrangement of the trichiatioii differs from that of Archi-
panorpd only in the complete loss of the archedictyoii, and in
the smaller size of both macrotrichia and microtrichia. As has
been already pointed out, if the macrotrichia on any area of this
wing be plotted out, they will still show, by their positions on
the membi-ane, the pattern of the lost archedictyon. There is a
well dcA-eloped series of strong cross-veins supporting the main
A'eiris at wide intervals.
;V;V;>;\Y;^^ fW ^'v''*'V,;'V^\; of the wing all show macro
^y!M^f^!y^ ^(M 'i"}\'^^U\\ trichia well developed .
Wm^i^^^xi^^:^^^ When we turn to the
the
specialised genera, we
great advance over
condition seen in Archi-
hanliodes. Take, for in-
still present abundantly all
over the wing, yet the mac-
lext-tig.-i-J. rotrichia have quite dis-
appeared, except round the fringe of the wing, and on the ptero-
stigmatic area.
An advance in a different direction is to be seen in the Sub-
order Raphidioidea, of which the genus Raphidia will serve as
an example. Here the mici-otrichia have completely disappeared,
but the irregular thickening of the wing-membrane remains as
"" Portion of a main vein and surrounding membrane from the wing of
Archichaidiodes giittiferm Walk., to show the trichiation; ( x 100).
BY R. J. TILLYARD. 637
an indication of that disappearance. The macrotrichia have also
disappeared from the niejiibrane, but are still present on the
veins. As a high s})ecialisation, only to be })aralleled in the
highest families of the Planipennia, we note also thai the- macro-
trichia appear for the first time upon the true cross-veins, thus
rendering them indistinguishable from veinlets except by the fact
that they are not preceded by tracheae in the pupal wing.
We may sum up the tendency of evolution of the trichiation
in this Order by saying that both njacrotrichia and microtrichia
appear to have been, from the \ ery first, of small size; and that,
although the most archaic genera still show the original arrange-
ment of the trichiation, preserved almost in full, yet reduction
set in very early, leading to the loss of one or other series oi
hairs in all the higher types of the Order.
The wing-trichiation of the Archetype of this Order will be
taken to be similar to that of the archaic genus ArchLchaull(/dt'^,
but with hairs of somewhat larger size.
Order PLANTPENNTA. (Text-figs. I'O, 2:), 24).
The oldest known types of this Order show a wing-trichiation
already specialised in certain directions. Ko traces of the arche-
dictyon are to be found, and macrotrichia have, in every case,
been eliminated from the membrane of the wing (Text-fig. 2 3).
This is easily understood, when we remember that the wing-
venation of the Order early underwent a unique specialisation,
in the form of a ra})id proliferation of sectors of the main veins,
and especially of the branches of the radial sector. This pro-
duced the "Prohemerobiid" type (still to be seen in recent Psi/-
chupsid(r) in which the whole of the wing is covered with numer-
ous parallel longitudinal veins, with few or no cross-veins sup-
porting them. In the very narrow spaces between these veins,
it was clearly impossible for the archedictyon to exist, and the
macrotrichia upon it probably disappeared with it.
►Starting, then, from a type having no macrotrichia upon the
membrane of the wing, we find a furtlier specialisation, in the
higher families of this Order, in the complete elimination of the
638 THE PAXOEPOID COMPLEX, ii.,
micrutrichia. Thus we may (H^•ide the Order into two groups,
as follows: —
(1 ). 'J'he more archaic families, with microtrichia still present.
These are the Ithoiildw, IIe7)u>rohiidie, Dilaridct, Sisyridcf, Conio-
pteiyyidfc, Fsychojjsidct!, Folystcechetidce^ and Osmylidce.
(2). The more specialised families, witli microtrichia absent.
These are the TrichomatklfP,, Bei'othkln', Chrysopidce, Apochrysidf^',
Mantisjjidff, Nynqjhidce^ Xemopieridie^ Mynnelpontidce, and Asca-
laphidw.
r-^ -_» '^
'rL-xi-ii-.-i;:;.
Portion of a main vein and sunoundint; membrane from the win*; of Mirro-
nms tasinrunrr Walk., to .show the triehiation: ( x 200).
Ill the first group, macrotrichia are never found upon the true
cioss \eins. In the second, we find in the Trirhvmatidw and
Bci-dfli'uhv forms showing the passage of the macrutrichia from
the main veins on to the cross-veins. For instance, in the geiuis
Stt'iiohlplla, the cross veins remain without macrotrichia, as in the
first group. In SpermophorHla, (jne or two isolated macrotrichia
may be found upon most of the cross-veins. In Trichoma, there
is an intense proliferation of mao'otrichia upon the main veins;
and they overflow, not only on to the cross-veins, but also, in
some places, even on to the membrane itself. .Vs this genus is
evidently specialised in its excessive hairiness, there can be little
doubt that this latter occurrence is in the nature of a ca:'nogenetic
development, and is not to be cjnsidered as an archaic survival
of macrotrichia from an original archedictyon.
BY n. .1. TILLYAKD.
639
A[)art from the two t'anulies dealt with in tlie i)recec]iijti- }»ara-
giaph, all those listed in the second group have the inacrotiiehia
well developed upon the cross-veins.
Another interesting specialisation in this Order is iJic dt-Nel-
opment of .scales, which occur in the family Berothido' Thev
occur upon a more or less restricted area of the hind wing, in the
females only, of the genus Spermoj^horella (Text-fig. 24), and upon
the fringe of the wing in the genus
Isoscelipteroit. They are clearly only
modified macrotrichia.
The Archetype of this Order, then,
may be defined as having had the
archedictyon and its macrotrichia com-
pletely eliminated, owing to the unique
specialisation of the venation : the
macrotrichia, consequently, only oc-
curring on the main veins and their
branches, but well developed : and the
microtrichia also present and of normal
size. From this original type, the line
of evolution runs forward in the direc-
tion of complete elimination of the
mici'otrichia, with subsequent invasion
of the cross-veins by the macrotrichia,
and with the occasional formation of scales (in two cases only).
Text-tig. 24.
Order MECOPTEKA. (Text-fig. 25).
In this (Jrder, microtrichia are always exceedingly well de-
veloped all over the wing. Macrotrichia are found upon the
main veins and their Ijranches, but never upon the true cross-
veins. The archedictyon has been completely eliminated, but its
macrotrichia still persist upon the wing-membrane. (Text-fig. 25).
OwiniT, however, to the small number of macrotrichia that can
* Portion of a main vein fioni middle of hindwing of Sptrmojihorefia
dUiitminata Till., ? , showing normal macrotrichia interspersed with seed-
Hke scales; ( X 90).
640
THK PANORPOID COMPLEX, 11.
l)e accoiiuiiodated in the somewhat narrow spaces between the
main veins, the original pattern of the archedictyon tends to
bc'cumc lost, and the maci'otriehia of the membrane tend to
become arranged in regular sets occupying the central part of
any closed area of the wing.
The line of evolution in this Order is towards a gradual
elimination of the macrotrichia from the membrane of the wing,
and a reduction in their size elsewhere. In the most archaic
families, such as .Ueropidce (Text-fig. 25) and ChoristidfP. (Plate
Ixix., fig. 13), these hairs are still to be found upon the membrane,
in almost every part of the wing. In the Panorjndce, we may
see the stages of gradual elimination from the basal areas of the
Text-fig. 25.
A single areole from the wing of M<rope tiilx r Newni., to show the Irichi-
ation; ( x 100). Note the absence of luacrotrichia from the two
cross- veins bounding the aieole.
wiiK^ while those left in the more distal areas become, for the
most part, very regularly arranged in rows parallel to the main
veins. In the Naivnochoristida'^ we see a further stage of reduc-
tion reached, the macrotrichia being here entirely absent from
the membrane of the hindwing: while, in the forewing, they only
remain upon the anterior portion of the pterostigma. Finally,
in the highly specialised Bittacidce, all the macrotrichia have
BY R. J. TILLVARD. 641
disappeared from the membrane, while tliose left upon the main
veins have become short, stont spines, resembling those found in
many Diptera.
As the true cross-veins are always weakly chitinised, and
entirely free from macrotrichia in this Order, we have, in the
macrotrichia, a valuable aid in the tracing out of the courses of
the main veins. In many genera, the extreme base of Cuj is
sharply bent up to join M not far from its origin, and so takes
on the appearance of a cross-vein. But the true nature of this
vein is still proclaimed by the pi'esence of well-developed macro-
trichia upon it.
To sum up, then, we may characterise the Archetype of this
Order as differing from that of the Protomecoptera only in the
complete loss of the archedictyon. While the microtrichia remain
constant throughout the course of evolution, the macrotrichia
undergo a gradual reduction in size and number.
Order DIPTERA.
(Plate Ixviii., fig. 12; Plate Ixix., fig. 14, and Text-figs. 26-29).
We take this Order next, since it appears to be, on the evidence
of the wing-trichiation, a direct evolutionary derivative from the
base of the Order Mecoptera.
Throughout the Order, microtrichia are present upon the wing.
Macrotrichia are to be found upon the main veins and their
branches; they tend to become reduced both in size and number.
In only one family that I have examined are macrotrichia to be
found upon the membrane, viz., in the Rhyphidcp (Plate Ixviii.,
fig. 12, and Text-fig. 26). In this family, the structure and size
of the macrotrichia are closely similar to those of the older
Mecoptera. The arrangement of the macrotrichia upon the wing-
membrane, however, shows the pattern of the archedictyon much
more definitely than in any existing Mecopteron; so that, on this
character, the Rhyphidce can only be derived from a very early
form of Mecopteron, if they are to be derived from that Order
at all. In this connection, it is interesting to note that the
family RhypJiidct combines within itself characters of both the
Nemocera and Brachycera. This accords well with the condition
642 THE PANORPOID COMPLEX, ii.,
of tlie wiiig-trichiation; and the two characters taken together
suggest that this family represents one of the most archaic types
of Dipteron still existing.*
Text-fig. 26.
Portion of two main veins and the included membrane from the wing of
Rhyphus hreris Walk,, to show the trichiation; ( x l.SO). (See also
Plate Ixviii., fig. 12).
From the type of trichiation still existing in the Rhyphidcf, all
other types to be found within the Order are derivable, by
elimination of the macrotrichia from the wing-membrane. There
are, however, two distinct lines of evolution to be traced, in one
of which the macrotrichia tend to become slender and elongated,
and finally develop into scales; while, in the other, they tend to
become short and stout spines, and also become gradually
eliminated from all except the anterior veins and costal border
of the wing.
As conspicuous examples of the first line of evolution, we may
take the Moth Midges or Psychodkhf. (Text-fig. 27). Here we
see the macrotrichia as slender hairs abundantly present upon
" In this connection, it should be noted that the short vein usually con-
sidered as a cross-vein connecting Cuj with M. in Rhyphn-s is reallj- the
basal piece of M^, since it carries macrotrichia. On the other hand, the
vein below it, usually considered to be the basal portion of Cuj , is a true
cross- vein and carries no macrotrichia.
BY R. J. TILLYARD.
643
all the main veins and their branches. Here and there these
hairs are seen to be slightly flattened out, forming delicate and
narrowly lanceolate scales. The culmination of this same line of
Text-fig. 27.
Two portions of a single main vein from the wing of Psi/choda sp.,
(Hornsby, N.S.W.), to show the triehiation. On the left, normal
maei'otrichia, one or two slightly flattened: on the right (distal end
of vein), narrow lanceolate scales; { x 875),
evolution is to be found in the Ciilicidce (Text-fig. 28), in which
all the macrotrichia, even those along the wing-border, have
become scales; and these scales, in most cases, are of broad form
and specialised sculpture.
Along the second line of evolution, which has been followed
by many families of Nemocera, and also by the whole series of
the Brachycera and Cyclorrhapha, we can follow out the gradual
elimination of the macrotrichia, through such forms as the
Tah(iuid<p, where they are still present in large numbers on the
more anterior veins of the wing, to the higher Cyclori'hapha,
wliei-e they finally cease to exist anywhere except upon tlie costal
border of the wing. Text-fig. 29 shows the type of short, stifle*
seta into which the macrotrichia develop in this line of evolution.
It should be noticed, also, that, concurrent]}' with the broadening
and strengthening of certain veins, the macrotrichia upon them
644
THE PAVORPOID COMPLEX, 11.,
liecome arranged two, three, or even four rows deep, so that the
whole vein takes on a spiny appearance. Concurrently with this
chanue in the macrotrichia, we find two lines of evolution of the
inicrotrichia. They either undergo
gradual reduction also, until they
become partly or entirely eliminated,
as in the family Syrphichf: or they may
remain well developed, with a differ-
entiation setting in in the size of the
two series on the upper and lower sur-
faces of the wing. In this latter case,
the microtrichia on the upjjcr surface
increase considerably in size, while
those of the lower surface remain fairly
constant. This de\'elopment takes
place in the Myiodaria, and is especi-
alty Nvell shown in the Muscidce. in
the Blowfly, for instance, the micro-
trichia of the upper surface of the %ving
are quite 30/x in length, and are set in
large, swollen bases, while those of the lower surface are only
about half as long, and have the usual small bases of insertion.
(Plate Ixix., fig.14).
Text-fig. 28.
Text-fig. 29.
Small portion of a main vein from the win^i; of Tabanu.^ sp., (Hornsbj',
N.S.W.), to show the trichiation: ( x 200).
While, therefoi-e, the main mass of the Diptera shows consider-
able specialisation in its wing-trichiation, we have to postulate
* Small portion of a main vein from the wing of a Mosquito, Mncidni<
alternans Westw., 9 , to show scales; { x 200).
BY R. J. TILLVARD.
645
for tliis Ordei", on account of the existence of the arcliaic Rhi/-
phidit', an Archetype similar to tliat of the Mecoptera, but with
the microtrichia somewhat smaller than in that Order.
Older TRICHOPTEHA. (Text-fios.:^(), :',1).
In this Order, the most archaic tj^pes, such as tlie Rhyacophil-
id(f, alread}' show considerable specialisation in their wing--ti'i-
chiation. The archedictyon is always absent, but its macrotrichia
are present upon the wing-membrane, and remain there, on the
whole with ever increasing abunrlance, as we pass to the highest
types within tiie Order. Microtrichia are present throughout
the Order, but are always small, averaging about 5//. in length.
Macrotricliia are present upon the main veins and their branches,
but never upon the few and weakly developed cross-^'eins, except
in the cases where the proliferation of these hairs upon the mem-
brane leads to a secondary invasion of all parts of the wing.
Text-fig. 80.
Portion of two main veins and the included menibiane from the win^ of
Rhyacopltila f/o/-s«//.9 Curtis, to show the tHehiation;( x 10")). (Micro-
trichia are slio;litly exaggerated in this figure).
Tn describing the macrotrichia of the Protomecoptera, we have
already mentioned that those found upon the archedictyon were
50
G46 THE PANORPOID COMPLEX, ii.,
somewlmt smaller in size than those upon the veins. This differ-
ence is also to be seen, somewhat more intensified, in the oldei-
types of Trichoptera, as may be seen in' the figure taken from
lihyacophihi dor^<ilis (Text fig. 30). Not only are the macrotrichia
of the veins larger, but they remain for the most part straight
and stifi:', whereas those of the membrane (derived originally from
the arched ictyon), are slenderer, and inclined to be slighth^
curved.
A single line of evoluti(m maybe said to characterise the whole
Order. It consists in the gradual greater and greater prolifera-
tion of the macrotrichia of the membrane, togethei' with various
specialisations of the macrotrichia upon the veins. Thus there
is produced that excessively hairy type of wing that is especially
associated with this Order. In many genera, the stiff bristles of
the veins become reduced, either in part, or altogether, to the
same type of slender flexible hairs that we find upon the mem-
brane; in others, certain sets of hairs, especially at the base of
the cubitus, become erect and bristly. In the Hydroptilid(€, all
the macrotrichia of the forewings become erect and thickened,
while the hindwing develops an exceedingly long fringe.
h
Text-lig.81.
Forms of scales found in Trichoptera : n, a narrow, lanceolate scale, with
sini^le longitixdinal stria, from wing of an unnamed Leptocerid
(Broken Hill, X.S.W.); ( x350): h, a lanceolate scale with three
longitudinal striie from wing of Plectrofarsns (jravenhori^ti Kol. ;
( X 400).
The slender, flexible macrotrichia of the membrane found in
the wings of Trichoptera are exactly comparable with those of
the PsijckodidcH in the Order Diptera (Text-fig. 27). Like them,
thev are easily converted into elongated lanceolate scales. The
simplest type of such a scale is one in which the flattening pro-
duces only suflicicnt width to allow of the development of a single
fiy R. J. TILLYARb. <^47
longitudinal stria (Text-tig.31/c.). Such scales may be found in
([uitc a number of isolated genera within the Order. Further
broadening leads to the addition of a second, third, or even a
fourth parallel stria. The best developed scales known to me in
this Order are those of the pecuhar genus Plpcfrofrirf<ufi(Text-i\g.
'U,/)), which always show three or four stria\
Certain groups of genera show a tendency to the reduction of
the hairiness of the wings, which is evidently of a secondary
nature. In the subfamily Macro neii tat iiixf, this culminates in
the exolution of a numbei- of forms with the wings almost devoid
of macrotrichia.
In conclusion, the Archetype of this Order must evidently
have had a wing-trichiation closely resembling that of Ehya-
cojjhila, i.e., with the archedictyon absent, its macrotrichia pre-
sent uj^on the membrane, and already differing considerably, in
their smaller size, and slighter, more flexible build, from the
larger and more bristle-like macrotrichia of the veins, Macro-
trichia were absent from the cross-veins, and the microtrichia
were already reduced to a small size.
Order LEPIDOPTEllA.
(Plate Ixviii., fig.ll, and Text-figs. 21, 32-34).
In this large Order, it will only be necessary to study fully
the most archaic types; in the higher families, only the general
trend of the evolutionary effort need be considered.
We take, then, first of all, the three families of the Suborder
Homoneura, viz., the Micropterygidce, Prototheoridoi, and Hepi-
alidce. In all (jf these, microtrichia are present, but exceedingly
minute, never exceeding 2/a in length. They have been called
(i.ndexp, in this Order, and an attempt has been made to class
together all those families, in which they occur, as the "Aculeate"
Lepidoptera. They occur in a number of the older families of
the Heteroneura, as well as in the Homoneura. Hence it will
be seen that this method of classification is faulty, and should
not be persisted in. In all the higher families, microtrichia are
no longer present; but it is evident that their loss has been
effected along many lines of advance; and that an association of
(348
THE PANORPOID COMPLEX, 11,
all '•iKJii-aculeate' funiisinto one Suborder would be as unnatural
a grouping as it would also be in the Order Planipennia.
The most striking character of the Order Lepidoptera is the
specialisation of the niacrotrichia as scales, throughout the whole
Order, from the lowest to the highest forms. Although, as we
have already seen, scales are to be found on the wings of other
Orders of Holometabola (Planipennia, Diptera and Trichoptera),
vet in each case they only occur spasmodically, as cases of high
specialisation in one or more isolated groups. But, in the Lepi-
tloptera, the scale-bearing habit is an ordinal character, the only
exceptions to which can be traced definitely to retrogression
(e.^., in the Fsyrhid(t').
%.:.Q.:.
V .
Text-%.32.
.Sniall |)ortion of a main vein and adjacent menilnane from tlie forewing of
rrofof/i«:ora pefrosemfi Meyr., (8. Africa), to sliow the trichiation;
( X 200 1. (From a partiall}^ descaled specimen).
In order to establish the proof that these scales are modified
macrotrichia, it is only necessary for us to study such an archaic
type as FrofotJwora''' (Text-fig. 3 2), in which the macrotrichia
upon the veins remain in their original state of somewdiat flexible
hairs, while the arrangement of the bases of insertion of the
scales upon the membrane can be seen to correspond wdth the
pattern of an original archedictyon. It will also be noticed that
* For the supply of material of this rare genus, I have to thank Dr.
Peringuey, Director of the South African Museum, Capetown.
BY n. J. TILLYARD.
649
the scales that lie closest to the veins are elongate and narrow;
and tluit, tlie further we go away from a vein, the hroader and
shortei- the scales become. Thus we can find, in this one insect,
all stages in the evolution (^f a scale from a macrotrichion present
upon different parts of the wing.
Similar variations in the form of the scales, (X)nnected with
their positions in relation to the veins, are to he found in the
Micropfj'ruyidit' (Text-iig..i.S); Init, in this family, the broader
scales ai-e much more abundant, and may frequently be seen (juite
close up to the veins.
a
Text-tig. 33.
Three types of scales found in the family Micropfery<jid<P', ( x 400) : a, from
Eriocrania ■sttnipurpvref /a Hteph.; h, from Mnemonica Hnhpurpurella
Haw,; c, from Sahatliica inroii(/nfe//fi Walk. (All three types may
be found upon any one genus).
In the JIf'pialidcc, both narrow, lanceolate scales and moder-
ately V)road, oval ones are scattered all over both membrane and
veins, interspersed with fairly runnerous, slender, flexible hairs,
or macrotrichia which have remained unaltered in form.
It vvill not be necessary, foi- the purposes of this paper, to
follow out the many specialisations to be found in the scales of
650 THE PANORPOID COMPLEX, ii.,
tlie Hetereiieura. Tliese liave already been very fully dealt with
by many authors.
Returning* to the Froiofheoridit, it is of the very greatest
interest to note that traces of the original archedictyon are still
to be found in this archaic family. In Text-fig. 34, and Plate
Ixviii., Hg. 1 1, J show the very definite archedictyon that is to be
seen above the radius. Here, not only do the macrotrichia
(which, in this region of the wing, are all scales, not hairs) show
very clearly the original arrangement of the mesh work, but there
is actually a sliglit thickening and darkening of the membrane
forming the archedictyon itself. Allowing for the difference in
the sizes of the two wings, this condition in Prototheora is closely
similar to that preserved in ArchijKinorpa. Though I have not
. X>""Vv.)»^:. k ^' ■■:■■ .A .9
fl
Cl
A
A
">'"'
^''
*<■
% .^'
9.
D
9
4 ;
;/>,
\
'•■^:
Ji
>-^
-K /
^
t
Text-fig. 84.
Portion of the aphantoneuric archedictyon preserved above the radius of
the forewing in Profotheora petrosema Meyr. ; ( x 180). (>See also
Plate Ixviii., fig. 11).
been able to find so good an example of an aphantoneuric arche-
dictyon as this, anywhere else within the Order, yet there are
numerous cases in which what appears to be the last remnant of
the archedictyon can be seen in proximity to the veins, usually
in the form of a series of slightly darkened and thickened spurs
projecting almost at right angles to the main vein upon which
they abut.
Reviewing the above evidence, it would seem that, in the
Lepidoptera, the foi-mation of scales must have begun befoi-e the
complete disappearance of the archedictyon. This suggests that
the scales in this Order must be an exceedingly archaic character,
originating in a specialisation of some type belonging to an
ancestral Order in which the archedictyon still persisted.
BY R. J. TILLY ARD. 651
Hence we conclude tliat tlie Archetype of this interestinij
Order must have possessed at least some portions of the aphanto-
neuric arcliedictyon; that it also possessed inicrotrichia of small
size; and that the maci'otrichia, though remaining as slender,
flexible hairs upon the veins, were largely interspersed with true
scales upon the wing-membrane, such scales being of elongate,
lanceolate form, with few stride.
We have now to review the above evidence, in order to dis-
cover what light it throws upon the Ph^^logeny of the Orders
included in the Panorpoid Complex.
First of all, it must be evident that, as far as the Wing-tri-
chiation is concerned, all six Orders may well have been derived
from a single ancestral Order characterised by the following-
points: presence of an aphantoneuric archedictyon, presence of
microtrichia all over the wing, and pi-esence of well-developed
macrotrichia upon the main veins and their branches, and upon
the archedictyon, but not upon the true cross-veins.
Such a type is preserved in the fossil Archipanorpa from the
Trias of Ipswich, Queensland. This fossil itself, however, cannot
have been the ancestral form, since the Mecoptera, Trichoptera,
and Planipennia, at any rate, are contemporaneous with it, while
the first of these three Orders goes ])ack also into the Permian.
For the actual ancestor we must postulate some unknown type
of the Lower Permian, or possibly even of the Upper Carbon-
iferous; and whether this ancestor is to be considered as belong-
ing to the Protomecoptera or not, will have to depend upon
other characters not dealt with in this Part.
Let us now follow the evolutionary changes of the various
structures of the wing separately.
The Archedictyon : — This undergoes complete suppres-
sion in all recent Orders, except only in a few Lepidoptera, where
distinct traces of the mesh work in an aphantoneuric condition
(but not so well marked as in Archipanorpa) are still to be found.
This is well shown above the radius in the forewing of Proto-
tkeora pHrosema Meyr. (Text-fig.34, and Plate Ixviii., tig. 1 I).
652 THE PANORPOID COMPLEX, ii.,
After the elimination of tlie arcliedictyon, the hases of inser-
tion of the macrotrichia left upon the membrane of tlie wing still
show, fairly plainly, the original pattern of the lost meshwork.
A study of the meshwork in the figures of Prototheora ali-eady
referred to will soon convince us that it is possible to reconstruct
the course of the meshwork with fair accuracy, by joining up
the bases of the macrotrichia along the lines which they still
indicate. The resulting figure will not be correct in all parti-
culars, since some of the cross- venules of the mesh do not carry
any macrotrichia at all, and would therefore be onntted in the
reconstruction; while, in a few cases, the positions of the macro-
trichia are more or less misleading.
In specialised cases where the macrotrichia proliferate freely
on the veins and membrane, as in the case of the higher Hetero-
neurous Lepidoptera, there are always more than sufficient macro-
trichia to ensure an accurate tracing of the original pattern of
the archedictyon. This is well shown in the figures of the Hawk-
Moth Cmqiiosa triangularis Don., (Text- fig. 21). Here it will be
seen that the meshwork, probably while it was still aphantoneuric,
tended to become arranged into almost parallel lines connected
by irregular cross-pieces. It is this tendency, carried to its
highest development, that gives the very regular arrangement of
the scales seen in the Butterflies. But in all cases, by careful
study, it can be seen that the arrangement is a modification of
an original meshwork, and not a series of unconnected straight
lines.
The evidence afforded by the archaic Bhi/phas in the Diptera,
and by both the Homoneura and the older types of the Hetero-
neura in the Lepidoptera, points definitely to the conclusion that
these two Orders, like the less highly specialised Orders of the
Panorpoid Complex, had their macrotrichia arranged upon the
membrane in the original positions which they occupied upon
the aphantoneuric archedictyon.
The M i c r o t r i c h i a : — Though absent in the higher types
of some Orders, ^.y., Megaloptera, Planipennia, Diptera (a few),
and Lepidoptera (the great majority), yet they are found to be
BY R. J. TILLYARD. 653
present upon the wings of archaic types of all the Orders of the
Panorpoid Complex. The line of evolution is towards reduction
in size in all the Orders, with the single exception of tlie Myio-
daria in the Diptera, where there is an increase in size of the
microtrichia upon the ujyper surface of the wing only, resulting
in the interesting condition found in the Blowfly and allies (Plate
Ixix., fig. 14).
The Macrotrichia : — A study of archaic types, includ-
ing the fossil Archijyamnya, shows us that these were originally
present upon both archedictyon and main veins, but not upon
cross-veins. Moreover, probably because of the comparative
weakness of the venules of the archedictyon, there was, from the
very first, a considerable difference in size between the macro-
trichia of the veins and those of the mesh work. The macrotrichia
of the veins also tended to be more erect than those of the mem-
brane; so that, in the fossil ArchijKinorpa, their bases of in-
sertion show the two concentric circles of the raised disc very
clearly, while those of the meshwork rarely do so.
We are thus led to expect the following lines of evolution,
which do, in fact, actually take place in the various Orders of
the Complex : —
(1) If the direction of the evolutionary effort be towards re-
duction, then the macrotrichia on the meshwork should be
affected first, since they are smaller in size from tlie first, and
inserted in less strongly built bases.
As a matter of fact, it will be seen that the tendency towards
reduction in the smaller macrotrichia becomes accelerated in the
course of evolution; so that, in many cases, they become quite
eliminated, while the macrotrichia upon the veins may even pro-
liferate, though usually undergoing some reduction in size as
well, as in the higher Diptera.
(2) If the direction of the evolutionary effort be towards the
formation of scales, then the macrotrichia of the meshwork should
be affected first, since they are in a more favourable condition,
both as regards their delicacy of structure and the condition of
their bases of insertion, to undergo this change.
51
654 THE PANORPOID COMPLEX, ii.,
The formation of scales in the Lepidoptera and Trichoptera
agrees with this; since, in archaic famiHes of the former Order,
and in all scale-bearing genera of the second, we can still find
some at least of the macrotrichia on the main veins remaining
in the form of hairs. Judging also from the tendency, in the
Lepidoptera, for the broadest (most highly evolved) scales to
appear furthest from the veins, while narrow lanceolate scales
are found nearest to the veins, we are led to the same conclusion.
Actually, the covering of scales is not needed so much upon
the veins as upon the membrane, and the original condition of
the macrotrichia is such that the need of the insect is to be met
along the easiest line of specialisation; i.f\, those macrotrichia
whose alteration into scales would most benefit the insect also
happen to be those in the best condition to undergo the change.
In the other Orders in which scales appear (Diptera, Plaiii-
pennia) it should be noticed that they ai'e a comparatively late
effort, which takes place after the macrotrichia have been com-
pletely eliminated from the membrane of the wing. The scales,
in these cases, are formed from the macrotrichia left on the
veins. If these are still somewhat bristly, as in the Planipennia,
a hard, seed-like scale is produced (Text-fig. 24); if, however,
they have become slender and delicate, as in the Psychodidoi and
CnlicidcB, then the resulting scales will resemble very closely
those of the Lepidoptera and Trichoptera.
The changes in position of the maci'otrichia, in connection
with the disappearance of the archedictyon, have been already
dealt with above.
Taking all the above evidence into account, we may reason-
ably come to the following conclusions, as far as the limitations
of this Part of our study permit: —
(1) The fossil Order Protomecoptera is undoubtedly the most
archaic type known within the Panorpoid Complex, as regards
the characters of its Wing-trichiation.
(2) A single line of descent from the Protomecoptera to the
Mecoptera, and from the very base of this latter Order to the
Diptera, is strongly indicated by the uniform type of trichiation
BY R. J. TILLYARD. 655
found in these three Orders, togetlier with tlie very definite
evolutionary trend towards reduction.
(3) A second line of descent is less definitely indicated, com-
prising the Megaloptera and Planipennia, in which the evolu-
tionarv tendency runs towards early complete suppression of
the inacrotrichia upon the membrane of the wing, and a later
invasion of the macrotrichia from the main veins on to the cross-
veins, culminating in the Kaphidioidea on the one hand, and in
the higher Planipennia on the other. Both Orders may well
have been derived from an early type of Protomecopteron, but
neither of them can be derived from the other. For the oldest
Planipennia have the largest microtrichia, while the oldest
Megaloptera still retain the macrotrichia upon the membrane, a
€ondition which is not to be found within the Planipennia.
(4) As the Trichoptera show a more archaic condition of the
macrotrichia upon the membrane than do most of the Mecoptera,
they can only be derived either from the very base of this latter
Order, or from the preceding Protomecopterous type.
(5) In the Lepidoptera there exist certain types that show
portion of the archedictyon still present in an aphantoneuric
condition: while, in many others, the pattern of the archedictyon
is well preserved. Botli the oldest Trichoptera and the oldest
Mecoptera are in advance of these types. Hence we can only
trace back the Lepidoptera to tlie Protomecopterous or some
similar, extinct type, and may not derive them either from the
^lecoptera or from the Trichoptera.
Tliough the conclusions to be drawn from the structures
studied in this Part are somewhat indefinite in thejnselves, they
will be found to be of considerable value wlien taken in conjunc-
tion with the rest of the evidence. Also, in the study of the
Phylogenies of the families of each separate Order, (which lies
outside the actual scope of this paper), a great deal of vevy valu-
able evidence may be gathered from the Wing-trichiation.
We may conclude this Part by giving, in taljular form, the
principal characters of the Wing-trichiation for the different
Orders —
656
THE PANOKPOID COMPLEX, 11.,
Table of the Condition of the Wino-trichiation
Panorpoid Complex.
* indicates well-developed.
in the Orders of the
present in reduced form, - absent.
Macrotrichia.
Orders and (t roups.
Micro-
trichia.
On main
veins and
On cross -
On arche-
dictyon or
Arche-
dictyon.
branches. .
membrane.
Protomecoptera
i
i
*
present,
aphanto-
neuric.
Megaloptera : —
t
Archetj^pe
*■
* —
*
-
Corydalkhf
.=
*
-
Sialidd'
*
_ _
(A)
-
Bap/tldilda ...
-
*
*
-
-
Planipennia: —
Archetype
*
-
-
-
Archaic types
* or X
* -
-
—
Higher types
"
(B)
-
Mecoptera : —
Archetype
*
:*.: —
*
-
Archaic types
>;:
•
* or X
-
Nannodioristid(f' \
and Bittacid(f j
*
*
(C)
-
Diptera :—
Archetype
^
•
*
-
Ehyphkbe
*
*
*
-
Psychodidfe
X
some scales -
-
-
Cidicidce
X
all scales
-
-
Other types
*, X or - ,
(1))
* or X
"~
"~
Trichoptera :—
Archetype ...
X
1
* ! —
*
-
Recent families
X
I
very abun-
dant, some-
times
~
scales. (F)
Lepidoptera :—
Archetype
X
*
hairs and
scales
at least
partially
present,
aphanto-
neuric.
Profotheorid"
X
* (hairs
and scales)
scales
partially
present,
aphanto-
neuric.
Other Homoneura ...
X
-
-
hairs and
scales
-
Archaic Heteroneura
X
,,
-
J J
-
Higher Heteroneura
scales
scales
Special References:— (A) Only present on fringe and pterostigma— (B) Some
scales developed on hindwing of SpennophoreUa 9 ? and fringe of Isosce/ipferon
— (C) A few present on pterostigma of forewing in Xanyiochori.^tida' — (D) Absent
in some Si/rphldip; enlarged on upper surface of wing in Myiodaria — (E) Absent
except as a late invasion from the membrane— 'F) Spar.se or absent in Macro-
nemaihxv and some other forms.
BY R. J. TILLYARD. 657
BIBLIOGRAPHY.
(Xofe. — The refei'ence-numbers are made consecutive from Part to Part,
hut only those referred to in any ^iven Part are printed with that Part).
5. Ttllyard, R. J., 1917. — " Mesozoie Insects of Queensland. No. 1.
Planipeimia, Trichoptera, and the new Order Protomecoptera. '"
These Proceedings, IHIT, xlii.. Part 1, pp.l7o-20(), PI. vii.-ix.
14. CoMSTOf'K, J. H., and Needham, J. G., 1898. -"Tlie Win^s of
Insects." American Naturalist, 1898, xxxii.
EXPLANATION OF PfyATKS LVII.-LXIX.
Plate Ixvii.
Fi2;.9. — Portion of forewin^ of tlie fossil Arrliipanorpa /nat/niffra Till.,
showing the aphantoneurio arohedietyon witli tubercles representing
the bases of insertion of the macrotriehia; also showing portion of
the main veins R„ and R^, with clearly marked l)ases of insertion
of larger macrotriehia, and ci'oss-veins Mithout tlie same; ( x 27).
Fig. 10.— Part of rig.9. further enlarged; ( X 4.S).
Plate Ixviii.
Fig. 1 1 . — Portion of forewing of Protofhconi petro-sema "Sleyr., (Lepidoptera;
S. Africa), showing the radius, with the aphantoneurio arohedietyon
above it, cai-rying scales (macrotriehia); from a cleared mount of a
partiallj'^ de-scaled specimen; ( X 50).
Fig. 1'2. — Portion of forewing of Rhypltu-^ Itreris Walker (Diptera), showing
distribution of microtrichia and macrotriehia; most of the latter
have become detached from their bases of insertion; ( X 400).
Plate Ixix.
Fig. l.*i — Apical portion of forewing of Chorlsia anst rails King, (Mecop-
tera), showing distribution of microtrichia and macrotriehia; notice
the absence of the latter from the cros.s- veins; ( X200).
Fig. 14. — Portion of forewing of Blowfij^ [Galliphora n/losa Desv. ; Diptera),
showing the two types of microtrichia; the larger ones are on the
upper, the smaller on the undersurface of the wing; ( X 60).
(All figures are from photomicrographs, Figs. 9- 10 from the actual fossil.
Figs. 11-14 from cleared mounts of wings*.
52
658
A STUDY OF THE EXTERNAL BREATHING-APPA
RATU8 OF THE LARV^ OF SOME MUSCOID
FLTES.
By John L. Froggatt, B.Sc.
(Plate Ixx.)
Heretofore, in the identification of different species of liies
from maggots collected, the perfect flies have had to be bred-oiit.
But, in the course of the work of the Government 8heep-Fly
Experiment Station, this method was often impossible, when
packets of maggots were received dead or rotten. By means of
the microscope, I have found that an examination of the spiracles,
in particular the posterior spiracles, gives a sure and certain
means of identifying the different species here dealt with. As
their description has not been published heretofore, the results
are here recorded. In every case, the tj'pes were procured from
eggs deposited by each species of fly, on meat in the field -
laboratory.
The species dealt with in this paper are: —
Anastellorhina augur {Calliphora oceanuf^).
Pollenia stygia {Calliphora villosa).
Pycnoso7na rufifacies {Calliphora rujifacies).
Pycnosoma varipes (Calliphora raripes).
Lucilia sericata.
Ophyra nigra.
The spiracles, in all probability, exercise a most important
function in the destruction of maggots by poison. Careful ex-
amination of the skin does not reveal the presence of any pores;
the organs of the maggot may be looked upon as being enclosed
in an india-rubber-bag, the only openings into which are the
cfuUet and anus — the extremities of the alimentary tract — and
the anterior and posterior spiracles— the extremities of the
respiratory system. These anterior and posterior spiracles are
connected by two main tracheae, from which branches ramify
BY JOHN L. FKOGGATT. 659
throughout the body. Absorption will, in all probability, take
place slowly through the skin; l)ut the action of some liquids is
far too rapid to admit of this happening, so that, in these cases,
attention must be paid to the openings into the body. I am not
at present in a position to point out more than the possibilities
of this point, but it is receiving further attention.
The absence of pores in the skin of the maggots of these species
is extremely interesting as well as curious, because the larvae of
most other insects, and even diptera, show definite, well-marked
pores.
The anterior and posterior spiracles are totally different from
one another, the most marked specific differences being given by
the latter.
The anterior spiracles are situated one on either side of the
body, on the first thoracic segment. In shape, thev resemble a
cap with "a sharp upper edge, and a row of tassels along this edge.
The caps are chitinised pi-ojections of the two main tracheie on
wliich they fit Ijy a thickened ring; and it is from this demarca-
tion-line that all measurements are taken. In some species, the
cap is differentiated into a thicker lower part, and a thinner
upper part. The tassels or tubules are hollow, the open ends
ha\'ing a much-strengthened chitinous rim to keep the apertures
open. The number of tubules varies in different species, and
within certain limits in the same species, and even between the
right and left anterior spiracles in the same maggot.
The tracheae terminate posteriorly on the anal segment in two
spiracular plates or stigmata, the posterior spiracles. These are
slightly raised above the sui'face of the truncated end of the
maggot. The position of these plates on the body is such that
the bases of the plate, l.t^., the spaces in the band, point obliquely
tow^ards the median perpendicular plane of the body, and with
the median horizontal plane of the body passing through the
lower portion of the plates.
As both spiracular plates are alike for each species, a general
description of the characters of one will serve equally for the
other.
660 BREATHING-APPARATUS OF LARViE OF MUSCOID FLIES,
The plate is siirroanded by a strong band of chitin, varying in
thickness and in outline in different species. At the base is a
space or opening in the band; in Opliyra nigra, however, this is
closed up: and, in Pycnosoma rnjifacips and P. raripcs, it is open,
forming a break in the band. This space appears to carry some
soft structure in life, but which is destroyed in preparation.
Within this band are three structures radiating slightly from the
base; on subjection to strain, it is seen that these are slits, the
openings being crossed by bands of chitin, either straight or
anastomosing. The side-walls of the slits are xevy much
strengthened with chitinous blocks, and the 'ends are always
rounded. Thej^ are also concave from side to side.
In some species here dealt with, there is an intermediate struc-
ture lying between the middle and inner slits — "inner" referring
to the side nearest the median line of the body. This is a
shallow groove, somewhat V-shaped, coming to a more or less
definite point towards the bottom, and rounded at the top. The
rounded end encloses a clear space, the margin of which shows,
under a low magnification, a "sun-ray" effect. This is due to
fine bands of chitin radiating outwardly to the band. The inner
ends of these chitinous bands project over the rim of the clear
space, and would seem to support some soft structure in life, but
which is destroyed by the preparation. In some species, also, a
spaQe resembling a blister occurs on the side of the slits; when
present, it is always on the outer side of the outer and middle
slits, and on the inner side of the inner slit. This "blister-
structure" also shows the "sun-ray" effect referred to in the
' ' intermediate-structure. "
The length and breadth of all the spiracular plates include the
width of the band.
I am indebted to Professor Wheeler, of Harvard University,
U.S.A., for the method of preliminary preparation of the maggots.
Full-grown maggots are dropped into water heated to SO'C. A
few seconds in water at this temperature is sufficient to kill
them, and has the great advantage of stretching the maggots to
their full length; by killing in alcohol, the maggots generally
6Y JOHN L. FROGGATT. 661
contract, and often curve the l>()dy. The maggots are then
placed in 35% alcohol, and, after about 14 hours, they are trans-
ferred to 75,/ alcohol, in which they can be preserved. Asa
general rule, the maggots are left perfectly white by this prepara-
tion, excepting occasionally when they have been feeding on
niaterial rich in blood. When ready for preparation as micro-
scopic specimens, the body of the maggot is slit along the median
basal line, and boiled in 10% KOH until only the integument
remains. This is then put through the usual process for such
preparations, and mounted in Canada Balsam. All measure-
ments are in fractions of a millimetre.
The bibliography on the spiracles of dipterous larvye is, so far
as I have been enabled to trace it, very scanty. Figures of the
spiracles of different species have been given on a number of
occasions; but when a description is given, it is brief and with-
out detail.
Nielsen(l) figures the spiracles of certain species of Tahanidie^
but goes no further.
Froggatt(2) figures the spiracle of a dipterous larva found
parasitic in locusts at Minimbah Station, near Singleton, N.S.W.
A curious feature in this species was an elongated, cone-shaped
tail fitting over the anal portion of the abdomen. This was
doubtless to protect the spiracles, and prevent the maggot from
being smothered when the legs and wings were folded. The
perfect tiy was never bred out, so that it could not be identified.
GurneyO) figures the spiracles of three species of fruit-Hies,
and states that the identification of the different species can be
thus determined, but gives no description.
Banks(4) gives a slightly more extended account of the spiracles
of the different species he deals with, but gives very little detail.
He refers to what I am calling the space in the band of the anal
spiracular plate, as a "button"'; in which I do not agree with
him. He does not say how the spiracles were examined.
As extreme forms of spiracles, the following may be given.
Eristalis tenax, " The Common Drone-Fly," an introduced species
common in gardens, is the parent of curious rat-tailed larvae
which can live not only in putrid but even in salt water. The
662 BREATHING-APPARATUS OF LARV^ OF MUSCOID FLIRS,
posterior spiracles are projected into a long tail in order to enable
it to breathe while completely submerged.
The larvte of Stratiomya chamc^ho, another European fly
which swarms in water, have a similar, elongated, tail-like pro-
jection of the spiracles. The larva? of a Tachinid fly {Gymnosoma
rotwiidatum) live in and feed on tiie body of a Shield-Bug, and, in
order to breathe, have an elongated, hook-like projection of the
spiracles, which is pushed through one of the spiracles in the
side of the body of the bug.
Anasteij>okhina auguk. (Plate Ixx., figs. 4, 4a).
Anterior S'plracles. — Cap differentiated into two parts. Tu-
bules very strong.
Posterior Spiracles. — The band strong, complete, nut markedly
thick, showing no differentiation into two parts as in P. rujifacies
and P varipes. The band is thickened round the space which
projects the contour outwards at this point. The band is drawn
in slightly between the slits, giving the contour a slightly scal-
loped appearance. The inner margin of the band projects very
sli'ditlv between the outer and middle slits, and less marked! v
between the middle and inner slits. The slits are fairly close
together at the base, but are well separated otherwise, and do
not run quite the full length of the plate. The outer and inner
slits are fairly straight throughout their whole lengths. The
middle slit is straight for about two- thirds of its length, and
then inclines inwards. "Intermediate structure'" always present
between the middle and inner slits. " Blister-structure ' well
marked on all three slits.
PoLLKNiA STYGIA. (Plate Ixx., figs.6, 6«).
^?i<erior tSpirac'/es. — Cap differentiated into two parts. Tu-
bules broad compared with tlieir length, and strong.
Posterior Spiracles. — Band strong and complete. Space at
base completely enclosed in the band, which is not projected
outward to such an extent as in A. augur. The band is only
slightly drawn inwards between the slits. The inner margin of
the band projects strongly downwards between the middle and
I
BY JOHN L. FROGGATT. 663
outer slits, and somewhat less strongly between the middle and
inner slits; in the latter case, this projection usually forms a
hollow above the space in the upper end of the " intermediate
structure." Slits fairly close together, and running practically
the whole length of the plate. Outer and inner slits fairly
straight througliout their whole lengths. The lower half of the
middle slit is fairly straight, from which point it bends slightly
inwards. " Blister-structure " usually situated just above the
point of curvature. " Intermediate structure " always present
between the middle and inner slits, and well developed. In a
few cases, a similar structure has been noticed between the outer
and middle slits, but then the " blister-structure" on the middle
slit is absent. " Blister-structure" well developed on the slits.
The "sun-ray" effect is particularly well marked on the " blister-
structures" and on the space in the top of the "'intermediate
structure."
Pycnosoma rufifacies. (Plate Ixx., figs.5, 5a).
Anterior Spiracles. — Cap differentiated into two parts. Tu-
bules long.
Posterior Spiracles. — Band very strong, often differentiated
into two parts, the chitinous material being thinner on the outer
portion. It would seem that, when the maggots are young, the
differentiation in the band is most marked, but as development
is carried on, this differentiation decreases until the two parts
become either fused or the outer becomes the thicker and stronger.
There is, of course, no space between these two parts in the band.
The band is broken at the base, leaving the space open and in-
completely surrounded. The band is always much thicker at
the base. The inner edge of the band projects downwards
slightly between the middle and inner slits, and less markedly
between the middle and outer slits. The outer margin of the
band is regular in outline. The slits run the full length of the
plate, abutting on the inner edge of the band at the top and
bottom of the spiracular plate. The outer slit, in shape, is fairly
straight on the inner side, the outer side being slightly convex,
following the outline of the inner margin of tlie band. The lower
part of the middle slit is fairly straight, and close against the
664 BREATHING-APPARATUS OF LARV^. OF MUSCOIt) FLlES,
base of the outer slit. About the middle of its length, it bends
inwards slightly. " Blister-structure, " when present, generally
situated about the point where the inward bend takes place. The
outer edge of the inner slit is fairly straight; the inner edge fol-
lows more the contour of the inner margin of the band. It is
slightly convex. " Blister-structure" at approximately the maxi-
mum point of curvature. The " intermediate structure ' occurs
between the middle and inner slits, but, in many cases, it is but
slightly developed. The "blister-structure" is generally present
on the inner side of the inner slit, and absent on the other two
slits. It may, however, be seen on all three slits.
Pycnosoma varipes. (Plate Ixx., figs. 3, Sa).
Anterior Spiracles. Cap differentiated into two parts. Tu-
bules much shorter than in F. rufifacies.
Posterior Spirxccles. — Band extremely thick and strong, and
showing the same differentiation into two parts as in P. rufifa'-ies;
the same remarks apply equally in this case as in the previous
one. The inner margin of the band projects slightly downwards
between the outer and middle slits. The band is broken at the
base, leaving the space open and incompletely surrounded. On
the inner side of the spiracular plate, the band shows a develop-
ment of chitin like a square flag —the minimum distance between
the plates has been taken from the edges of these flags. The slits
run the whole length of the plate. 'J he outer slit is fairly
straight and regular throughout its whole length, although the
outer margin conforms more or less to the contour of the inner
margin of the band. The "intermediate structure" is only very
slightly developed between the middle and inner slits, and is
apparently only noticeable in the later stages of the growth of the
maggot. The "blister-structure" on the slits is absent.
LuciLiA sericata. (Plate Ixx., figs. 2, 2a).
Anterior Spiracles. — Cap not differentiated into two parts.
Tubules short and slender.
Posterior Spiracles. — Band fairly thin and entire, pinched in
between the slits as in A. augur. The space in the band is com-
pletely enclosed by the band, which is thickened at this spot;
Bt JOttN L. FROGGATT. 6(>5
the space generally tends to project the inner side of the band
inwards. Tliere is no projection of the inner margin of the band
between the slits. The slits radiate out from the base, and fill
the greater part of the spiracular plate, but do not run the full
length of it. In shape, the slits are all very similar, being, as a
rule, fairly straight and regular. The "intermediate structure"
is well developed, and always present between the middle and
inner slits. The "blister-structure" is always present on the
slits, and well developed. The blister seems, more noticeably in
this species, to be contained in a sheath which generally runs a
long way along the slit on either side of the blister: this is
generally more pronounced on the middle slit.
Ophyka nigra. (Plate Ixx., ligs.l, \a).
Anterior S]) ir acles.— Csup not divided into two parts. Tubules
long and slender.
Posterior Spiracles. — Band complete, and comparatively thick
and strong. It is differentiated, in the earlier stages of the de-
velopment of the maggot, into two parts, the inner part being
the broader and stronger. It is regular in outline. The space
is not in the band, but is surrounded by a development of cliitin
at the base of the plate, resting against the band. The space is
almost completely filled by a development of cliitin. The bases
of the inner and outer slits rest against this ingrowth of cliitin.
Compared with the size of the spiracular plate, the slits are small,
and appear to be well separated from one another. This is due
to each slit being surrounded by an envelope of hyaline material,
which envelopes are, however, lying up against one another. The
inner margin of the band does not, as a rule, project downwards
at all between the slits The outer and inner slits are usually
fairly straight and regular to almost the upper end, where they
bend over slightly outwards an3 inwards respectively. Thev
may, however, present a slightly wavy appearance. The middle
slit is always wavy in outline, with the tip of the upper end
bending slightly over. "Intermediate* and "blister-structures"
are absent.
666
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BY JOHN L. FROGGATT. 667
BIBLIOGRAPHY.
(1) Nielsen, J. C. — "Om Gyniuosoiiia. Aiteners Biolo^n (Dipt. Tachin).
Undersogelser over entoparasitiske Muscidelaiver lios Aithropodur
vi.
(2) Fko(;(;att, W. W.— " Eastern Plaj^^ue-Locust.'" Aj,aic. (Gazette N. S.
AVales, June, 1907.
(3) GuRXEY, W. B. — "Fiuit-FIies at Xaiaia." Aoiic. Gazette N. S.
Wales, 1912.
(4) IJanks, N. — "Structure of certain species of Dipterous Larwe, witli
particular reference to those in human Foods." U.S. Department
of Agriculture, Entoni. Bureau, Januarj% 1912.
EXPLANATION OF PLATE LXX.
Anterior and posterior spiracles of Ophyra iwjra (Figs. 1, la) — Lacilia
xe/'/c«/a(Figs.2, 2«) — Pycuo.soma raripes (Figs. 3, 8a) — Anastellorhina
f(n;/nr (Figs. 4, ia) — Pyriioxoiini rii/ifdcles (Figs. 5, oa) — Pollenia
■^fyijki (Figs, (j, G(/).
668
NOTES ON AUSTRALIAN SAWFLIES
( TENTHRhWIXIDJ:).
By Walter W. Fruggatt, F.L.S., (Iovernment Entomologist.
The Sawtiies are well represented in Australia by a number
of handsome insects belonging to very distinctive genera peculiar
to our insect-fauna.
While enormous numbers of the gregarious larva? of several
species of the genus Per^a are common at certain seasons (March
and April in particular), feeding upon the foliage of young gum-
trees {Eucalyptus) wherever there is plenty of young growth,
the perfect insects are comparatively rare. In breeding-out
specimens from the pupae, one finds that a large percentage of
them never reach maturity, because they are attacked in the
larval stage by many dipterous and hymenopterous parasites
which develop after the sawfiies have pupated. A great number
also fall victims to a mould-fungus, which destroys them in the
cocoons.
Perga dorsalis Leach : The Steel-Blue Sawfly.
Zoological Miscellany, iii., p. 11 7, t.l48, fig.l, 1817.
This handsome sawfly, one of the largest and best known
species of the typical Australian genus Perya, has an extended
range round the coast, and is common in Victoria and New
South Wales.
The gregarious larvje feed at night, and rest during the day,
clustered together in an oval mass, on the stem of the gum-tree
upon which they are feeding. When disturbed, they exude a
sticky yellow substance from the mouth, at the same time raising
the tip of the body, and tapping it down on the foliage. The
leaves are devoured from the top of the young gum-trees; and,
when the larvae are full fed, they crawl down the stem to pupate.
I have found them fully developed in the middle of April; but,
BY WALTER W. FROGGATT. 669
when they descend from their resting-place, thev wander about
over the grass for several days before thf^v finally select a place
in which to pupate, generally the softer soil against a tree trunk.
Into this they burrow to a depth of three or foui- inches, mass-
ing their large, oval cocoons in rows, one against the other I
watched several large swarms feeding upon the Peppermint-gums
{ Encalyptnii novce-angliff') at our Experiment .Station at Uralla,
and afterwards in their erratic wanderings over the grass; and
marked down their final resting place and dug up the cocoons.
At Binalong, in April, I observed two large swarms marching in
massed formation : the heads of the hind rows always rested
upon those in front as they moved along steadily together.
Every now and then, the front rank came to a dead stop, when
they all rested for three or four minutes; then a number began
raising up and tapping down the tip of the abdomen, whereupon
the whole band took up the motion: the leading ranks made a
fresh start, and all moved along again. In the largest band, T
counted two hundred and tifty caterpillars
Larvfe that pupated in the soil in the middle of April, pro-
duced the perfect sawflies in the Insectarium in the early part
of October.
The pupation of this sawfly is very remarkable. Under natural
conditions, the long, oval cocoons are formed one above the other
into a solid mass like honeycomb. When the full-fed hawse are
placed in a jar of sawdust, each forms a separate oval cocoon up
to ]J inches in length and J inch in diameter. The walls
are very thin, composed of a tough, sticky, papier-mache-like
material, the inner surface smooth and black, with the anterior
end rounded, and the apical cut off from a false end (containing
the cast larval skin loosely attached to the true cocoonj by a
slightly convex partition or cap of a greyish-brown colour, except
for the black centre just behind the head of the semipupa: the
outer side of this cap is fiat, black, and thickly impressed like
the surface of a thimble.
The first moult leaves a very wrinkled, pink and yellow, naked
pupa, with all the outlines of the larva, and the abdominal seg-
ments curled in underneath the hindlegs. This stage may
670 NOTES ON AUSTRAMAN SAWFLIES,
remain from two to three months, wlien the pupa undergoes a
second transformation. A fine, almost black, soft but closely
felted, fibrous, inner cocoon is spun, within which the pupa,
now an elongate, very wrinkled, white creature, without an}'
outlines of head or appendages, is enclosed. There must be a
third change, when the typical, true pupa develops, but this has
not yet been w^orked out. Probably, like some moths, this will
not take place until a month or two before the emergence of the
perfect sawfly.
Pterygophorus bifasciatus Brulle.
Hist. Nat. Insect., Hymen., Vol.iv., p.660, PI. 46, fig.l, 9, 1846.
This handsome species is easily distinguished from all the other
species by the dark marking on the forewings. The type, a
female, was described from Tasmania. Mr. Rowland Turner
informs me that it was unique; he had never seen this insect
until I sent specimens to the British Museum. My specimens
were collected in the pupal state: a colony, containing about
twenty cocoons imbedded in soft wood from the stem of an un-
determined tree, was sent by Mr„ Harold Brooks, from Dungay,
Tweed River, N.S.W. The larvie, when received in the cocoons,
were in a semi-pupal state, but showed that they were typical of
the genus Pterygophorus. The perfect sawflies emerged from
the cocoons early in September.
X. Shining blue-black, variegated with dull yellow. Length,
}. : exp. wings, | inch. Head and thorax, with the exception of
a yellow blotch on the sides of the latter, shining dark blue.
Legs black, variegated on the tarsi. Basal half of the dorsal
surface of the abdominal segments black, with the whole of the
ventral surface and apical portion of the dorsal surface yellow.
Forewings hyaline, richly variegated with chocolate-brown, form-
ing a clouded costal nervure; hind and marginal band, and an
outer transverse baud crossing the centre of the wing, so that
there are three semiopaque areas on the basal half of the wing
enclosed in clouded bands. Hindwings slightly fuscous.
9. Larger than ^J; of the same shining blue-black colour, but
having the face, a large blotch on the shouldeis, scutellum, post-
BV WALTER W. FKOGHATT. 671
scutelhmi, and tlie whole of the abdomen dull yellow. Antennae
composed of eleven joints; 1st and 2nd small, 3rd-10th rounded,
bead-shaped. Eyes large, projecting. Centre of the median
]oV)e of the mesothorax with a lateral depression. 8cutelluni
large, rounded. Length, ^: expanse wings, i inch.
Philomastix MACLEAii Westwood.
Perga macleaii Westwood, Proc. Zool. 8oc., 1880, p. 372, PI.
XXXV., f.2, (J. Philomastix ylaber Froggatt, Proc. Linn. 8oc.
N. S. Wales, (2;, Vol. v., p.489, 1890.
The type was described from a male specimen whose antennae
were wanting, and thus Westwood made the mistake of placing
it in the genus Perga.
I have collected a large series of both sexes, with the curious
double-tailed green larvae, upon the wild raspberiy-plants on the
Tweed River, N.S.W. When visiting the British Museum in
1908, I went through the cabinets of sawflies, and recognised mv
species under the name of Perga macleaii.
Pterygopiiorus analis G. Costa.
Ann. Mus. Zool. Napoli, Vol. ii., p.66, 1864.
A number of cases have been recorded from the Roma and
Mitchell districts, Southern Queensland, of the death of cattle
that have acquired the abnormal habit of eating the larv?e of
these sawflies. The sawflies appear in the early summer in the
open forest-country in enormous numbers, and deposit their eggs
on the foliage of the ironbark-trees. The resultant, long-tailed,
slender, green larvae are so numerous, that they completely strip
all the foliage off the trees over a large extent of the ironbark
forests. When full}' fed, they crawl or fall to the ground, and
congregate at the base of the tree-trunks in regular heaps.
The cattle running in infested country have acquired the habit
of licking up these moribund larvne; and quite a number of the
young stock, in these particular areas, have died from their par-
tiality to this change of diet.
Mr. Moore, the owner of "The Peaks," Marbango, Southern
Queensland, has sent me the following notes. " These cater-
672 NOTES ON AUSTRALIAN SAWFLIES.
pillars mature on the leaves of the Iron bark-trees, and, when
numerous, as thev were last season (1917), do not leave a leaf
on the trees. When full\^ matured, they come down and die
all round the trunks of the trees: and it is at this stage that
cattle lick them up: an overdose is fatal, particularly to young
stock, such as weaners, nine to twelve months old, and calves.
I fancy that acute inflammation of the bowels causes death: and
the beast appears to be in terrible pain towards the end. I am
inclined to think that a good deal of sand is licked up with the
caterpillars, and this may add to the irritation."
" The adult sawtlies emerge and are active all through April,
the caterpillars feeding upon the foliage through the winter. Tn
August, the full-grown caterpillars come down the trunks of the
trees, and die in heaps; and, for about three weeks, are a danger
to the young cattle in the paddocks. I think it must be a craving
for salt that attracts them, and we have laid rock-salt round the
trees, but once they have acquired a taste for the caterpillars
they will rush round the Ironbarks to lick them up.'
" r have had to remove all my cattle into the open country
away from the iron bark-forest, or my losses would have been
very heavv. As it was, I lost eighty head out of a mob of four
hundred, sixty weaners and twenty cows. Twenty per cent, is
very heavy in a week, and all the beasts that died were in
splendid condition; in fact the fattest seemed to suffer most."
In my opinion, this caterpillar-pest is going to prove a very
serious matter; and the only remedy will be to ringbark the
ironbark-trees in some of the paddocks, so that the cattle can be
kept away from the infested areas.
[Printed off, l)eoeml>er Utli, 1<>IS.
r XZ:^
.S.N.S.W. 1918.
16 9^
Carbouiferous Trilobites of Australia (P/u7fi>s(V/, (inffitlu<l,
PIS N.8.W. 191P
Cai'boniferouB Trilobites of Australia (P/n7/ii'!<i«) ,
"^.L.S.N.S.W, 191
CarbuiiUerous TriluLite.s of Australia (Griffithidcs, PhiUipsia).
PIS N.S.W. 191P
Carboiiit'erous Trilobites of Australiu (PhiHiiii<ia) .
L.S.N.S.W. 1QI8f
PL XLVIM
Carbouil'eroixs Trilobites of Australia (Griffithides, fhillipsia).
i
P.L.S.N.S.W. 19
l6Cx6)
I2M
13 ,
ii-bouiferous Trilobites of Australia (PltiUip>ii(i, Bniclnnnctopiiti) .
5(x4)
P.L S.N.S.W. 19I8-,
C'ai'bouiferoiis Trilobites of Australia [PhiUi))sia,'-Bra(:]i]iinetoi>us).
P...S.,N.S,W. t'J;3.
CarVjoiiiferous; Triloljites ui Avistialia (FltUliiinia, Brachyinetopus).
P L.S N.S.W. 1919.
3'
Cai'bouiferous Trilobites of Australia (PhilUpsiu, Biuchymetopus, Griffitliidesj.
8
Carboniferous Trilobites of Australia (Griffitkides, Brachynutoinis, Pldllipsia).
^
fL.S N.S.W. 191S.
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P.L.S.N.S W. 1918.
Volvocacese.
P.1.8 N.S.W. 1918.
Volvocaceae and Flagrellatse.
P.L.S.N.S.W. 1918.
Protoooccoideae.
'L.S N.S.W. 1918.
Spirogyra neylecta : a study iu variation.
"x
"^
Vi
,/
.L S.NS.W.
1, Ccelospermumpaniculatuiii F.v.ai.: (x24). 2. Milhtia australis F.v M.; (xl7).
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P.L.S.N.S.W. 19li
7. Bryonia laciniosa Linn.; (x-23). 8, Wistaria chinensis DC.; {XiO) ,
P.L.S.N.S.W. 1918.
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9. Fitis stercuZi/olia F.T.M.;(x40). 10. _6ougauifiltea spectabih's JusB,;(x23),
P.L.S.N.S.W. 1918.
vb. ''WiM
12
11, Bou'jainvillea lateritia Hort.; (x90). 12. B. 6andrrinn(L v. variegata Hort.; {x90).
^.L.S.N.S.W. iai8.
14
13. Piper nigrum Linn.; {x4^J) . 14. Galeola cassijtlioides Reichh.; (xl9)
'.L.S.N.SW 1918.
^
9.-10. ArchirKitiorixt ninginjicn TiU.
P.L S.N.S.W. 1918.
"^12
II. Profotheorapetrosema Meyr., (Lepiduptera) . 12. Rhyphris l>revis Walli.. (I>ii'tera),
P.L.SN.S.W. 1913.
13. ChoristaaustralisKlng, (Mecoptera). 14. CaZhp/iora r (7 ?oso Desv., (Dipteru).
p.L.s.N.SiW. loia..
E. H. Zeck, del.
Spiracles of larvse of Muscoid Flies.
673
NOTES ON THE NATIVE FLORA OF NEW 80UTH
WALES.
Part x. The Federal Capital Territory.
By R. H. Cambage, F.L.S.
(Plates Ixxi.-lxxiv.)
(Continued from Thesp Proceedings, 191^2, p.6ol.)
Synopsis.
Area and Situation .. .
Early Explorers in the Locality ...
Elevation and Topography
Geological Formations
Climate and Rainfall
Absence of Trees from Canberra Plains .
Insect- Action on Twigs and Buds
Plants absent from the Federal Territory
General Remarks on Various Species
List of Plants
Comparison with Tasmania
PAUE
. 673
. 673
. 679
. 680
. 682
. 684
. 688
. 688
. 689
. 701
. 709
Area and Situation.
The Federal Capital Territory contains an area of about 900
square miles, and is situated between latitude 35° and 36° south.
Its greatest length north and south is upwards of 50 miles, while
its average width is under 20 miles.
Early Explorers in the Locality.
The first explorer to reach the Federal Capital Territory was
Charles Throsby, of Bong Bong and Glenfield, and he was pro-
bably accompanied by Joseph Wild, a constable of the district of
Argyle, and a notable bushman, who discovered Lake George, not
many miles distant, on 19th August, 1820.
^3
674 NOTES ON THE NATIVE FLORA OP N. S. WALES, X.,
BY H. H. CAMBAGE. 675
In October, 1820, Lake George was visited by Governor Mac-
quarie, who then named it after His Majesty George IV. He
also named Goulburn Plains during the same visit. The Gover-
nor arrived at Lake Bathurst, from Parramatta, with Deputy
Surveyor-General James Meehan, Charles Throsby, Joseph ^^'ild
and others, and was there joined by Commissioner Bigge, Siii'-
veyor-General John Oxley, and Charles Fraser (Colonial Botanist),
who had journeyed from Bathurst up Campbell's River and
across the Abercrombie River.* The party travelled to Lake
George from Lake Bathurst, which latter was discovered by
James Meehan and Hamilton Hume on 3rd April, 1818 (Field
Book 143, Lands Department).
On 28th October, 1820, the Governor and party ascended a
high hill to the eastward of Lake George, evidently EUenden,
and viewed the high land in the southern portion of the Capital
Territory, though, owing to an intervening range, they would
not have been able to see the plains at the northern end. That
they were not impressed with the potentialities of the future
Federal Territory is evident from the following entry made by
Oxley : — "The whole extent between the S.E. and West may be
properly described as rocky, broken, and mountainous, and no
feature or object in the prospect offered any reasonable expecta-
tion that a good or even tolerable country could have existence
in those quarters." How little could they foresee that they
were gazing over what \Vas to become, not only the spot where
William James Farrer 'was to carry out his great wheat-giowing
experiments, which would do so much towards producing
drought-resisting wheats, and revolutionise wheat-production in
a dry climate, but that they were also viewing portion of the
site of the future capital of all Australia.
Macquarie refers to "anew-discovered river," and mentions
that " Mr. Throsby tried to get hold of some of the natives of
this part of the country to serve as guides to conduct them to
the new River Murrumbidgee," but the Governor's party could
* See Governor Maequarie's Journal — Report by Commissioner of In-
quiry Bigge (Mitchell Library). Oxley's Field Book No, 17"2, Lands De-
partment,
676 NOTES ON THE NATIVE FLORA OF N. S. WALES, X.,
not find time to visit it, though Throsby did. They were under
the impression that the new river flowed towards the south-east
into the ocean, which suggests that, although they were aware
of the presence of the river, no white man had actually visited it.
It was in April, 1821, that Throsby visited the locality of the
present Federal Territory, and, after going southerly from Lake
George, he wrote :— " I passed over two rivers, exclusive of the
one I discovered and on the banks of which I passed a night at
the time the Governor was at Lake George."
The two rivers referred to would be the Molonglo and Quean-
beyan, and the one he discovered previously, the Murrumbidgee.
He met with vast quantities of limestone, and a good quantity
of open forest and plains.*
On the 31st May, 1823, Captain Mark John Currie, Brigade-
Major Ovens, and Joseph Wild reached the vicinity of what is
now known as Queanbeyan, and, Currie writes, "encamped by
the side of the South Fish River (as called by our attendant,
Joseph Wild), on the edge of Lime-stone Plains," They travelled
thence south-westerly towards the Morumbidgee (as it was
usually spelt in those days), and named the Isabella Plain after
Governor Brisbane's daughter. This plain is largely included
within Portions 190 and 203, Parish of Tuggeranong. They
followed up the right bank of the Morumbidgee, finally crossing
the Umaralla, thinking it was the Morumbidgee, and discovering
the Monaro Plains or Downs. On the way up, they mention
having seen pine-trees of about 2 feet in diameter. These would
be the species now known as Callilris calca7'ata, Black or Moun-
tain-Pine.
In returning, the party for some distance kept to the eastward
of the Federal Territory, and, on the 8th June, 1823, "met with
large rocks of limestone," discovering what is now known as
London Bridge, " a natural bridge of one perfect Saxon arch,
under which the water passed."
Settlement followed this visit within the next few months, for
among the records of the Chief Secretary's Department is a letter
* The Australian Magazine, 1821, Vol. i. (Public Library).
BY K. H. CAMBAGE. b/ I
from Joshua John Moore, a retired Lieutenant, dated 16th De-
cember, 1826, in which he expresses his desire to proceed with
the purciiase of 1000 acres, " situated at Canberry, on the east
bank of the river, which waters Limestone Plains, above its
junction with the Murrumbeeja." He mentions that he had
been in possession of the land for upwards of three years. This
appears to be the first reference, in an official document, to the
name, which, with a slight alteration, was to be selected for the
capital city of Australia.
In a letter dated 14th September, 1831, Moore says :— "It is
called and known by the name of Canburry, and is bounded on
the south by the Molongoo River, and on the west by Canburry
Creek."
This area is now Portion 52, Parish Canberra, County Murray,
and permission to purchase was granted by Sir Thomas Brisbane
on 3rd May, 1825. There seems no doubt that the original was
a native name, but its meaning is unknown.
What is now Portion 58, of 4000 acres, Parish Canberra, and
known as Duntroon, was promised by Sir Thomas Brisbane on
the 18th May, 1825, and Portion 181, of 1000 acres, by Lieu-
tenant General Darling on the 22nd March, 1830, to Robert
Campbell, pursuant to instructions from the Right Honorable
Secretary of State for the Colonies, in part compensation for the
loss of a certain ship called " The Sydney '' while employed by
the Government of the said territory in the year 1806.
Portion 51, of 640 acres, Parish Canberra, was promised to
John McPherson on or before the 10th September, 1831, as a
Primary Grant. The name of the farm was Spring Bank, and
it was stated to be at Canbury, Limestone Plains
An area of 2560 acres, said to be at Yarralumla, now Portion
4, Parish Narrabundah, was promised by Governor Darling to
Edward Weston, of Horsely, Liverpool, on or before the 5th
September, 1831.
An area of 2560 acres, said to be at Yarralumla, now Portion
5, Parish Narrabundah, was promised to Henry Donnison on or
before the 15th August, 1828, by Governor Darling, but finally
granted to Terence Aubrey Murray and Thomas Walker.
678 NOTKS ON THE NATIVE FLORA OF N. S. WALES, X.,
Portion 203, of 2000 acres, Parish Tuggeranoiig, at Isabella
Plain, was promised to Peter Murdock on or before tlie 14tli
February, 1827; and Portion 190, of 2560 acres, adjoining, to
John McLaren, un or before the 21st February, 1829.
In May, 1829, Surveyor Robert Dixon traversed the Molonglo
River from near Queanbeyan, across the " Limestone Plains to
the Morumbidgee River." (Field- Book 317, Lands Department).
The position of the junction of the Queanbeyan and Molonglo
Rivers is shown, and on one side of the former is written
" Medora C/reek," and on the other "Quinbean,"
The position of "Tim Beards' Station" is shown on the south
side of Molonglo River near " commencement of plains."
To the west of where Canberra Church now stands, Dixon
shows the position of a fence, and the initials J.J.M., which
latter evidently refer to Joshua John Moore, the first owner of
of Portion 52, Parish of Canberra.
The entry, "Taylor's Hut," appears near the junction of
Yarrolumla Creek and Molonglo River.
On the 15th May, 1832, Surveyor Robert Hoddle commenced
a survey at " Malonglo River for the purpose of measuring land
at Limestone Plains." (Field Book, No. 375). Portions were
measured for Robert Campbell, George Thomas Palmer, Joshua
John Moore, John McPhei'son, John Stephen, and Edward
Weston.
He refers to Majoura, Ainslie's Hill, Black Mountain, Queen-
beeann, Pialligo, and II. Campljell's Cattle Station on Portion 181.
On the 16th May, 1832, when measuring J.J. Moore's Portion
52, of 1000 acres, Hoddle noted, on page 50, the position of
several "huts" and some " limestone rocks " at a bend in the
river at the spot since named Acton,* and, near them, entered
the name Canburry, which he afterwards altered to Canberry,
the latter name appearing on his plan. The site near the huts
is now occupied by an old stone cottage, built in the early days
* These identical lime.stone rocks are shown on the extreme left in
photograph No.l, in the "Report on a Geological Reconnaissance of the
Federal Territory" by D, J. Mahony, M.Sc, &c., and T. (Iriffith Taylor,
B.Sc, &c. (1913).
BY R. II. CAMBAGE. 679
of settlement. On the same day. lie traversed the creek which
flows south-westerly past the eastern slopes of Black Mountain,
and, on pages 52 and 53 of his Held-book, entered the name as
Canbury Creek.*
Elevation and Topography.!
The elevation of the Federal Capital Territory above sea-level
ranges from something under 2,000 feet along the valleys in the
northern portion, including part of the proposed Federal City of
Canberra, to upwards of 6,000 feet in the south-western portion,
the highest point being Mount Bimberi, which reaches an eleva-
tion of 6,264 feet, giving the Territory a range of elevation ex-
ceeding 4,000 feet.
The western boundary follows a range northerly from Mount
Bimberi to Mount Coree or Pabral, the elevation of the latter
being 4,657 feet Dividing the Cotter from the Murrumbidgee
River is another range running north and south, one of the
highest points of which is Tidbinbilla, said to be a slightly altered
native name which signified a snow-capped mountain, and which
reaches an elevation of 5,115 feet. These high points are all
visible from many spots on the north side of the Molonglo River.
The area east of the Murrumbidgee, which is regarded as a
fault-block, is generally lower than that on the western side,
and ranges between about 2,000 and 2,800 feet above sea-level,
the vegetation being of an open forest character. Around the
Federal City site and Duntroonare the Canberra Plains, formeiiy
known as the Limestone Plains, naturally almost treeless, and
through which the small Molonglo River flows.
The southern portion of the Territory is largely composed of
a network of broken mountains, though, in a few places, as near
* In 1837, Surveyor Hoddle had charge of the laying-out of the City of
Melbourne, and in 1851 became the first Survej^or-General of Victoria.
t See "Notes on the Physiography of the Southern Tableland of New
South Wales," by C A. Siissmilch, F.G.S., Journ. Proc. Roy. Soc. N. S.
Wales, Vol. xliii., p.331 (1909). Also "The Physiography of the Proposed
Federal Territory at Canberra," Commonwealth Bureau of Meteorology,
Melbourne, by Griffith Tajdor, B.Sc. (1910).
680 NOTES ON THE NATIVE FLORA OF N. S. WALES, X.,
Gudgeiiby, there are fairly level interspaces showing little or no
dissection, and in some cases containing swampy areas.
If we block out a model in the form of a great irregular
wedge, and consider the Molonglo River as the northern edge of
the wedge, a horizontal section the full width of the Territory
and extending south to the southern boundary would give us the
length of the wedge, the length of the base would be the width
of the Territory in the south, while the width (depth in this
case) of the base would vary from perhaps 3,000 to about 4,300
feet in the south-west corner. In viewing this great irregular
so-called wedge, which is higher along the western side, we find
it is scored longitudinally into more or less deep ravines, along
which flow the Cotter, Paddy's, Gudgenby, and Murrumbidgee
Rivers. The deepest of these gorges is occupied by the Cotter
River, which at Thomas Oldfield's (Portion 2, Parish Fergus,
County Cowley) is roughly 3,600 feet above sea-level, so that
the river has here, under Mount Bimberi, entrenched itself to a
depth of nearly 2,700 feet. The Murrumbidgee occupies the
shallowest of these channels, and, in parts of its upper portion,
around Tharwa, flows at the eastern foot of the mountains
through an almost mature valley, so far as the eastern side is
concerned, while it has cut gorges of varying depths down
stream.
Geological Formations.*
Quoting from Mr. Pittman's map, it may be said that the rocks
within the Federal City Site consist of Upper Silurian sand-
stones, quartzites, shales, tuffs, clay-slates, and several outcrops
of limestone, while the igneous rocks are crystalline tuffs and
lavas, quartz-porphyries, and quartz-felsites.
In the western and southern portions of the Federal Territory,
a considerable area is composed of granite of a fairly siliceous
character. Granite rocks are common around Tharwa, Boo-
* See a detailed "Geological Survey of the Site of the Federal Capital
of Australia," by Edward F. Pittman, A.R.S.M. (1910). Also, a "Report
on a Geological Reconnaissance of the Federal Territoiy," by D. J.
Mahony, M.Sc, &c., and 't. Griffith Taylor, B.Sc, &c. (1913).
feY R, H. CAMBAGE. 681
roomba, the Gudgenby River, in places alternating with slate, on
the range separating the Upper Gudgenby waters from those of
the Cotter, and on the summit of Birnberi Peak or Mountain.
No evidence of glaciation was noticed on Birnberi, but it was
not specially searched for. The absence of a suitable gathering
ground, however, would alone probably be sufficient to account
for its absence.
The valley of the Cotter, in the vicinity of Bimberi, and the
side of Bimberi, up to at least the 5,000 feet level, are of slate
formation, probably Upper Silurian, and this accounts for the
great denudation which has been effected by the Cotter River
operating on the softer rocks, for it was noticed that the highest
hills in the locality are granite.
The summit of Tidbinbilla was found to consist of quartzite
and slate, the former supplying the resisting qualities.
The central-eastern portion of the Territory consists largely
of quartz-porphyries, and produces an open forest vegetation.
Mount Coree is composed of igneous rocks, a specimen from
the summit being considered by Mr. G. W. Card, A.R.S.M.,
without critical examination, as of the rhyolite or rhyolite-tuft'
class. This rock splinters in a remarkable manner, and under
the bluff at the south-west end, narrow strips may be seen up to
10 feet long.
A few miles south of Coree, and also where the main road
crosses Coree Creek, the forn)ation is slate.
A good example of the effect of geological formations on the
vegetation may be seen from the City site. To the north east
are the slopes of Mount Ainslie (2,762 feet), composed largely of
volcanic tuffs, and clothed with open forest, while to the north-
west is the coarse sandstone-hill known as Black Mountain
(2,658 feet). The name of the latter was suggested by the dark
appearance of the more dense foliage of this hill as compared
with that on the surrounding elevations, but this mass of vege-
tation is a direct response to the more siliceous sandstones of
wliich the eminence is composed.
682
NOIES ON THE NATIVK FLORA OF N. S. WALES, X..
^ Climate and Eainfall.*
If tlie flora of the area be classified under the heading of either
a warm- or a cool-country flora, its proper place is certainly under
the latter. There are a few western or warmth-loving plants
found there, one in particular {Casuarina Luehmaniii) raising
an interesting point in the study of distribution, but the great
bulk of thera are such as may be expected in our cool mountain-
areas. Judging from the native vegetation, therefore, the
climate of the Federal Capital Territor}^ may be designated as
cool.
According to the Commonwealth Bulletin No. 7, and further
information kindly supplied by Mr. H. A. Hunt, Commonwealth
Meteorologist, and Mr. D. J. Mares, Divisional Officer, Sydney,
the following are the mean temperatures at a few stations, and
the average annual rainfalls at some localities in and around
the Federal Territory, including, for comparison, Cootamundra
towards the foot of the western slopes, and Milton to the east
on the coast.
Wettest
Driest
Years.
Rainfall
Years. "'
Mean
month.
month.
in inches.
temp.
Braidwood
January
September
39
27 3
55 '4
Buiigendorc ...
January
May
27
23 0
— '
—
Carwoola
June
February
27
24-9
—
—
Collector
—
. —
17
26-4
—
—
Cooma
February
August
02
191
44
54-2
Cootainu'idra ...
Jiuie
February
28
22-9
16 '
59-6
Duiitroon
June
February
19
20 1
4 :
56-6
Goulburn
JaiRiary
April
-yZ
25-0
46
56 1
Gudgeiiby
January
February
27
31-8
—
—
Guudaroo
June
February
40
24-2
—
Kiandra
June
February
42
63-8
— '
44-4
Lake George ...
j June
February
33
26-4
19
58-1
Majura...
January
February
30
24 1
— ,
—
Milton
i
25
44-2
—
—
Quean Ue^-au ...
January
August
40
22-3
—
561
Uriarra
—
15
32-9
i
—
From the above, it may be seen that January and June are
* See Bulletin No. 7, "On the Climate of the Yass-Canberra District,"
by H. A. Hunt, Commonwealth Meteorologist. (1910).
BY R. H. CAMBAGE. 683
the wettest months, while Fehruai-y is usually the driest. The
annual rainfall at the Capital City site may be estimated at
somewhere about 21 or 22 inches.
The effect of climate upon the native vegetation is largely
regulated by the question of aspect. Broadly speaking, Eastern
New 8outh Wales has two dominating aspects, the eastern or
moist, and the western or dry.
The rain comes chieHy from the eastern or ocean-side, and is
precipitated by the cooling or ascending clouds on the mountain-
sides and summits; when the clouds pass beyond the summits
and commence to descend, the precipitation is reduced. This
applies also to clouds coming from the west. The result of this
natural law may be seen in the Braid wood district, about 40
miles nearer the coast than Canberra, and at a distance of only
35 miles from the ocean, for Braid wood is largely shut off from
full coastal influence by a mountain-range a dozen miles away
and known as Budawang, to the east of which the rainfall is
(juite 40 inches, while at Braid.wood it is only about 27 inches.
Similar conditions are found at many places towards the eastern
edge of the plateau in this State.
A great portion of the Capital Territory, especially along the
depressed iMurrumbidgee valley, is sheltered by north and south
ranges on either side, as well as for the most part on the south,
the result being that, to a large extent, it is the descending clouds
from east, west, and south which pass over this area, and the
rainfall in the valley is, in consequence, less than on the hill-
sides. Evidence of this may be seen in the resultant flora.
Moreover, the clouds from the coast, after passing over the
Territory, encounter the high range known in part as Brinda-
bella Mountain, forming the western boundary extending from
Bimberi to Coree, and on its slopes deposit much of their
load. Asa result, this is the area in which is found the most
robust and comparatively luxuriant vegetation in the Capital
Territory, including splendid examples of tree-ferns (Dicksoiiia
antarctica), ten feet high, which, at about 3,000 feet above sea-
level, are nestling under the shelter of Mount Coree, and facing
the eastern or moist aspect.
684 NOTES ON THE NATIVE FLORA OF N. S. WALES. X.,
My view is that the luxuriant forests on the mountain side
are tlie result of a high rainfall induced chietly by the position
of such mountains, rather than that the rainfall is a consequence
of the presence of the forest. Judging by the vegetation on the
higher land towards Bimberi and Coree, it is probable the annual
rainfall there reaches 40 inches, and on Bimberi itself, which
intercepts clouds from every direction, it possibly amounts to
quite 50 inches
Absence of Trees from Canberra Plains.
The reason why many thousands of acres of almost level or
slightly undulating land should be naturally destitute of trees is
difficult to explain. The question is a universal one, and Aus-
tralian examples have been much discussed between botanists.
The instances are many, and include those from the extensive
plains of the interior with a meagre rainfall, whei-e treeless
stretches of upwards of fifty miles are not uncommon, to those
on the highlands with a rainfall, in exposed situations, as around
Kiandra, of sixty inches per annum, and also such spots as those
on the upper Dorrigo, where the brush or jungle in places ceases
abruptly and forms a fringe on the edge of an open plain.
My own observations lead me to suggest that the explanation
will yet be found in many cases by an examination of the soil,
or, in other words, that it is from this source that we shall obtain
our best knowledge of the subject, studied in relation to topo-
graphy, rainfall, and aspect. Whether the feature is chiefly
regulated by the chemical constituents or the physical properties
of the soil is difficult to decide, but it is probably in some degree
the result of both factors.
In These Proceedings (1909, p. 3 10), I discussed the question
of the absence of trees fiom the Monaro Tableland, and pointed
out that, where the hills were composed of siliceous formations,
they were more or less tree-clad, while those made up of basic
soils were bare, except for some scattered trees of Eucalyptus
coriacea and Acacia ruelaHoxylou on a few basalt-summits.
BY R. H. CAMBAGE.
685
T am indebted to Mr. J. C. H. Mingaye, F.C.S, for the follow-
ing analyses of soils and quartz-porphyry from the Canberra
Plains,
A. From crest of low, treeless elevation at 6;^ miles from
Queanbeyan.
Mechanical Analysis.
Coarse particles of stone ... ... ... 19 *o8 per cent.
Stone left on 20-mesh sieve 9 "87 ,,
Stone left on 30-mesh sieve 3 '92 ,,
Stone left on 60-mesh .sieve 7 '83 ,,
Root-fibre 0-59
Chemical composition of soil passing through 60-mesh sieve
soluble in warm hydrochloric acid : —
Insoluble in acid
82-04%*
tFerric oxide alumina ...
10-56%
Lime (CaO)
0-06%
Silica iSiOo)
0-25%
Magnesia ( Mg( ) )
0-16%
Potash (K^O)
0-34%
Soda (NaaO)
0-13%
Phosphoric anhydride (P., Og )
0-095%
Manganous oxide (MnO)
trace
Water
5-70%
Organic matter
0-81%
100 145
* Containing silica 78-21%.
t Including a small amount of Titanium dioxide (TaOj).
B, From shallow valley at 5 miles from Queanbeyan.
Mechanical A nalysis.
Coarse particles of stone 0*11 per cent.
Stone left on 20-mesh sieve 3 '07 ,,
Stone left on 30-mesh sieve ... ... ... nil
Stone left on 60-mesh sieve 9-38 ,,
686
NOTES ON THE NATIVE FLORA OF N. S. WALES, X.,
80-11%*
5-13%
012%
0-61%
0-29%
0-88%
0-13%
0-()7%
trace
3-38%
0-51%
100-23
Chemical composition of soil passing through 60-mesh sieve
soluble ill warm hydrochloric acid : -
Insoluble in acid
fFerric oxide alumina ...
Silica (SiO.,)
Lime (CaO)
Magnesia (MgO)
Potash (K2O)
Soda (Na.,0)
Phosphoric anhj'dride (P„Oji)...
Manganous oxide (MnO)
Water
Organic matter ...
* Containing silica 76-62%.
t Including a small amount of titanium dioxide (T.^O.J.
The major portion of the soil which passed through a 60-mesh
sieve, and used for the analysis, consists of fine particles of
broken-up stone
No. 2124/18. Quartz-porphyry, Canberra Road, on treeless
plain 2| miles from Queanbeyan.
Chemical Composition.
Moisture at 100°C
Water above lOO'C
Silica (SiOa)
Alumina (AloO-)
Ferric Oxide (Fe.,03)
Ferrous oxide (FeO)
Manganous oxide (MnO) ...
Nickel and cobalt oxides
(NiO-CoO)
Calcium oxide (CaO)
Magnesium oxide (MgO) ...
Barium oxide (BaO)
0-52
Soda(Na„0) r37
2-66
Potash (K.,0) 2-76
64-64
Lithia (LoO) absent
14-49
Titanium dioxide (TiOa) ... 0'65
2-30
Zirconium dioxide (ZrOa).. absent
3-69
Iron sulphide (FeS 2) ... absent
0-12
Sulphur trioxide (SO3) ... 0-08
Phosphoric anhydride (PoOe) O'll
001
Vanadic oxide (VoOy) minute trace
3-42
Chromium sesquioxide (Cr^Os ) ditto
2-93
Chlorine (CI) absent
0-06
Strontium oxide (SrO)
. present"
99-81
* Spectroscopic reaction only. Specific gravity of rock -2-735.
The soils of these plains around the Federal Capital are sili-
ceous rather than basic, and the feature which is noticeable at
Canberra, and other similar plains, is, that if there are any con-
BY R. 11. CAMBAGE.
687
siderable elevations, of say, 200 feet or upwards, rising above
the plain, then such elevations produce trees. The inference is,
therefore, that there is some difference, either chemical or physi-
cal, between the soils on the well drained hills, and the soils of
Text-tig. 2.
Buds of Eiiralypftt.s dea/hafa enlarged by dipteious laivie.
the lowland; and it may be that, even though the geological
formation is the same from which the soils of hill and valley are
originally produced, certain salts are leached out from the high
land and carried down to the lower, thus differentiating the
characters of the two soils.
688 NOTES ON THE NATIVE FLORA OF N. S. WALKS, X.,
Tt is usual, of course, for both valley and height to produce
trees, the differences in soil accounting perhaps for different
local species, but this does not apply to the Canberra Plains,
which, for some unexplained reason, are for the most part tree-
less, though all the considei-able elevations around are clad with
forest-growths.
Insect Action on Twigs and Buds.
Some gouty swellings were found at the Cotter River on fruit-
ing twigs of Encalyptus hcemaatoma (Brittle Guni), which, Mr.
W. W. Froggatt, F.L.S., informs me are caused by the attack
of small chalcid wasps (Chalcididse), some of wiiich are plant-
feeders and deposit their eggs just under the bark. The effect
on these twigs was such that, in some cases, for a length of 15*3
cm. (about 6 J inches), they had been increased in diameter from
2"5 mm. to 16 cm , or about 6^ times their original diameter.
In November, 1911, great numbers of very interesting insect-
galls were found within the City site on manv trees of Euca-
lyptus dealhata (Red Gum). The flower buds were aborted by a
dipterous larva which Mr. Froggatt has kindly identified as
belonging to the family Agromyzidae. By the action of these
larv£e, the buds had been increased in diameter from 2 mm. to as
much as 1'2 cm., or six times their original size (Text-fig. 2). In
some cases, five out of six buds in the umbel were affected. The
effect of this swelling of the buds so greatly increased their
weight, that broken branches were to be seen in many directions,
reminding one of the result of a snow-storm.
Plants absent from the Federal Territory.
Owing largely to climatic reasons, several groups of plants
are absent from this district, the locality being too cold for
them, though, in more northern latitudes, they may ascend to
greater elevations than 2,000 feet. No species of Angophora,
the coastal Apple-Tree, was seen, and its absence from the south-
western district was commented upon by Hume and Hovell, in
their overland-journey in 1824 {ibid., p.87).* No representative
* For previous remarks in regard to distribution of this species, see
these Prooeediugs, 1905, xxx,, p. 207.
BY R. H. CAMBAGE. 689
was seen of that large genus Melaleuca, which generally prefers
a wanner climate. The whole of the Ironbark-trees are absent,
these forming a group which avoid the cold, no species of tiue
Tronlmrk occurring in Tasmania. No species of White Box was
noticed, not ev^en Eucalyptus <dhens, which creeps up the western
slopes wherever it can find sufficient warmth, and may be found
in isolated cases near Yass. The absence of E rostrata, the
Murray or River Red Gum, was noticed, but I was informed
that it ascends the Murrumbidgee to Umburra, some few miles
below tlie Federal Territory, though it is unable to face the cold
within the Territory itself. E. globulus, the Tasmanian Blue
Gum, was not seen, although the climatic conditions are suitable,
and it occurs lower down the Murrumbidgee, at Burrinjuck, and
may possibly yet be found in the valley of the Cotter.
Genkkal Remarks on various Species.
The notes for this paper were obtained during short visits to
the locality in November and December, 1911, and January,
1912. In addition to the area immediately surrounding the
City site, the routes examined were the following : — Canberra
to Queanbeyan and along the Bungendore Road; the Gundaroo
Road; Canberra to the junction of the Cotter and Murrumbidgee
Rivers, and up the Cotter just above the dam; Canberra to the
summit of Mount Coree; to Tharwa, Booroomba, and the summit
of Mount Tidbinbilla; Tharwa to Gudgenby, thence up Middle
Creek across to the Upper Cotter, and to the summit of Bimberi
Peak. It will be seen that there were many spots not visited,
so that the list of plants must be regarded as incomplete.
Mount Tidbinbilla. — Within a radius of 20 yards around the
actual summit of Tidbinbilla (5,115 feet), the following plants
were noticed : —
Gramine^e : Poa crespit'jsa (Snow-Tussock or Snow-Grass).
JuNCACEiE : Luzula campesfris.
LiLiACEiE : Bulbiiie bulbosa, Dianella tasmanica.
OrchidacEjE : Caladenia dimorpha.
CARYOPHYLLACEiE : SteUaria pwni/eus, Sderanthns bifiorus.
54
690 NOTES ON THK NATIVE FLORA OF N. S. WALES, X.,
LEGUMiNOSiE : Acacia penninervis (Mountain Hickory), Oxy-
lobium procu77ibens{1), Daviesia ulicina, Hovea linearis.
RuTACK^ : Eriostenion myoporoides.
ViOLACEiE : Viola hetonicoifolia (Native Violet).
THYMELiEACEiE : Pimelea sp.
MYRTACEiE : Eucalyptus coriacea (Snow-Gum), Kunzea pechm-
cularis, Callistemon lophanthiis, Bceckea Gunniana.
Epacridace^ : Leucopogon hiflorus, L. Fraseri (prostrate),
Acrotriche aggregata.
ScROPHULARiACEiE : Vcronica perfoliata.
GooDENiACEiE : Goodcnia hederacea.
CANDOLLEACEiE : Candolha serrulata (Trigger-Flower).
C0MPOSIT.E : Br achy come sp., Helipterutn incanurti, Microseris
Forsteri.
Bimberi Peak. — Writing from memory, the summit of this
granite-mountain is approximately a quarter of a mile long, by
about 200 or 300 yards wide, the highest point being 6,264 feet
above sea-level.
On viewing the flora of this elevated spot on the 15th January,
1912, the feature which impressed me most was the high colour-
ing of the flowers and their great numbers. Masses were to be
seen of flowering examples of Brachycome scapigera (a Yellow
Daisy), and these were blended with others of Senecio, Podolepis,
and Helichrysum, the hill being charmingly crested and bright-
ened with a profusion of yellow and white, distributed amongst
a groundwork of countless, graceful, grey flowers of the Snow
Grass.
The only species of Eucalyptus found on the summit was F.
coriacea (No. Si70), occurring as spreading, dwarfed trees of from
1 0 to 20 feet high, and flowering, the branches being intensely
glaucous. This species grows at a slightly higher level on
Kosciusko, but it is doubtful if any other Eucalypt grows at an
elevation exceeding that of Mount Bimberi. This was also the
only Eucalyptus found on the summits of Tidbinbilla and Coree.
Helichrysum ledifolium was seen only on the summit, and this
species, which occurs in Tasmania, had not been previously re-
corded for New South Wales.
BY R. H. CAMBAGE. 691
The following is a list of plants noticed on Bimberi, and in
most cases also collected, above the 6,100 feet level, though doubt-
less several species were overlooked : —
PoLYPODiACEiK : Polystichum aculeatum (a common fern in the
cold areas of Southern New South Wales).
GRAMiNEiE : Poa ccespiiosa (Snow-Tussock, or Snow-Grass, in
flower all over the summit).
RESTiONACEiE : Festuca Hookeriana, Hypolcena lateriflora (a
weak, straggling, wiry plant of a few feet high, growing in masses
in damp or swampy places, and slightly resembling Caiistis flex-
uosa, a common Sydney plant).
ORCHiDACEiE : Thelymitra venosa (a blue Orchid).
PROTEACEiE : O^ntes lancifolia, Grevillea aitstralis (a very nar.
row-leaved form. This is the only Grevillea which occurs in Tas-
mania, the genus being one which favours a warm climate).
PiTTOSPORACEiE : Mariauthus procumhens.
PoLYGALACEiE : C omesperma retusum.
THYMKLiEACEiE : PimeUa ligustrina.
MYRTACEiE : Eucalyptus coriacea (Snow-Gum), Callistemon
Sieberi (the flowering was just over), Bceckea Gunniana.
Umbellifer^ : Aciphylla si77iplici folia.
EPACRiDACEiE : Epacris paludosa, E. microphylla, Richca
Gunnii.
Labiat^e : Prostanthera cuneata (with whitish flowers).
RuBiACEiE : Asperula oligantJia.
CoMPOSiTiE : Olearia stellulata, Celmisia longifolia (Silver
Daisy), Brachycome scapiyera, B. discolor, Podolepis longipedata,
Leptorrhynchos squamatus, Helichrysiwi scorpioides, H. ledi-
folium, Erechtites quadrideutata (a broad-leaved form), Senecio
pectinatus, Microseris Forsteri.
Thirteen species of ferns were found within the Territory, the
most of them being in sheltered portions of the mountain-slopes.
Of the Graminese or grasses, twenty-one species were noticed,
four of which were naturalised. Probably several native species
escaped notice during my hurried visits. Boa ccespitosa, the
Snow-Grass or Snow-Tussock, is an interesting plant in view of
692 NOTES ON THK NATIVE FLORA OF N. S. WALES, X.,
its very wide range in Australia and New Zealand, and its
adaptability to environment. Along the coast it is often known
as " White Tussock," because of its pale grey colour, and is
regarded as an indication of good forest-land; it may occur
within a short distance of the ocean, and sometimes, in favoured
situations, grows into robust plants of three to four feet high.
It shows its disregard for climatic effect b}^ climbing from sea-
level to upwards of 6,000 feet, where, owing to the influence of
more rigid conditions, it becomes matted and dwarfed, losing-
much of the tussocky form, and in summer is most useful as
sheep-fodder.
Among the Liliacese, the somewhat succulent little plant,
Bulhine bulbosa, was found in various places, including the
summit of Tidbinbilla. This species has an exteirded range in
Eastern Australia and Tasmania, and in addition to being found
at elevations of 5,100 feet, as in this case, its yellow flowers are
conspicuous every spring in the much drier interior at such
places as the Macquarie and Lachlan Rivers.
The little terrestrial orchid, Caladenia dimorpha, was only
noticed within a few yards of the actual summit of Tidbinbilla,
while C. alba was seen a few hundred feet lower,
Casuarina stricta (She-Oak) was found on various hills, in-
cluding Ainslie, Majura, Stromla (2,560 feet), and Mugga Mugga
(2,662 feet), near Tharwa, and also to the west of the Naas River
on the ascent to Gudgenby, where it was growing on the northern
or warm side of granite-hills at elevations up to 3,000 feet. I
have not found it at an altitude exceeding this.*
Casuarina Cunnhighamiaiia (River-Oak) occurs along the
banks of the Murrumbidgee up to within about four miles of
Tharwa; above this point, the country is evidently too cold for
it. This is an attractive-looking tree and always grows within
reach of fresh water, but is restricted in the extent of river it
will follow, by the degree of cold in the highland and warmth
in the lowland. This Oak-tree may be found on the rivers from
Tropical Queensland southerly to the Murrumbidgee and its
For previous remarks, see These Proceedings, 1909, Vol. xxxiv., p.;^26.
By k. h. cambage, 693
tributaries, but is not recorded from Victoria. The first refer-
ence to its absence from the Murray or Hume River is tliat by
Hovel] and Hume in the report of their exploration fi'om Goul-
burn to Port Phillip.* When referring to the river, their note
reads : " but there was no swamp-oak, the tree so universal on
the rivers to the northward and eastward " (p.44).t
A small clump of Casuariua LueJnnanni^ the Bull Oak of the
interior, was found near where the Gundaroo Pvoad, at about 2^
miles from Queanbeyan, passes the Molonglo River. The trees
were confined to a high, steeply sloping, volcanic tuff" bank of from
40 to 70 feet deep, on the southern side of the River, and, while
facing a northern aspect, were well sheltered from the cold south-
erly influence (Plate Ixxi.). Although the branches and stems
of some of the Oak-trees reached above the bank, it was noticed
that their bases were all below the summit, which signifies that,
in the early seedling-stage, the plants require shelter to allow
them to become established. -The finding of these trees in the
Territory was a matter of great surprise, for the nearest locality
where they are known to me is between Cootamundra and
Temora, a distance of about 80 miles in a direct line, though
some may possibly have occurred at intervening spots along the
valley of the Murrumbidgee. The species is one having a very
wide distribution, extending from the south-eastern portion of
South Australia, across part of Victoria and the whole of Central
and part of AVestern New South Wales, continuing along the
eastern portion of Queensland at least as far as Biboohra, west
of Cairns, in latitude 17°; and, in places, pushing through low
gaps in the mountains, towards the east coast, but usually select-
ing a warm climate. It comes through the Cassilis Geocol and
down the Hunter Valley to near Ravens worth and Pokolbin,
where it overlaps the coastal salt or brackish-water Swamp-Oak
(C. ylaiica), and it may be seen from the train between Bunda-
berg and Gladstone in Queensland. In 1899, a few trees of
* Journey of Discovery to Port Phillip, N. S. Wales, in 1824 and 1825,
by W. H. Hovell and H, Hume.
t For some previous remarks, see Journ. Proc. Ro3^ fSoc. N. S. Wales,
Vol. xlix., p.399(1915). Also, These Proceedings, 1901, Vol. xxvi., p. 685.
694 NOTES ON THE NATIVE FLORA OF N. S. WALES, X.,
dwarfed Bull oak were seen on the southern bank of the Fish
River, near O'Connell, in the Bathurst district, and these were
growing under somewhat similar conditions, and at approxi-
mately the same elevation, about 2,100 feet, as those under dis-
cussion on the Molonglo River, which are also rather diminutive
specimens, ranging from 12 to 20 feet high, with a stem-diameter
up to 9 or 10 inches in a few cases.
The question that naturally arises is, how did these trees find
their way to this spot so far froui their congeners 1 Dispersal
of seeds by wind can probably be ignored in this case, and dis-
persal by birds, though much more likely, can scarcely be shown
to account for it, although this possibility calls for consideration.
There are very few spots within the Federal Territory where
seedlings of this species Avould survi-ve without special care,
owing to the coldness of the winter climate. The birds which
chiefly feed on Casuarina seeds by tearing open the small cones
are of the cockatoo-family, Calyptorhynchus viridis, the Glossy
Cockatoo or so-called Macaw, and there would be nothing re-
markable in finding that one of these birds had visited both the
Cootamundra and Queanbeyan districts within a few days. At
the same time, there is the coincidence to be accounted for that
a seed should be deposited in one of the few spots which would
result in the production of a mature tree. It must also be borne
in mind that this is a dioecious species, the male and female
flowers occurring on separate trees, so that it would be necessary
that seeds, producing a pair of trees, should reach the same
locality before the species could become established.
It may be considered possible that the seeds were conveyed
by natives, but the seeds ripen in midsummer and fall out of the
cones within a few days (usually two), after being gathered, and
the natives would, therefore, have required closely woven bags
to retain them.
The genus is known to be an ancient one, having been identi-
fied in fossil form in the Tertiary flora. ''^ This particular species
* Ettingshausen, " Contributions to the Tertiary Flora of Australia,"
p. 107.
nv R. H CAMBAGE. 695
is probably old, considering its wide distribution, and if repre-
sentatives had been growing in this vicinity, at the time of the
Eastern Australian uplift in late Tertiary time,* which seems
quite possible, some plants may have survived, during the gradual
upheaval, and their descendants, on finding the new conditions
too cold, would gradually have become restricted to the warmer
and more sheltered nooks where the geological formation was
favourable to their growth, and would, thereby, come under the
heading of relics or stranded plants. A difficulty about accept-
ing this explanation is that this Oak is a warmth-loving species,
and, at an elevation of 2,000 feet, would have found it difficult
to survive the Pleistocene glacial period which is generally re-
garded as of subsequent date to the uplift that formed the pre-
sent mountains. The fact is clear, however, that these particular
Oak trees are now growing near the Molonglo River, and although
they certainl}^ appear to be stranded plants, the process by which
they reached this spot must remain unsolved, at least for the
present.
Grevillea juniperina is one of the most attractive shrubs
within the Federal Territory, chiefly because of its beautiful,
red, spider-like flowers. It was noticed on the right bank of the
Murrumbidgee near its junction with the Cotter, growing as
thick spreading bushes up to 10 feet high (Plate Ixxii.). It was
flowering in November, and among the birds, which were evi-
dently after its honey, were the Leatherheads {Tropido7'hy7ichus
corniculatus). At Mongarlowe, near Braidwood, this species
was seen in November, 1908, with yellow, as well as red flowers,
and in several cases both colours were noticed on the same plant.
The feature has been observed by others.
Batiksia marginata (Honeysuckle) was seen in many portions
of the Capital Territory, ascending to an elevation of 4,000 feet,
and occurring chiefly in the granite or somewhat siliceous areas.
The trees appear toreach greater dimensions than do those of
this species in the Sydney district, and examples were seen with
* "Geographical Unity of Eastern Australia," by E. C. Andrews, B,A.,
Journ. Proc. Roy. Soc. N. S. Wales, Vol. xliv., p. 420(1910). —Presidential
Address by C. Hedley, F.L.S., These Proceedings, 1911, Vol. xxxvi., p. 13.
696 NOTES ON THE NATIVE FLORA OF N. S. WALES, X.,
trunks up to two feet in diameter and a heiglit of twenty feet.
Some of the finest are growing in granite-formation around JBoo-
roomba (Plate Ixxiii.). Although this Honeysuckle always
avoids soils derived from basic rocks, it has a wide range, and
is doubtless the species referred to by Hume and Hovell as
occurring on the Yass Plains in 1824.
The genus Loranthus (Mistletoes) appears to be only sparsely
represented in the Territory, and is practically confined to the
levels below about 3,000 feet. The Loranthus is not a lover of
extreme cold, and its general absence from part of the southern
hiiihlands was noticed some vears af^o,* while there is no record
of the genus occurring at all in Tasmania.
Two species of Drosera (Sundews) w^ere collected, and from
the paper in which they were pressed, it has since been noticed
that when drying, specimens of D. peltata from Gudgenby at
4,500 feet, dyed the paper pink leaving the impression of stems
and flowers.
Of the family Leguminosie, fifteen species of Acacia (Wattles)
were seen, and next after the Eucalypts this was the greatest
number of species found of any genus.
The plants identified as A. obtusata, from Black Mountain,
sometimes reach 7 or 8 feet high. Around Mount Coree, they
are locally known as Blue Wattle, from the slight colouring some-
times appearing on the leaves and stems, though on many plants
the bark is reddish-brown.
Acacia pravissima grows to a height of from 6 to 10 feet, with
somewhat pendulous branches. The pods ripen during the latter
part of December.
Oxylobium alpestre was seen as spreading shrubs, semi-pros-
trate, above the 5,500 feet level on Bimberi, and it was noticed
that the womhats {Phascolomys mitchelli), which are fairly plenti-
ful at many places on these highlands, had undermined many of
these plants, upon the roots or possibly root-nodules of which
they evidently feed.
* " Eastern Monaro," by R. H. Cambage. These Proceedings, 1909,
Vol. xxxiv,, p. 331.
CY R. H. CAMBAOE. 697
Tetratheca ericifolia was found near Coree and Booioomba.
It has been noticed, over many years of collecting, that flowers
of this species and its variety thijmifolia keep their pinkish
colour for years, oi- ver}^ much longer than those of the majority
of Australian plants, and it seems evident that these flowers
contain some dye of a fixed nature which is probably worth
investigating.
Five species of Pomaderris were seen, distributed over various
portions of the Territory. Unless some disinfectant is used in
herbaria, the flowers of most plants are attacked by insects,
orchids, for instance, being devoured in a very short time; but
plants of the genus Pomaderris seem to possess some resisting
qualities, as its flowers may remain intact for several years with-
out any special care.
Brachy chiton ])0)>ul7ieiis (Kurrajong) is not common within
the Capital Territory, the locality being rather cold for it. There
is one fairly large tree, however, on the summit of a quartzite
hill within the Capital Cit^ site, and from which the hill has
taken its name. This species is a lover of limestone-formation,
and in places on the Western Slopes may occupy almost exclu-
sively certain areas where there is a considerable outcrop of
limestone. It is remarkable, therefore, that it should flourish
on this quartzite-hill, the rocks of which contain only a trace of
lime, according to Mr. J. C. H. Mingaye, but have a high per-
centage of silica, a constituent which this plant does not favour
when present in large quantities. A few Kurrajcmgs were
noticed at other spots, notably between the Naas Iliver and
Gudgenby, in which locality they were chiefly on the north or
warm sides of the hills, and by this means were able to grow at
higlier altitudes than usual in this latitude, one tree being seen
at an elevation of about 2,600 feet above sea-level.
Viola betonicce/olia and V. hederacea, commonly known as
Wild Violets, were seen in many shady nooks throughout the
Territory, and the beautiful dark blue flowers of the former were
noticed as far up as between the 5,500 and 6,000 feet levels
on Bimberi Peak.
698 NOTES ON THE NATIVE FLORA OF N. S. WALES, X.,
Of the Eucalypts seen within the Federal Territory, none has
so great a vertical range as E. coHacea, which extends from the
City site at about 2,000 feet, to the summit of Bimberi at 6,264
feet. On the lower land, it is often known as Scribbly Gum
from the insect-markings which appear like scribbles on the
bark, a feature not confined to this species, but on the high
mountains, where it is dwarfed and grows as a spreading plant
with several stems, it is usually known as Snow-Gum.
The largest trees within the Federal Territory are E. ijigantea
and E fastigata, and both are common on Brindabella Mountain
near Coree, the former being known as White, and the latter as
Black Mountain Ash.
E. giyautea was described by Hooker,* but there seems no
doubt that, when doing so, he had in his mind the present tree,
and also a previously described Stringybark, E. ohliqua L'Herit.,
both occurring in Tasmania, the latter being the more common
of the two. When the identity of E. ohliqua was afterwards
placed beyond doubt, E. gigantea was accepted as a synonym.
In 1900, Mr. R. T. Baker, F.L.S , described this tree under the
name of E. Delegatensis, from Southern New South Wales,! and
pointed out its specific characters. In 1913, Mr. Maiden repro-
duced Hooker's figure of E. gigantea as given in the Flora of
Tasmania (Vol. i., p. 136), and showed how the confusion between
E. obliqua and E. gigantea had arisen. J In Hooker's figure, the
fruils depicted appear to be those of the Mountain Ash, and not
those of E. ohliqua.
The bark of this Ash for about half-way up the trunk is
fibrous, while the upper portion and the branches are smooth
and white, hence the prefix, white, before the name of Ash or
Mountain Ash. The timber of this species, though very valu-
able, is known to be light and fissile, and, on examining seedlings
of 4-5 feet high, on Brindabella Mountain, it was noticed that
the wood was exceptionally soft and would snap with only a
* Lond. Journ. Bot., vi., 479 (1847).
t These Proceedings, 1900, Vol. xxv., p.305.
X " Forest Flora of New South Wales," Part li.
BY R. H. CAMBAGE. 699
gentle pressure. So far as I know, the most northern tree of this
species is about three miles north of Mount Coree.*
Eucalyptus fastiyata is very plentiful along the mountain sides
under C'oree and Tidbinbilla, and as its trunk and large branches
are covered with fibrous, brown bark, it is, in contradistinction
to the White Ash, called Black Mountain Ash. Its great affinity
with the giant gumtree of Victoria, E. regnans F.v.M., is well
known, and since E. fasiigatawa^s described by Deane & Maiden,
the latter has expressed the view that it is only a form of the
former.! At the same time, it appears as a distinct tree when
seen in the forest, for while the Victorian and Tasmanian repre-
sentatives of E. regnans are tall gumtrees with fibrous bark for
only 10 or 20 feet at the base, E. fastiyata. wherever it has been
seen over its wide range in this State, has fibrous bark on its
trunk and large branches.
E. dives and E. maculosa are often found in association, and
both will thrive in soils heavily charged with iron.
E. macrorrhyyicha (Red Stringybark) is not uncommon, and
with E. Juemastoma (Brittle Gum), and a few trees of Exocarpus
cnpi-essi/ormis (Wild Cherry), takes possession of the higher por-
tions of the Black Mountain, all three being lovers of a siliceous
formation.
Eucalyptiis j^ohjanthemos {Ked Box) is fairly common through-
out the lower levels, and is the form (E. ovalifolia R. T. Baker)
with smooth gum-tree bark, except that, in many cases, the bark
is flaky for a few feet at the base, and as forest-trees are dis-
similar to the rough-barked ]led Box of Victoria and around
Albury. The great lasting qualities of Red Box posts are well
known throughout the Federal Territory.
E. elceophora {E. Cambagei, Mountain- Apple, No. 3000) occurs
at various points, and around Tharwa and Booroomba appeared
almost white in November, with its intensely glaucous fruits
and branchlets.
* For previous remarks ou this species under the name of E. ddeyateusis,
occurring near Tumbarumba, see These Proceedings, 1904, Vol. xxix.,
p,690.
t See "A Critical Revision of the Genus Eucalyptus," by J, H. Alaiden,
Part vii. (1905).
700 NOTES ON THK, N\TIVK FLOKA OF N. S. WALES, X.,
E. camphora (8wamp Gum, No. 3342) was seen only on Coree
Creek, near S. A. Shannon's, though it has a considerable range
on the highlands of New South Wales, southwards from the
Kylstone district. In January, 1913, it was found at various
points on the Omeo-Mount Hotham Road, in Victoria, between
the 3,000 and 4,000 feet levels (No.3682).
The plants identified as L6ptos2Jermum aUenuatutn were seen
up to 12 feet high, chiefl}^ on the banks of streams, and have
hard, firm, grey bark, and not scaly or tlaky baric such as is
found on plants recognised as of this species occurring around
Sydney and on the Blue Mountains.
Kuiizea Muelleri was observed in the driei- portions of swamjw
areas on the highlands around the Upper Cotter, growing in
small masses of about a foot high, and having whitish flowers.
These plants are associated with Aciphylla simplicifolia, Epacris
paludosa, Juncus falcatus, and perhaps Celmisia loiiyifolia (Plate
Ixxiv.).
Kunzea ^^edunciUdris was seen up to 15 feet high, with some-
what flaky bark, on the river-banks, and it occurs on some of the
mountain-summits as a tough, stunted plant of a few feet. Mr.
John Blundell, of Coree, informed me that the blacks formerly
split pieces of the wood of this highland form, which they called
Budawang, and, after hardening it by fire, used it as needles to
pierce holes in the skins of various animals so that such skins
might be sewed together for use as rugs.
Richea Guniiii was found only on the summit of Bimberi Peak,
and is an alpine plant with very beautiful clusters of flowers
somewhat resembling those of Dracophyllnin secundum, which
grows along the coastal districts.
Pomax umbellata is growing on the northern slopes of Black
Mountain, selecting a similar highly siliceous formation to that
which it favours around Sydney and on the Blue Mountains.
Wahlenberyia yracilis (Blue Bell) and Candollea serrulata
(Trigger-Flower) were seen at their best at elevations between
(4,000 and 5,000 feet. Their colours were deeper than usual, a
common feature with plants at high altitudes, and the two species
formed separate masses of most charming deep blue and red, the
BY R. H. CAMBAGE. 701
distant blending of wliicb gave a brightness to the open forest,
and formed a beautiful setting, in one of Nature's many artistic
designs.
Olearia argophyUa (Musk-Tree), which grows to a height of
over 20 feet, and is regarded as the largest Composite in the
world, is growing in the sheltered portions of Mount Coree,
associated with Dicksonia antarctica and Bedfordia salicitia, tiius
showing that the locality is not exposed to the western or drv
atmosphere, but is in a zone of moisture resulting from a good
rainfall on the mountain-side. In this, and similar sheltered
situations within the Federal Territory, the Lyre Bird, Menura
siiperba, has its home.
List of Plants.
The following is a list of plants seen within the Federal Capital
Territory : —
HEPATlCEiE : Marchanfia polyiHorpJia L., (a cosmopolitan
species).
GvATHEACE.E : Dicksouia antm'ctica Labill., (Tree-ferns on
Mount Coree).
POLYPODIACE^ : Dryopteris punctata (Thunb.) C. Chr., {Poly-
podium puuctatnm Thunb. j, Polyslichum acnleatum (L.) Schott,
[Aspidium aculeatum Swartz), Aspleuium flahellifolium Cav.,
Pleurosorua i uti/oliics (H.Br.), Jj/echuwm cartilagineuni iiw., B.
discolor (Forst.) Keys., {Lomaria discolor Willd), B. penua-
marina (Foir.) Kuhn, [Lonuwia alpiiia Spreng.j, B . capeiise (Li.)
Schlecht., {L. capensis Willd.), Cheilauthes tenuifolia Sw., (plants
up to 1 foot 9 inches on Black Mountain), Adiantum cethiopicum
L., (Maiden-Hair Fern), Pteridium aquilinum L. Kuhn., {Pteris
aquiliiia L., Bracken), Pulypodiiun diver sifolium Willd., (P.
scandeiis Labill).
PiNACEiE : Callitris calcarata R.Br., (Black or Mountain Pine).
TypHACEiE : Typha augitsti/olia L., (Bullrush, in Canbury
Creek and other streams).
PoTAMOGEToNACEiE : Fotamoyeton tricarinatus F.v.M. & A.
Benn., [P. natans Benth., non L ), P. perfoliatus L.
JuNCAGiNACEiE ; Tviglochin proceva R.Br.
702 NOTES ON THE NATIVK FLORA OF N. S. WALES, X.,
ALlSMATACEiE : AHsma plantago L., (along the banks of the
Molonglo River, and flowering in December).
GRAMiNEiE : Andropogon ajjiiiis R.Br., A. refractus R.Br.,
ThfAneda Forshalii Hack., {Anihisiiria ciliata Benth., Kangaroo
Grass), Panicum effusu^n R.Br., Stipa scahra Lindl., (Silver
Grass), Uchino^jogon ovatus Beauv., (Saw Grass), Calainagrostis
cemula Steud., {Deyeuxia Forsteri Kunth), Holcus lanatus L.,
(naturalised), Aira caryoy)hyllea L., (Fairy Grass), Danthonia
carphoides F.v.M., D. penicillata F.v.M., var. semiannul avis
F.V.M., Cynodon dactylon Rich., (Couch Grass, common in hot
and some temperate countries), Chloris truncata R.Br,, (Umbrella
Grass), Pappophomm commune F.v.M., Phragmites commimis
Trin., (^Arundo Phragmites L.), Koeleria phleoides Pers., (natural-
ised), Poa ccespitosa G. Forst., (White or Snow Tussocks), Fes-
tuca Hookeriana F.v.M., [Schedonorns Hookeriana Benth.), F.
hromoides L., (naturalised), Bromus maxim,us Desf., (naturalised),
Hoi'deum mnrinum L., (Barley Grass; naturalised).
Cyperace^ : Kyllingia hiterinedia R.Br., (A", brevi/olia
Rottb.), Cyper2is sanguineo-fuscus Nees, C . Guiinii Hook., (C.
lucidus R.Br.), Eleocharis acuta R.Br., E. cylindrostachys Boeck.,
Scirpus cernuus Vahl, {A. riparius Benth.), >S'. inundatus Poir.,
S. stellatus C. B. Clarke, (S. cartilagi7ieus Benth.), jS". lacustris L.,
S. polystachyus F.v.M., Carex te^-eticaidis F.v.M., C. appressa
R.Br., C. Gaudichaudiana Kunth, C. p)seudo-cyperus L.
RfiSTiONACEiE : Restio ausfralis R.Br., (at Gudgenby), Hypo-
Icena lateriflora Benth.
JuNCACEiE : Luzula campestris DC, Juncus hufonius L., J.
plebeius R.Br., (J. homalocaulis F.v.M.), J. paUidus R.Br., J.
radula Buch., J. vaginatus R.Bi-., J. polyanthemos Buch., J. pi'is-
m,atocarpus R.Br., ./. Fockei Buch., J. lamjyrocarpus Ehr., J.
falcatus E. Mey.
LiLiACEiE: Aiiguillaria dioica R.Br., ( Wurinhea dioica F.v.M.),
Bulhine hulhosa Haw., Thysanotus tuherosus R.Br., (Fringed
Violet), Arthrojjodiura panicnlatum R.Br., Tricoryiie elatior
H.Hr., Stypandra glauca R. IJr., Dianella tasvianica Rook., D.
revoluta R.Br., Xerotes longifolia R.Br., X. multifiora R.Br.,
X. filiformis R.Br., X. glauca R.Br., Xanthorrhcea sp. (Grass-
Tree, at Lower Cotter and Booroomba).
BY R. H. CAMBAGE. 703
AMARYLLiDACEiE : Hypoxis hygvometrica Labi)].
Orchid ACEiE : Gastrodia sesamoiaes R.Br., I'helymitra
venosa R.Br., Diuris maculata Sm., (Spotted Orchid), D. sul-
phured R.Br., Prasophyllum fuscum R.Br., Q.) Pterostylis cu7'ta
R.Br., {l)P. ohtusa R.Br., (near Gudgenby, at 4,700 feet), P. rufa
R.Br., (at Booroomba, a form with very short points to the
sepals and petals), Caladenia testacea R.Br., C. alba R.Br., (on
side of Tidbinbilla), C. dimor'pha Fitzg., (on summit of Tidbin-
billa).
Casuarine^ : Casiiarina stricta Ait., (She-oak, C. quadri-
valvis), C. Luehmanni R. T. Baker, (Bull-Oak), C. Cunning-
hamiana Miq., (River-Oak).
Urticace^e : Urtica incisa Poir., (Nettle), Australina pusilla
Gaud.
PROTEACEiE : Persoouia chamcepeuce Lhotsky, (Prostrate Gee-
bung, at Gudgenby), Orites lancifolia F.v.M., (on Mount Bim-
beri), Grevillea lanigera A. Cunn., G. jwinperina R.Br., (at junc-
tion of Murrumbidgee and Cotter Rivers), G. ausiralis R.Br., (a
very narrow-leaved form), Hakea sericea Schrad., (//. acicularis
R.Br.), H. inicrocarj^a R.Br., Lotnatia longifolia R.Br., Banksia
marginata Cav., (Honeysuckle).
SANTALACE.E : Exocarpus cupressifo7"mis Labill., (Native
Cherry), E. stricta R.Br., Choretrum spicatum F.v.M., Ompha-
comeria acsrba A. DC.
LoRANTHACE^ : Lovauthus pendulus Sieb., (Mistletoe, with
Eucalyj^tus dives as host).
PoLYGONACEiE : Ruwiex Brownii Campd., E. acetosella L.,
(Sorrel; naturalised), Polygonum 2^'^'ostratum R.Br., P. minus
Huds., (Smart-Eye, in bed of Murrumbidgee and other streams).
CHENOPODiACEiE : Che7iopodium triangulare R.Br.
CARYOPHYLLACEiE : Sileue gallica L., (naturalised), Cerastium
vulgatum L., (Mouse-ear Chick-weed, naturalised), Stellaria
pungens Brong., Spergida7'ia rubra Camb., Tunica prolifera
Scop., (Dianthus, naturalised), Scleranthus bijlorus Hook.,
(Cushion- Plants forming green compact mats).
RANUNCULACEiE: Clematis aristata R.Br., C. microphylla DC,
Ranuncidus aquatilis L.(?), H. lappaceus Sm., (Buttercup), R.
704 NOTES ON THE NATIVE FLOIJA OF N. S. WALKS, X.,
AiHws Banks and Sol , (K. plebeiv.s R.Br.), R. rivularis Banks and
Sol., R. parvifloi'ns F..
Magnoliace^ : Drimys aromatica F.v.M., (Peppercorn).
MoNlMiACEiE : Iledycarya angiistifolia A. Cunn., {H. Cmniivg-
hamii Tul , Native Mulberry).
LaukacEjE : Cassytha pJuwlasia F.v.M., (Dodder), C. melaiifha
R.Br.
Crucifer.^ : Cardamine hirsuta L., var. tenuifolia F.v.M., (C.
teniiifo/ia Hook. ).
Duoserace^ : Drosera peltata Sni., (Sundew or Fly-catcher),
D. miriculata Backh
CkassulacE/E : Tillcea verticiUaris DC.
PiTTOSPORACEiE : Marianthus procumbe'HF! Benth., Biirsaria
spmosa Cav., (Whitethorn), Billardiera ^canden^ Sm., (Roly-Poly
Vine).
RoSACEiE : Rubus parvifoliiis L., (Redherry), Acrcna ovina
A. Cunn , A. sangidsorbcB Vahl, (Burr).
LEGUMiNOSiE : Sub-family Mimosoideae: — Acacia lanigera A.
Cunn., var. vemdosa, {A. venulosa Benth., on Black Mountain),
A. siciiliformis A. Cunn., (at Tidbinbilla and Gudgenby), A.
diffusa Edw., (Prickly Wattle), A. armata R.Br., (Kangaroo-
Thorn, on Mount Ainslie), A. vertdcijiua A. Cunn., A. peiini-
nervis Sieb., (Mountain-Hickory), A. obtusata Sieb., (on Black
Mountain and Coree), A. rubida A. Cunn., (Red-leaved Wattle),
A. buxifolia A. Cunn., A. pravissima Y.v.M., (Cotter River near
the dam, and on the divide between the Gudgenby and Cotter
waters, at 4,500 feet), A. Dawsoiii R. T. Baker, (near Gundaroo
Road from Queanbeyan), A. melanoxyloit R.Br., (Hickory or
Tasmanian Blackwood), A. implexa Benth., A. decurrens Willd.,
var. mollis, (Green Wattle, seen only in the northern or lower
portion of the Territory), A. dealbata Link, (Silver Wattle).
Sub-family Papilionatse : — Trifoliutn arvense L., (Hare's-foot Tre-
foil, naturalised), Oxylobium ellij^licum R.Br., var. alpinum, (on
Tidbinbilla at 4,500 feet, and on Bimberi at 5,300 feet), 0.
alpestre F.v.M., 0. Pulteneoi DC, 0. procumbens F.v.M. (?),
Mirbelia oxylobioides F.v.M., Gompholobium Htiegelii Benth.,
G. grandijlorura Sm.('?), G. unciiiatum A. Cunn., Daviesia corym-
BY R. H. CAMBAGE. 705
hosa Sm., D. nlicina Sm., PiiUeiupa Muelleri Bentb.C?), (on Brin-
dabella Mountain), P. j^rocumhens A. Cunn., (at Boorooniba, on
granite at 3,700 feet), P.fascicuJata Bentli., (at 4,500 feet, Giid-
genby to Cotter River, rare in New South Wales), Dillwynia
eriQifolia Sm., var. iihylicoides^ Platylohiiim formoHum Sm.,
(around Coree), Bossicea buxifolia A. Cunn., (at Booroomba), B.
Walkeri F.v.M., (5 feet high, with a cluster of from 30 to 40
stems; at junction of Murrumbidgee and Cotter), Hovea linearif<
R.Br., Lohis comiculatush., (Upper Cotter), L. aust7'alis A ndr.,
(Upper Cotter, flowers purple to pink). Indigo/era australis
Willd., (Indigo), Psoralea adscendens F.v.M., Swainsoita tephro-
tricha F.v.M., Zornia diphylla Pers., Desmodinm varians Endl.,
Glycine clandestiiia Wendl., G. Latroheaiia Benth., (with blue
flowers among the grass, on the City site), Hardenheryia mono-
phylla Benth., (False Sarsaparilla).
GEKANiACKiE : Gevaiiunn dissectum L., Urodimn cygnorum
Nees, E. cicnfarium Willd., (naturalised). Pelargonium australe
Willd.
OxALiDACEiE : Oxalis corniculata L., (Sour Grass).
RuTACEiE : Boronia polyyalifolia Sm., Eriostemon myoporoides
DC, Phebaliam squmnulosiim Vent., var, alpinum, fat 4,500
feet on Coree), Correa s])eciosa Andr.
Tkemandkace.*: : 7'etrathec<i ericifulia Sm.
PoLYGALACE.^^. : Gotnesperma velnsnrri Labill.
EuPHOKBiACEiE : Phyllauthus thymoides Sieb., Euphorbia
Drammondii Boiss., E. Lathyrus L., (naturalised), Poranthera
microphylla Brongn., Bertya ohiefolia Planch., (near Murrum-
bidgee and Cotter junction).
STACKHOUSlACEiE : Stackhousia linariifolia A. Cunn., [S.
monogyna Labill ), S. viminea Sm.
Sapindack.e : Dmhuiea viscosa L., D. attenuata A. Cunn.
Rhamnace^e : Pomaderris elliptica Labill., (on Black Mount-
ain), P. apetala Labill., P. prunifoli(t A. Cunn., P. raceinosa
Hook., P, pliylicifoHa Lodd., Cryptandra spinescens Sieb., Dis-
caria anstralh Hook., (low prickly bushes, with sickly-sweet
flowers j.
Malvaceae : Playianthus pulch'Uus A. Gray, also var.
55
706 NOTES ON THE NATIVE FLORA OF N. S. WALES, X.,
tomentosns Hook., Malva rotundifolia L., (naturalised), Modiola
multifida Mcench., (naturalised).
Sterculiace.^^ : Brachychiton populyieiis R.Br., (Sterculia
diversifolia G. Don, Kurrajong).
Dilleniace^ : Ilibhertia stricta R.Br., H. sevpyllifolia R.Br.,
H. linearis R.Br., var. ohtusi.folia.
Guttifer^ : Hypericum japonicnm Thunb., also var. arami-
neum F.v.M.
Violace^ : Viohi betonicce/oha 8m., (Native Violet), V.
hederacea Labill.
THYMELiEACE^E : Pimeleci glauca R.Br., a form with very
narrow and acute involucral bracts, P. colorans A. Cunn., P.
linifolia Sm., P. ligustrina Labill., also var. hypericina Benth.,
P.paiiciflora R.Br., P. curviflora R.Br., (No. 3473, three feet
high, near Gudgenby).
Lythrace.¥. : Lythrum salicaria ]j., (a swamp-plant), Z.
hyssopifolia L.
Myjitace^ : Eucidy])ius steUulata Sieb., (Sally), E coriacea
A. Cunn., (Snow-Gum), E. amycjdcdina Labill., (Peppermint or
Messmate), E. fastiyata Deane and Maiden, (Black Mountain
Ash; around Mounts Coree and Tidbinbilla), E, dives Schauer,
(Peppermint), E. gigantea Hook , (E. Delegatensis R. T. Baker,
White Mountain Ash), E. inacrorrliyncha F.v.M., (Red Stringy-
bark), E. hfemastoma Sm., (Brittle Gum), E melliodora A. Cunn.,
(Yellow Box), E. jwlyarUhemos Schauer, (Red Box), E. rubida
Deane and Maiden, (a White Gum, with orbicular reversion-
foliage), E. maculosa R. T. Baker, (a gum-tree, slightly spotted,
but distinct from the Spotted Gum of the coast), E. camphora
R. T. Baker, (on Condore Creek near Mount Coree), E. aggregata
Deane and Maiden, (a few trees seen on a flat on western side
of Bungendore Road, between the 48 and 49 mile posts from
Goulburn),^. eheophora F.v.M .,(£". Cambayei Deane and Maiden,
Mountain Apple), E. Bridgesiana R. T. Baker, (Apple-Tree or
WooUybutt; recognised by Mr. Maiden as E. Stuartiana F.v.M.),
E. inminalis Labill., (White or Manna Gum), E. Blakelyi Maiden,
(Forest Red Gum), E. dealbata A. Cunn., Leptospermum, Jiaves-
cens Sm., var. obovatura F.v.M., (near summit of Mount Coree),
BY R. *H. CAMBAGE. 707
Tj. scopariitm Forst., (Tea-tree, the narrow-leaved form), L. lani-
rjerum Sm., (along the banks of creeks; trees 30 feet high along
the damp gullies on the slopes of Bimberi, just above the 5,000
feet level), L. stellatn,m, Cav., L. attenuatnm Sm., L. Diyrtifolhtm
Sieb., L. triloculare Vent., (with silky-hairy calyx-tubes and
leaves, on Black Mountain), Kimzea Muelleri Bentli., (at 4,500
feet, oji divide between Gudgenby and Cotter Rivers), K. parvi-
folia Schauer, (between the 7 and 8 mile posts, Queanbeyan to
Tharwa Road), K. peduncularis F.v.M., (on banks of Murrum-
bidgee and other streams, also near the summits of Coree and
Tidbinbilla; in full flower early in December), Callistetnon palu-
dosus F.V.M., in bed of Molonglo, Murrumbidgee, Paddy's and
Cotter Rivers, flowering early in December, flowers creamy-
purple), C. lophanthus Sweet, (near the summit of Mount Tid-
binbilla), 6'. Sieberi DC, near the summit of Mount Bimberi),
Bceckea Guiiniana Schauer, Ccdycothrlx {Calythrix) tetragona
Labill.
OENOTHERACEyE : Epilobitim (jlahdhim G. Forst., Oenothera
hieihiiis L., (Primrose, naturalised).
Halorrhagace/E : Halorrliagis titragyna (Labill.) Hook., //.
)iucrantha (Thunb.) R.Br., Myriophyllum propinquu'm A. Cunn.
Araliace^: Tieghemopanax samhucif liusl^. V^iguier, [Panax
samhucifolius Sieb.), Astrotricha ledifolia DC, (on Black Mount-
ain and Coree).
Umbellifer.e : II ydrocotyle laxijiora DC, (//. Candollei
F.v.M. ), Didiscas hiimilis Hook., {Trachymene huniilis Benth,),
Trachyiiiene Billardieri F.v.M,, [Siebera Billardieri Benth, on
Tidbinbilla), Oreoinyrrhis atidicola Endl., (Native Carra way-seed),
Aciphylla simplicifolia F.v.M,, (in damp spots on the highlands
around Gudgenby and Bimberi), Daucus br'achiatiis Sieb.
EpACRiDACEiE : Styphel la triflora Audv., Melichrus iirceolaluH
R.Br., Lissanthe strigosa Sm., Leucopogon laiiceolatus Ji.Br., L.
virgatus R.Br., L. llookeri Sond., L. bi^orus R.Br., L. Fraseri
A. Cunn., Monotoca scoparia R.Br., Acrotriche agyregata R.Br.,
A. serr^datal^,.\^v., lirarJiyloma daphnoides 'Benth., E]jacris palu-
dosa R.Br., E. brevi/olia Stapf, (at 3,000 feet on Mount Coree),
K, serpyllifolid R.Br., E. microphylla R.Br., Richea Gu7t7iiiIiook.
708 NOTES ON THE NATIVE FLORA OF N. S. WALES, X.,
Pkimulace^: AnngnJlis arvfiisis L,, (Pimpernel, natiualised).
Ghntianace.e : Erythrcna australis R.Br., LiinitaiiAheiimni
creuatinn F.v.M., (an aquatic plant, with beautiful yellow, large,
fringed Huwers: in Molonglo Hiver).
Convolvulace.e : Couvolvidus eruhescens Sims.
BoKRAGiNACEiE : Ci/uoijlossntn australe R.Br.
Labiate : Mentha laxijiora Benth., J/, aastralis R.Br., (Penny
Royal), JJ.8aturejoides J\.hi\, Salvia verbenacea L., (naturalised),
BruneUa {Frunella) vulyaris DC, Scutellaria hmnilis R.Br.,
Prostanthera lasianthos LabilL, (Native Lilac, called Turpentine-
Bush at Coree), F. cuneata Benth., Wesfriugia ereniicola A.
Cunn., (white flowers; opposite junction of Murrumbidgee and
Colter), Ajnya aastralis R.Br.
►SoLANACE.E : Solauum uhjrum L., *S'. simile F.v.M., S. ojjacunt
A.Br., (naturalised). Datura stramoniuvi L., (naturalised).
ScROPHULARiACE.E : Mimulus moschatus Dougl., (naturalised),
Gratiola Feruviana L., (Brook-Lime), G. nana Benth., Veronica
perfoliata R.Br., (on the highlands, with beautiful blue flowers),
r. Derwentia Littlej., V. (jracilis R.Br., V. calycina R.Br.,
Eaphrasia Fyroionii F.v.M., (on Coree, and at 4,500 feet on Tid-
binbilla), Vcrhasemii hlattaria L., (naturalised).
Lentibulariace.e: Utricnlaria dichotoma LabilL, var. unijlora
Benth.
Plantaginace^ : Flantago varia R.Br.
RuBiACEiE : Coprosnia hirtella LabilL, (at 4,000 feet, on
granite, at Booroomba, also towards the summit of Tidbinbilla),
Foutax muOellata Sol., Asperula oliyantha F.v.M., also var.
scoparia F.v.M., Galiani niubrosani Sol.
Caprifoliace.e : Sambncas Gaudichaudiana DC.
CucuRBiTACE.E : Cacunds my?'iocai'pas Naud., (small melons,
naturalised).
Campanulace^ : Lobelia dentata Cav., L. pedunculata R.Br.,
Isotomajluviatilis F.v.M., Wahlenheryia yracilis DC.
Goodenlvce/e : Velleia paradoxa R.Br., Goodenia hederacea
Sm., G. pinnatijida Schlecht.
BY R. H. CAMBAGE. 709
Candolleace.t: (Stylidiaeete) : CandoUra sprrulafa Labill.,
[StylidiuAn tjraniiitijhflnnt Sw., Trigger-Flowei").
CoMPOSiTiE : Olearia mcAjalophylla F.v.M., (around Coree and
Tidbiubilla), 0. chrijsophylla^iinih., 0 . anjopht/lht Labill., (Musk-
Tree), 0. stellulaia DC, Cehn'isia loiKjifolia Cass., (Aster celmisia
F.v.M., Silver Daisy), VUtadinia australis A. Rich., Calutis
scabiosi/olia .Sond. and F.v.M., var. intec/ri/olia, Lagenophora
Billardierl Cass., Brachyctmie scapiyrra DC, B. scapn/onnis DC,
(on Coree and near Gudgenby at 4,500 feet), />'. Sieberi DC.(?),
B. discolor C 8tuart, Cotala Jilicida Hook,, CeiUipeda C anuing-
hamii F.v.M., Craspedia Richea Cass., (Bachelors' Buttons),
Casslnia amdcata R.Br., (at Boorooniba and Coree), C lonyifolia
R.Br., (at Gudgenby), C quinquefaria R.Br., Podolepls loiaji-
pedata A. Cunn., also var. robusta Maiden and Betche, P.
cauescens A. Cunn. Lepjtorrhynchos squamatus Less., HeHchrysuDi
scorpioides Labill., II. Incidum Henck., and vai-. viscosimi, (H.
bracteatum Willd., "Everlasting Flower"), H. apiculatuin DC,
//. semipcqjposuin DC, II. ledifolium Benth., //. rosinari'ulfolimn
Less., var. thyrsoldeum Benth., (at 5,000 feet on Bimberi), II.
Stirlingii F.v.M., (at 4,000 feet on Bimberi), //. baccharoides
F.v.M., Heliptevimi anthemoides DC, H. incannm DC, //. di-
morplwlepis Benth., Gnaphalium japonicmn Thunb., G. purpa-
reniu L., Staarfiana Aluelleri Sond., Erechtites luixta DC, B.
qiradridentata DC, Seiiecio pectiuatus DC, S. dryadens Sieb., [S.
<iii.stralis A. Rich.), /^ed/ordia salicina DC, (near Mount Coree,
and locally called Adam's Flannel from its tomentose, Hannel-
like leaves), Cyinbonotus Laivsonianus Gaud., Centaurea calci-
^/Yi/X6L., (Star-Thistle, naturalised), C. solsf it i(dis J^., (imtuvAliiHid),
Microseris Forsteri Hook., (a yam, with yellow flowers), Hypo-
chmris radiataJ^., (Dandelion, naturalised), Ficris hierat-ioides 1j.,
(naturalised), Cardnus pyciiocephalus L., (naturaliseti).
Comparison with Tasmania.
Although the plant-associations found on tlie high points appear
to disclose certain facts, it has to be remembered that an assem-
blage of plants is not only regulated by climate, but by geological
formations as well. In discussing those plants noticed on Tid
710 NOTES ON THE NATIVE FLORA OF N. S. AVALES, X ,
biiibilla at 5,100 feet and on Bimberi at 6,200 feet, some inter-
esting features are brought out. On Tidbinbilla, the Families
with most representatives were Myrtacete and Leguminosa?, with
four species each, while Compositjc had three species. Curiously
no species whatever uf Leguminosjc was noticed on the summit
of Bimberi, while Myrlacea^ had three, but Compositai had
eleven.
Before conclusive deductions could be drawn from a record
such as this, it would be desiral)le to have a more systematic
examination carried out than time permitted me to make, and
over a greater number of levels. It is not remarkable, however,
that this limited record shows that the Composita\ which are
considered to be the largest Family among flowering plants, should
be the most numerous at the higher levels, for they are known
to be able to resist the cold. On the other hand, masses oi a
few species of Composit«e may be seen on the hot western plains
of this State, in places forming the dominant vegetation.
In coiniection with distribution, it is of interest to make a
comparison with Tasmania. There the Family Composite pre-
dominates, and contains more species than Leguminoste and
Myrtaceie combined, the figures being approximately — Compositfe
110, Leguminosie 60, and Myrtaceie 40*
From the list of plants collected by me within the Federal
Capital Territory, the Families, according to numbers of species,
are arranged in the following se([uence — Leguminoste 13 species,
Compositte 42, and Myrtaceie ol. These figures maybe modified
by further discoveries, and the first two Families may even
change places.
The genera are represented in the following order : Eucalyptus
(Myrtaceiti) 19 species. Acacia (Leguminosa?) 15, Juncus (Jun-
cacete) 10, and Helichrysum (Compositie) 8.
Out of 27 species noticed on the summit of Tidbinbilla, at
5,100 feet, 15 of these, or 55%, occur in Tasmania; while on
Bimberi, at 6,200 feet, 22 out of 30 species, or 73%, are found
in Tasmania.
* "The Tasinanian Flora," by Leonard Rodway, C.M.G., (1903).
BY R. H. CAMBAGK. 711
As an evidence of considerable similarity in climate, it is
pointed out that, in the whole of tlie Federal Capital Temtoiy,
'^>C) 1 native species were noticed, of which 233, or 65%, are also
indigenous in Tasmania.
T wish to express my indebtedness to Mr. J. H. Maiden,
F.R.8., the late Mr. E. Betche, and Mr. E. Cheel for assistance
and corrolwi^ation in the identification of plants. To Mr. C. R.
Scrivener, I.S.O., Director of Commonwealth Lands and Sur-
veys, 1 am grateful foi- having afforded me the opportunity and
facilities for visiting the various points witliin the Fedei'al Ter-
ritory; while to Messrs. Charles H. McKeachnie, of Booroomba,
and Marmaduke W. Lee, of Gudgenby, my thanks are due foi-
hospitality and personal guidance to the summit of Tidbinbilla,
and the head of the Cotter River respectively; also to Mr. John
Blundell for his escort to the summit of Mount Coree. T have
also to thank Mr." A. J. Hare, Under Secretary for Lands, and
Mr. E. B. Harkness, Under Secretary, Chief Secretary's Depart-
ment, for information concerning the early records of settlement
in the locality.
EXPLANATION OF PLATES LXXI.-LXXIV.
Plate Ixxi.
Ca/^iiarina Lnthmanni R. T. Baker; growing under shelter of bank on
Molonglo River.
Plate Ixxii.
Grej'illea jitnipe:nn(( R.Br.; near Cotter and Murrumbidgee Rivers.
Plate Ixxiii.
Baiiksia marguKita Cav. ; in open forest granite-country at Booroomba.
Plate Ixxiv.
iSwanipy plain, with Kniizta Muelleri, Aciphylla mmp/lcifo/ia, Epacris
pa/udoi^a, J uncus falcafus, and Eucalyptus coriacea; between Gud-
genby and Cotter River, at about 4,500 feet level.
712
OEDINARY MONTHLY MEETING.
November 27th, 1918.
Professor H. G. Chapman, M.I)., B.8., President, in the Chair.
Before proceeding with the formal business of the Meeting,
the President reminded Members of the supremely important
events that had transpired in the interval since the last Meeting
^the signing of the Armistice presaging a victorious peace, and
the vindication of the strenuous efforts of the Allied Nations to
re-establish Right and Justice, as opposed to Might and Cunning,
as the basic principle of International relations.
The President moved the following resolution, which was
carried by acclamation, the audience rising: — "That the Members
present at this Meeting desire to atiirm their loyalty to the King
— to record their thankfulness for the cessation of hostilities, as
the prelude to a triumphant peace : their gratitude for, and ap-
preciation of, the services rendered to the cause of humanity by
the Allied Armies and Navies (including those who have so
worthily represented the Commonwealth), and by those, both
women and men, who have co-operated with them: and their
sympathy with the bereaved."
The National Anthem was sung, and cheers given for the King.
A letter from Lieutenant G. Goldfinch (written from Havre,
18th September, 1918) returning thanks to Members for greet-
ings and a friendly message of sympathy when he was in hospital
in England, in the early part of the year, was communicated to
the Meeting.
The list of the names of Members on the Honour Roll — only
three of whom had yet returned — was read by the Secretary, the
audience standing. Mr. F. Turner and Dr. C. Hall expressed
the hope that the Honour Roll would take a permanent form —
NOTES AND EXHIBITS. 713
a matter that lias not been overlooked, but, at present, the records
available are incomplete.
Mr. J. H. Campbell, Hon. Treasurer, said that, as tlie only
way open to the Society in its corporate capacity, to help the
cause of the Allies, it had subscribed to all the War Loans, the
total amount beins: £37,000.
The President reminded Candidates for four Linnean Macleay
Fellowships, 1919-20, that the 30th inst. was the last day on
which applications would be received.
The Donations and Exchanges received since the previous
Monthly Meeting (October 30th, 1918), amounting to 3 Vols.,
38 Parts or Nos., 7 Bulletins, 1 Rt^port, and i Pamphlets, received
from 33 Societies, etc., and one private donor, were laid upon the
table.
NOTES AMD EXHIBITS.
Mr. Fred Turner exhibited a teratological specimen of the
European Plantago lanceolata Linn., from Chatswood, in which
all the spikes or heads had developed a number of secondary
spikes, in a way that he had not previously observed in this
species in Australia.
Mr. E. Cheel exhibited a species of Callisf*'inon pachyphylhi.^
showing the delayed dehiscence of the fruits, corresponding, to a
certain extent, to those of C. rigidus described by Professor A.
J. Ewart (Annals of Botany, xxi., p. 135, 1907). The specimen
exhibited was taken from a plant cultivated at Ashfield, and is
7 feet long. It was originally raised from seed collected at
Bullahdelah, in August, 1911. The seed was sown on 29th
October, 1912, germinated on 6th November, 1912, and the plant
flowered for the first time in October, 1914, but no fruits were
set. In October, 1915, it flowered for the second time, and
several fruits set; these, in October, 1918, were apparently fully
developed, and measured 5 mm. in diameter. In October, 1916,
the plant flowered for the third time, and again in April of the
same year; the individual fruits of both of these clusters measure
50
714 NOTES AND EXHIBITS.
from 7 to 9 mm. in diameter. In October, 1917, a normal crop
of flowers was produced; and a few spikes appeared in December,
and again in April of the same year, so that there were three
flowering-periods. In October, 1918, the plant had a profusion
of spikes, and a branch (exhibited) was cut below the fruiting-
spike of 1915. It will be seen from this, that the fruits were
normally delayed for three years; and those of 1916 for two
years, as the valves were unopened; but, ten days after the
branch was cut, the seeds freely escaped, and some of them,
when sown, readily germinated. The fruits of the October, De-
cember, and April (1917) flowers were not sufticiently matured,
as no seeds have fallen from the fruits. It is interesting to note
that the character of the delayed dehiscence is similar in all
species of Callistemon, except C. viminalis, in which species the
fruits are fully developed about ten months after the flowering-
period; and the seeds germinate freely as soon as the twigs are
cut, and the seeds liberated. Prof. Ewart's observations were
made in connection with C. rigid^is, cultivated in Melbourne; but,
so far as the exhibitor knew, C. rigidus is confined to the Port
Jackson district, and is not a native of West Australia, as stated
by Prof. Ewart.
15
DESCRIPTIONS OF NEW SPECIES OF AUSTRALIAN
COLEOPTERA. Part xiv.
By Arthur M. Lea, F.E.S.
LUCANID^.
LiSSOTES C4RAMMICUS, 11. Sp.
(J. Black, shining; parts of appendages obscurely diluted with
red. Sides and undersurface very sparsely clothed, the legs
moderately setose.
Head gently and almost evenly convex, a fairly large notch on
each side, front incurved to middle and almost impunctate; with
large, round, dense punctures at sides and about eyes, becoming
smaller towards middle of base. Mandibles not very large and
almost simple. Eyes small, round, completely enclosed, and four
in number, the lower ones slightly smaller than the others.
Antennje with three apical joints moderately large, the apical
one truncated. Prothorax with sides and base margined, sides
lightly sinuous, dilated to near apex; with large, round, dense
punctures on sides, becoming smaller towards middle, the middle
itself almost impunctate and evenly convex. Elytra scarcely
longer than head (including mandibles) and prothorax combined;
each with four, rather wide, shining, impunctate lines from base
to beyond the middle, elsewhere with crowded and comparatively
small punctures, but a few larger ones at sides of the smooth
lines Front tihice with two, strong, obtuse, apical teeth, and
three smaller and still more obtuse ones. Length, 16 mm.
ffab.~New South Wales: Bodalla (Dr. R. H. Pulleine).
Unique.
No other Australian species of Lissotes, except L. Inteus {which.
by the figure looks like a Lissotes, and was doubtfully referred
by Westwood to Dorcus; it was omitted from Masters' Catalogue),
has been described as having four eyes; Westwood regarded the
716 NEW SPECIES OF AUSTRALIAN COLEOPTERA, xiv.,
type of that species as a male, despite the feminine appearance
of the mandibles: but it was described and figured as having six
impressed striae on each elytron; on the present species there are
no striae, their places being taken by smooth, shining lines not
interrupting the general convexity; there are also many other
differences in the head, antennae, etc. On account of its eyes,
the species should perhaps liave been referred to Lissapterus
{how lit anus of that genus has sometimes been referred to Lissofes),
but the head and mandibles are very different from those of the
species at present referred to that genus. The mandibles of the
type are touching at their tips, and enclose a top-shaped space;
each has an obtuse swelling about the middle of the upper
surface, and is obtusely bicuspidate at the apex.
SCARAB^ID^.
LiPARETRUS MA.TORINUS, n.Sp.
Black, shining; elytra and appendages reddish-castaneous.
Front of head, pronotum (including disc), and elytra at base and
near suture, with rather long, erect, and rather sparse whitish
hair; undersurface, hind-parts, and legs with dense clothing.
Head wntli crowded (but not confluent) and not very large
punctures, becoming larger and sparser on clypeus, sides of the
latter conspicuously narrowed and sinuous to apex, which is
evenly and moderately incurved to middle. Antennae nine-
jointed. Prothorax with strongly rounded sides, hind angles
rounded off, front ones produced and acute, median line vague
and traceable only towards base; punctures of somewhat uneven
size, but mostly rather large, and not very crowded. Elytra
with punctures moderately large, becoming smaller and more
crowded posteriorly, geminate striae feebly defined. Hind-parts
with rather dense punctures, smaller and denser on propygidium
than on pygidium. Front tibicB strongly and obtusely tridentate;
hind tarsi with basal joint distinctly shorter than second.
Length, 9 mm. •
Hah. — Queensland (unique).
The erect pilosity is continued on to the elytra from the pro-
notum, but is not very don.se there; regarding the species, how-
BY A. M. r.£A. 717
ever, as belonging to Group 1, of Blackburn's Tal)le,* it would be
associated with L. fuh-ohirtus, from wliicb it differs in being
larger, with the prothoracic punctures considerably larger, and
the surface more shining: regarding it as belonging to Group 2,
it would not fit into either F or F¥, as the clypeus is neither
truncate nor rounded in front. It is larger than any previously
described black species with red elytra from Queensland. Some
parts of the pronotum are very obscurely diluted with red; the
elytra of the type are without a membranous fringe.
LiPARETRUS ACUTANGULUS, n.sp.
(J. Black: elytra (base narrowly black), hind-parts and append-
ages (most of femora excepted) bright reddish-castaneous.
Undersurface and legs with long, pale hair, a fringe of similar
hair on each side of prothorax, but becoming darker in front,
rest of upper surface glabrous.
Head with rather small and crowded, but not confluent punc-
tures, becoming larger and less crowded on clypeus; sides of
clypeus lightly elevated and strongly narrowed, apex strongly
elevated, lightly emarginate and acutely produced at sides.
Antennae nine-jointed. Prothorax with sides strongly rounded,
hind angles widely rounded off, front ones acute and produced,
median line very feeble; with dense and sharply defined but
rather small punctures, less numerous along middle than else-
where. Elytra with fairly large punctures, becoming crowded
towards sides and smaller posteriorly; geminate-stria? well-de-
fined. I/ind-parts with punctures as on pronotum. Front tibice
strongly but obtusely tridentate; front tarsi rather thick, basal
joint keeled internally, claws thickened at base; hind tarsi with
first joint conspicuously shorter than second. Length, 7-7| mm.
^. Difters in having the clypeus short, truncate in front, with
the sides not produced, abdomen more convex, legs shorter, and
front tarsi thinner.
i^a6. — Queensland : Brisbane (T. McGregor).
Belongs to Blackburn's Group 4, and there would be referred
to JJ, but the median line of the pronotum, although faii'ly
* Trans. Roy. Soc. S. Aust. 1905, pp, 287-296.
718 NEW SPECIES OF AUSTRALIAN COLEOPTERA, xiv.,
distinct, is feeble even at the base; but regarding it as belonging
to K, it would be associated with L incerius^ from which it
differs in the acutangular clypeus of the male, larger punctures
of pronotum, and glabrous hind-parts; if referred to KK, it
would be readily distinguished from L. vicarius by the clypeus.
Its front tarsi and clypeus are much as in L. phodyiicopterus, of
Group 1 (with which it would be associated in Macleay's system),
but which has very different clothing. On the male, the club
is slightly infuscated; on the female, it is no darker than the rest
of the antennae; on the female also, the whole of the abdomen
and legs are reddish; on the male, manv of the elytral punctures
are transversely confluent towards the sides, but, on the female,
this appearance is less evident. On both specimens, there are a
few hairs at the tip of the pygidium, but, except for these, the
hind parts are quite glabrous; both specimens are without a
membrane at the apex of the elytra.
LiPARETRUS MIXTUS, n.Sp.
(J. Black; elytra and appendages (parts of the legs deeply in-
fuscated) bright castaneous. Undersurface with long, pale hairs,
a fringe of similar (but darker) hair on each side of pronotum;
hind-parts with short, dense, erect set;e in addition to long hair.
Head with rather small, crowded, and more or less transversely-
confluent punctures, becoming sparser, non-confluent, and slightly
larger on clypeus; clypeus with sides moderately elevated,
strongly decreasing and incurved to apex, apex rather strongly
but obtusely tridentate. Antenna? nine-jointed. Prothorax
with sides rounded and rather strongly produced towards base,
hind angles widely rounded off, front ones slightly produced
and somewhat acute, median line feeble; punctures small and
rather sparse, but more numerous towards sides and front angles.
Elytra with punctures of moderate size and not very numerous;
geminate-strite fairly well-defined. Uiiid-parts with dense sub-
asperate punctures of moderate size. Front tibicB strongly and
acutely tridentate; basal joint of hind tarsi slightly shorter than
second. Length, 6 mm.
Hah. — New South Wales (unique).
6Y A. Jt. LEA. 719
At first glance, the second joint of the hind tarsi appears to
be distinctly longer than the first, but in reality it is very little
longer. Regarding the species as belonging to Blackburn's
Group 4, it would be associated with L. dii<tans^ which is a much
larger species with all parts more or less reddish; if not to Group
4, it could only be referred to Group 7, and there associated with
L. iridipeiiiiis, from which it differs in its bright red elytra and
clothing of hind parts (the setae of these are very short and
partially concealed by the hairs, but they are very distinct from
the sides): each lateral fringe of the pronotum is partly pale and
partly dark, and although Blackburn tabulated Z. iridipeniiis
as having the fringe whitish, it is almost as often dark or partly
dark as whitish; at first glance, it appears to be close to L.
perkinsi of Group 10, but the clypeus has less acutely projecting
teeth (the type is certainly a male), and the hind parts are
densely clothed; L. hihiherculatus (also of Group 10) also differs
in the hind parts and in the considerably longer basal joint of
hind tarsi. Most of the upper surface is brightly iridescent; the
femora are almost black; the elytra (of the type) are without an
apical membrane.
LiPARETRUS INTERMEDIUS, n.Sp.
^. Black; elytra (except for a narrow space at base), most of
tarsi, parts of front tibiae, antenna) and tarsi, more or less
castaneous. With long hair on most parts, but elytra glabrous
Head with crowded and small punctures, but a few of larger
size scattered about; clypeus with sparser punctures than between
eyes, and smaller than the large ones there; sides moderately
elevated and decreasing (with curved outlines) to apex, which is
strongly but obtusely tridentate. Antenna? nine-jointed. Pro-
thorax with sides strongly rounded, hind angles widely rounded
off", front ones subacute and scarcely separately produced; with
comparatively large and not very dense punctures, mixed with
smaller ones; median line represented by a feeble basal depression
only. Elytra with not very dense punctures, no larger than the
larger ones on pronotum, and becoming smaller and more crowded
on sides ; geminate-striaj fairly well-defined. Hind-parts with
720 NEW SPECIES OF AUSTRALIAN COLEOPTERA, xiv.,
rather dense punctures, slightly larger on pygidiurn (where they
are as large as on pronotuni) than on propygidiuni. Front tibUn
strongly tridentate; basal joint of hind tarsi distinctly longer
than second. Length, 7 mm.
Ilab. — Queensland : Cunnamulla (H. Hardcastlei; unique.
Belongs to Group 8, of Blackburn's Table, but could hardly be
associated with L h-identaius, as tlie clypeal teeth are much less
acute and less produced: it differs from that species also in the
clothing of the pronotum being quite as dense at the base as in
front, and in the clothing of the pygidium; the punctures of the
pronotum are also sparser and considerably larger. The clothing
of the head and base of prothorax is much paler than on most
of the prothorax, where it is black or blackish (it is uniformly
coloured on Z. pa7'tn(i ens a.nd L. ohhisideiis)] irom L. parvideits h
differs also in having the clypeus more conspicuously tridentate
(both sexes of that species are before me), and from L. obtusidens
in having rather larger punctures (on the elytra as well as on
other parts). The antennae are paler than the other reddish
parts, but the outer parts of the club are lightly infuscated. The
elytra are terminated by an extremely short membrane.
LiPARETRUS QUINQUELOBATUS, n.Sp.
(J. Black; elytra, antennye and palpi bright flavo-castaneous,
legs of a darker red. Sterna with rather long pale hair, abdomen
sparsely clothed, clypeus with a few hairs, a pale fringe on each
side of pronotum. and extended on to sides of front margin, rest
of upper surface glabrous.
Head with crowded and small, non-contluent punctures, an
irregular row of larger ones near clypeal suture; clypeus with
sparser and larger punctures than between eyes, sides strongly
narrowed and suddenly incurved near apex, which is strongly
and acutely tridentate. Antenn<v? nine-jointed. Prothorax with
sides strongly rounded, hind angles widely rounded off, the front
ones acute, median line shallow: with dense and sharply defined
but rather small punctures. Elytra with rather sparse punc-
tures of moderate size, becoming smaller and denser at the sides
and apex; geminate-stria? well-defined, apical membrane narrow
BY A. M. LEA. 7^1
but conspicuous, lluid-parts with dense punctures, much as on
pronotuni, but a few of larger size scatteretl about. Front tibui^
strongly but obtusely tridentate: basal joint of bind tarsi con-
spicuously longer than second. Length, 8-8 1 nnn.
Ilab. — Queensland: Cloncurry(H. Hacker).
In Blackburn's Table, would be associated with L. perkinsi,
but ditfers from that species in being much larger, non-iridescent,
prothoracic punctures considerably larger and niore sharply
defined, and elytral punctures larger. L. trideiUahis, which has
a sornewhit similar clypeus and is similarly coloured, has the
front half of the pronotum densely clothed; from L. inifirmedius,
it differs in being larger, clypeus strongly incurved before the
apical teeth, prothorax glabrous on disc, and with smaller and
much denser punctures, elytra entirely pale, etc. The clypeus is
conspicuously tridentate in front, but the sides near the apex are
strongly incurved, so that, when viewed obliquely from behind,
it appears to consist of five almost equal lobes; immediately
behind its suture, the surface is shining and sparsely punctate,
with large punctures marking the position where the sparse and
dense punctures meet. On the two specimens before me, the
hind parts are entirely glabrous, except for a few hairs on the
pygidium.
Haplonycha coi.ossa, n.sp.
Bright reddish-castaneous, elytra paler. Undersurface and
legs with dense, golden hairs, a few similar hairs at sides of eves,
and forming a thin row in each lateral gutter of pronotum; mem-
branous fringes of elytra very short; pygidium glabrous.
Head with rather small but sharply defined punctures, becom-
ing somewhat larger and more numerous (but not confluent)
about clypeal suture ; front face of clypeus with numerous
setiferous punctures on sides, but aetse confined to a single row
across middle. Antenna? with fourth joint slightly longer than
third, the five following joints forming a club. Maxillary palpi
rather long, penultimate joint slightly longer than antepenulti-
mate, and scarcely shorter than apical, Prothorax more than
thrice as wide as long, sides rather strongly rounded and feeblv
arcuate to base and apex, front angles somewhat produced, hind
722 NEW specik:s of Australian coleoptera, xiv.,
ones obtuse and not completely rounded off; with rather small
but distinct punctures, sparser in middle than elsewhere. Elytra
moderately dilated to about the middle: punctures fairly numer
ous, except between the geminate strite (these very close together);
suture very feebly mucronate. Fygidium shining, lightly con-
vex; with rather small, scattered punctures. Basal joint of hind
ta7'si distinctly shorter than second. Length, 32 mm.
Hab. — Western Australia.
I have had the type for many years under the name of H.
gigmitea, but although looking like a large specimen of that
species, it may be at once distinguished by the maxillary palpi:
on ff. gigaiitea, the antepenultimate joint is slightly longer than
the penultimate, hence Blackburn referred it to his Group 5.
On the present species, the penultimate is slightly the longer,
hence the species must be referred to CC, of his Group 4, and
there associated with N. nobilis, from which it differs in having
the prothorax more transverse, and the fifth joint of the antennae
(instead of the sixth) the first of the club. The rami of the club
are decidedly long, but as that of its first joint is only about half
the length of the second, the type appears to be a male. From
some directions, the pronotum appears to have a fine, iridescent
bloom.
Nova PUS parvus, n.sp.
(J. Keddish-brown, head and parts of legs black. Under-
surface, legs, and upper part of pygidium with dense, rusty-red
hair, upper surface glabrous.
Head with crowded and irregular punctures: with a sharp,
short, oblique, simple horn. Antennae ten-, club three-jointed.
Prothorax about one-fourth wider than long, hind angles rounded
off, front ones produced and acute, with a large discal excavation,
its front part with transverse sculpture; with punctures of
moderate size, but very irregularly distributed. Elytra with
sutural and lateral striie well-defined, but the others represented
by feeble depressions or oblique scratches; punctures small and
sparse, but becoming immerous at apex and sides. Pygidiiiin
with numerous rather small punctures, becoming larger and
crowded at base. Length, 15 mm.
6y a. m. lea. 723
Hab. — Western Australia: Swan River (A. M. Lea); unique.
Much smaller than any previously described species, with the
prothoracic excavation unusually small and shallow (it scarcely
occupies one-third of the width of the prothorax), and the
cephalic horn very small and simple (scarcely half the length of
that of N. simplex). At first glance, the type bears a strong
superficial resemblance to the males of Isodon pecuarius, but the
horn is on the head, not on the prothorax.
Cryptodus aberrans, n.sp.
Black, not very highly polished (the elytra subopaque;, parts
of undersurface and of legs obscurely diluted with red, club of
antennae paler. Upper surface almost glabrous, undersurface
sparsely and irregularly clothed, pygidium with a few short set?e.
Head with rather large but not very dense punctures; clypeus
with front margin rather strongly elevated and truncate, lateral
inargins lightly elevated and oblique, suture well defined towards
sides but obsolete in middle, where the surface is slightly elevated;
iiientum large, with large, shallow punctures, depressed in front,
base rather lightly notched, with a few setfe and long bristles.
Antenna? apparently nine-jointed, club three-jointed. Prothorax
rather strongly convex, about once and one-half as wide as long,
sides strongly rounded, base bisinuate, front angles obtusely pro-
duced, the hind ones rounded ofl:*, median line rather vague; with
fairly large but nowhere crowded punctures. Elyti-a at base the
width of prothorax, slightly dilated to beyond the middle; surface
finely shagreened, with well-defined rows of fairly large punc-
tures towards suture, but becoming smaller and irregular towards
side and apex. Pygidium with fairly large but rather shallow
punctures. Four hind tihicH strongly serrate or digitate at apex,
each notch with a seta; front claws simple. Length, 11 mm.
Hah. — Northern Territory: Darwin (N. Davies); unique.
In its comparatively small size, convex body, and general
appearance, the present species certainly does not look a
CryjHodus;* but the wide mentum concealing all the mouth-parts
* Neither does C. yrossipes, at first glance, appear to belong to the genus,
but its remarkable mentum is almost exactly as in C. caviceps, which is
quite an ordinary species of Cryptoda-^.
724 NEW SPKCIES OF AUSTKALIAX CoLEOPTERA, xiv.,
is like tliat of Cryptodus, and utterly different from that of any
otiier Australian genus of Dynastides: the projection in front
of the prosternuni, and the triangularly dilated liasal. joint of the
antennae are also as in Cryptodas\ the four hind-tihiie are digitate
instead of truncate at apex, and are certainly not fringed with
setse (as on normal Dynastid'is), but, at the base uf each notch,
there is a seta; smaller and less distinct setie, however, are pre
sent on several other species of Cryptodus (e.g., C. tasmaaiamis)^
although they need to be closely looked for. The base of the
mentum is less deeply notched than is usual in those having it
notched, in this respect agreeing with 6'. gly(f'S: in C. tasmanianus,
it is truncate: and, in C. carioeps and C. grosslpes, it has a long
and almost vertical process ; the apex of the basal joint of
antennae is less conspicuously produced over the following joints
than is usual in the genus, but, from some directions, it entirely
conceals the two following joints. The elytra, at first glance,
appear to be glabrous, but, on close examination, some very fine
setie become visiVjle; there are three or four interstices on each
elytron slightly more prominent than the others, but not one is
conspicuously elevated. The sex of the type is doubtful, as many
males of Cryjytodus have the front claws simple.
CORYNOPHYLLUS INTEROCULARIS, n.Sp.
^. Black: antennae, most of legs and of undersurface more or
less castaneous-brown. Undersurface and legs with rather
dense, rusty-red hair; upper surface and pygidium glabrous.
Head concave, and with irregular (but not very dense) trans-
versely-contluent punctures between eyes ; clypeus moderately
long, punctures more or less confluent, apex and sides rather
strongly elevated, basal carina strongly elevated (sub-tuberculate)
in middle; mentum gently convex. Antennae ten-, cluV) three-
jointed, rami large, about as long as head is wide. Prothorax
not twice as wide as long, sides strongly rounded, apex bisinuate,
front angles produced, hind ones rounded off', a rather small
excavation in front, the middle of its front margin with a small
tubercle, basal gutter distinct at sides, but not traceable across
middle; with small and sparse punctures, becoming more numer-
BY A. M. LKA. 725
ous on sides and larger in excavation. Elytra the width of pro-
thorax; with irregular rows of large punctures in distinct striae,
hut becoming very irregular about apex. Pygidinm with vario-
lose punctures, almost absent along middle, but crowded in upper
corners. Spurs of hind tibice stout, and very unequal. Lengtii,
15|-17 mm.
Hab. — ^ew South Wales (Dr. E. AV. Ferguson), Jenolan (J.
C. Wiburd).
With the general outlines of C. modestus, but elytra dark and
club of antennse considerably smaller than in the male (although
much larger than in the female); from the dark form of C. fvrf-
numi, it is at once distinguished by the very different clypeus
and single cephalic elevation; C. andersoni has the clypeus dif-
ferent, and the club much larger. One of the specimens before
me has the upper surface entirely deep black, but, on the other,
it is obscurely diluted with red; on the latter specimen, the
clypeus is distinctly bilobed in front, but, on the other, it is
almost simple there. Some of the elytral stria^ are irregularly
EUCNEMID^.
MiCRORHAGUS RUFICOLLIS, n.sp.
Black ; prothorax and legs red, tarsi paler, antennse dull
piceous-brown, the two basal joints somewhat brightei-. Some-
what irregularly clothed with depressed, more or less stramineous
pubescence.
Head with dense, partially concealed punctures; with a fine
transverse carina near the base, and a very feeble longitudinal
one near each eye; these large and prominent. Antennae long
and thin, second joint very short, third slightly shorter than
first, and slightly longer than fourth, fourth to sixth somewhat
wider than the others, fourth to tenth slightly produced on one
side at apex, eleventh very thin, and conspicuously longer than
tenth. Prothorax at base almost twice as wide as the median
length, front angles rounded, hind ones strongly produced and
acutely carinated: front margin carinated, the carina on each
side with a short spur extended towards but not meeting the one
726 NEW SPKCIKS OF AUSTRALIAN COLEOPTERA, xiv.,
on the basal angle ; with a vaguely impressed median line,
becoming carinated at base; punctures moderately dense. Elytra
parallel-sided from near shoulders almost to apex; with punctures
of moderate size about base, becoming smaller posteriorly, but
about tip decidedly coarse. Length, 3 J mm.
ZTaS.— N.S.W. : National Park (A. M. Lea); unique.
The second joint of antenna^ has a curious appearance as of
being forced out of alignment. The elytral punctures are more
or less lineate in arrangement, but not placed in stride, the
sutural stria (and that only from about the middle) is the only
distinct one on each elytron; elsewhere there are but vague
remnants of striation, or none at all. The prosternal sulci are
deep, parallel-sided to near the base, and somewhat narrower than
the propleural parallelograms, which are about once and one-half
as long as their basal width.
Hemiopsida longicornis, n.sp.
Dull castaneous-brown; head, basal joint of antennae, junction
of prothorax and elytra, sterna, and hind coxse, black or blackish.
Rather densely clothed with moderately long, stramineous
pubescence.
Head densely granulate punctate; with a subfoveate impres-
sion between antennary sockets; clypeus widely depressed in
middle. Antennae long, slightly passing elytra, second joint
very short, third slightly longer than first and distinctly longer
than fourth, fourth slightly shorter than fifth, fifth-tenth sub-
equal in length, eleventh almost as long as ninth and tenth com-
bined. Prothorax strongly convex, front angles rounded, hind
ones acute and obliquely produced on to shoulders, with a very
feeble median line; with dense, rugose punctures. Elytra slightly
wider than hind angles of prothorax, parallel-sided to beyond
the middle; with rather dense and irregular, but sharply defined
punctures, becoming crowded about base; striation well-defined
throughout, but especially on apical fifth. Abdomen with a deep,
conspicuous, hairy depression on each side of middle, extending
from tip of first segment to tip of fourth. Length, 6-6^ mm.
Hah. — Queensland: Mount Tambourine (H. Hacker's No.895).
BY A. M. F.EA. 727
Readily distinguished from all other species, except H.
ventralis, by the abdominal depressions; the antennae are also of
unusual length; H. ventralis has the abdominal depressions com-
mencing nearer the base of the first segment, and not continued
beyond the third, and its antennae are shorter and more con-
spicuously (although not strongly) serrated.
Dyscolocerus porosus, n.sp.
(J. Black; undersurface, antennae, and legs obscurely reddish.
With very short pubescence.
Head with crowded punctures, with a very feeble depression
on clypeus. Antennae rather stout, eight basal joints densely
punctate, second joint slightly longer than fourth, third slightly
longer than two following combined, fourth to eighth equal and
strongly transverse, ninth about as long as three following joints
combined, and conspicuously wider, slightly longer and wider
than tenth and much shorter and distinctly wider than eleventh,
three apical joints as long as the seven preceding combined.
Prothorax with sides rather strongly rounded in front, hind
angles acute, with the outer side of each somewhat oblique,
median line rather shallow but well-defined; with crowded punc-
tures of moderate size. Elytra parallel-sided to beyond the
middle; punctures at base as on prothorax, becoming somewhat
smaller, but almost as crowded posteriorly; striation well-defined
throughout, becoming deep posteriorly. Length {$<^), 7-1 1mm.
9. Diifers in being slightly more robust, antennae shorter, the
three terminal joints distinctly shorter than the seven preceding
combined, the ninth scarcely shorter than the eleventh, and the
fourth-eighth not transverse.
Hab. N.S. W.: Forest Reefs.— W. A.: Swan River (A. M. Lea).
The antennae and sterna are somewhat darker than the abdo-
men and legs, but no parts are conspicuously reddish; the front
of the prothorax of several specimens is very obscurely diluted
with red. The pubescence of the upper surface is black and
very short, but on the head, and base and apex of prothorax, it
becomes longer and greyish, on the undersurface it is uniformly
pale. On one specimen, the head appears to have a feeble median
728 NEW SPEC'IKS OF AUSTRAIJAN fOLEOPTERA, xiv.,
line, but it is quite absent from the five others before me. The
Swan River specimen has the sterna and femora quite black. In
some respects, the female is close to the description of Lycaon
ater, but the fourth joint of the antennse is no shorter than the
fifth, and the prothorax has a conspicuous median line. The
types were taken in cop.
Dyscolocerus rubriventris, n.sp.
(J. Black ; abdomen and legs bright red, antennae reddish,
becoming darker towards base, with the basal joint black.
Clothed with blackish and ashen pubescence, becoming palei-and
more uniform on the undersurface.
Antenufe with second joint slightly longer than fourth, third
slightly longer than fourth and fifth coml)ined, fourth-eighth sub-
equal in length, ninth-eleventh as long as first seven combined,
ninth slightly wider than the following ones, almost as long as
the four preceding combined, about one-third longer than tenth,
and about two-thirds the length of eleventh. Length, 5J-6mm.
9. Differs in having the fourth-eighth joints of antennie slightly
wider than long, the ninth-eleventh scarcely as long as the seven
preceding combined, and the eleventh very little longer than the
ninth.
//rtfc.—N.S.W.: Jenolan (J. C. Wiburd).
The description of the sculpture of the preceding species,
except of the antenna?, applies exactly to the present species, but
besides the conspicuously red abdomen (in striking contrast to
the black sterna) it differs from that species in having the three
terminal joints of antennae longer (in both sexes) and the fourth-
eighth joints of the antennte of the male much less conspicuously
transverse.
DiCTYEUCNEMIS, n.g.
Head moderately large, antennary sockets comparatively small
and widely separated. Mandibles large, prominent, strongly
curved, their hind outline straight. Antennae thin and rather
long. Prothorax moderately transverse, hind angles small, pro-
duced slightly outwards but not backwards, and not embracing
the elytra. Scutellum subquadrate. Elytra strongly convex,
BY A. M. LEA. 729
distinctly wider than elj^tra, parallel-sided to near apex, epi-
pleural fold narrow but traceable almost to apex. Prosternum
with propleural triangles each with a narrow carina internally,
and a still finer one externally, the two touching the apex at a
slight distance (about equal to the length of the second joint of
antennte) from each other. Metasternum with episterna narrow
and parallel-sided. Abdomen with first .segment at the side
about as long as the fifth along middle. Legs rather long: inner
half of hind coxa? moderately long (about half the length of
.second segment of abdomen), then strongly narrowed to sides;
tarsi moderately long, second, third, and fourth joints of exactly
the same shape but decreasing in size; claws each with an obtuse
swelling at base.
The prothorax is very aberrant for the family, but the com-
bination of entire absence of a visible labrum, mandibles closely
applied to the breast and concealing the palpi within the buccal
cavity, intercoxal process of prosternum narrow and received
into a deep groove in the mesosternum, and abdomen with five
segments, forbid its being placed in any other family. The face
is vaguely suggestive of some females of the Rhijndoceridcc. In
Blackburn's Table, the genus would be associated with Lycaon
( = Hemiopsida), with some features of which it agrees, but the
base of the prothorax is at once distinctive from that, as from
all other Australian genera. There is nothing at all approaching
it in the Plates accompanying Bonvouloir's monograph. The
punctures of the head and prothorax are remarkable. The ex-
ternal face of the mandibles ^s densely punctate, the punctures
(except towards the base) within a depression enclosed by shining
carinse, which meet near the tips.
DiCTYEUCNEMIS MIRUS, n.sp.
Blackish-brown, or castaneous-brown, appendages somewhat
paler. With not very dense, and very short, whitish pubescence.
Head with large, shallow, net-like punctures, margined by fine
carinse, and with the inner part of each puncture flat and
shagreened; antennary sockets almost as far apart as the length
of the three basal joints of antenna?. Antennae extending almost
56
730 XEW SPKCIKS OF AUSTKALIAN COLEOPTERA, xiv.,
to hind coxse, first joint not carinated, and slightly longer than
third, second about half the length of fourth, third about as
long as fourth and fifth combined, fifth to tenth very feebly
decreasing in length, eleventh slightly longer than tenth, and
about the length of ninth. Prothorax truncate in front, front
angles rather strongly rounded, sides thence feebly undulated to
base, a transverse impression near base, marking off a wide and
short median lobe, with vague remnants of a median line, aud a
vague foveate impression on each side of middle; punctures as
on head. Scutellwin with apical half polished and almost im-
punctate. Elytra about one-fourth wider than prothorax, and
about five or six times as long; with numerous distinct punctures
and small round granules, becoming more crowded about base;
striation well defined. Frosternum with moderately large punc-
tures, each with a central pit, on middle portion, the propleural
triangles with punctures much as on upper surface, but less
defined. Apical segment of abdomen with dense, asperate punc-
tures, each side with a shallow depression. Legs densely asperate-
punctate; first and fifth joints of tarsi of equal length, and each
about as long as third and fourth combined. Length, 7-lOJmm.
Hab.—W.A.: Mullewa (IMiss J. F. May).
The two specimens taken by Miss May are evidently con-
specific, but the larger one is much darker than the other, with
some parts almost or quite black.
Nematodinus, n.g.
Head short, mandibles strongly sinuous posteriorly, antennary
sockets short and widely separated. Antennse short, first and
third joints long. Prothorax subquadrate, entirely concealing
head from above. Scutellum subquadrate. Elytra with an epi-
pleural fold at apex, and with suture armed. Prosternum with
a vague longitudinal impression towards each side, the impres-
sion bounded outwardly by a fine carina (representing the pro-
sternal suture), then with a narrow, almost parallel-sided space
between the carina and margin of pronotum (this represented by
a thin but not continuous carina). Metasternum elongate ;
episterna very narrow. Abdomen evenly convex, apex evenly
BY A. M. LEA. 731
rounded. Legs not very long; hind coxse moderately long near
where they touch, suddenly narrowed, and then almost parallel-
sided to sides; tarsi thin, fourth joint very small and simple;
claws thin, with a slight basal swelling.
The only species known has the top-heavy appearance of
Nematochfi and 7ricjo7io2)hu?'us, but it differs strikingly from
these genera in the prosternal sclerites, and in the tips of the
abdomen and of the elytra. Although the prosternal sutures
are not deeply impressed (they are, however, almost parallel with
each other) as they are in Microrhagus and Entomophthalmus,
the side-pieces may be regarded as propleural parallelograms, as
in those genera; whilst, in Nematodes and Trigonophuriis, the
side-pieces are decided triangles. Tn Blackburn's Table, the
genus would be associated with Hypoccelus, from which it is at
once distinguished by the elytra. The only genus at all ap-
proaching it in the tips of the elytra is Galba, with which it
has scarcely anything else in common. In catalogues, it may
be placed near Nematodes.
NeMATODINIIS ARMIPENNIS, 11. sp.
Of a dull castaneous brown, legs and antennae somewhat paler.
Moderately clothed with short, stramineous pubescence.
Head with crowded but not very large punctures. Antennae
not passing middle coxae, second joint very short, third about as
long as the three following combined, fourth slightly longer than
wide, fourth-tenth subequal in length but feebly dilated till the
tenth is feebly transverse, eleventh scarcely wider but distinctly
longer than tenth. Prothorax about as long as wide, front
straight across middle, with a fine marginal carina curved at
each side, and then continued parallel with the prosternal suture
till it obliquely diverges to margin the hind angle, with a feeble
medio-basal carina, each side of base depressed ; densely granulate-
punctate. Elytra with outlines continuously parallel with those
of prothorax to near the apex, with crowded granulate-punctures
about base, somewhat sparser elsewhere; tips with an epipleural
fold from about level with base of fifth segment of abdomen,
densely granulate and each with an oblique projection at the
suture; striation feeble.
732 NEW SPECIES OF AUSTRALIAN COLEOPTERA, xiv.,
^«6.— Queensland: Cape York (H. Elgner).— Northern Terri-
tory: Darwin (N. Davies).
Seen directly from behind, the tips of the elytra appear to
enclose the tip of the abdomen, and each to have a short semi-
upright process at the suture. On one specimen there is a con-
spicuous carina on the prosternum extending from the left eye
to the middle of the intercoxal process, but it is accidental, as it
is not represented on the right side, and is absent from two other
specimens.
Arisocephalus, n.g.
Head wide, antennary sockets large and moderately close
together, clypeus sinuous in front, its edge finely carinated.
Antennse v^ariable. Prothorax moderately transverse, basal
angles not very long but acutely carinated. Elytra subparallel-
sided, with a wide epipleural enlargement from base to hind
coxae. Prosternum with a conspicuous carina marking the suture
on each side from front coxa to level with middle of eye, pro-
pleural triangles bounded externally as well as internally by a
conspicuous carina. Metasternum with episterna rather narrow
and parallel-sided for some distance, but dilated posteriorly. Ab-
domen with fifth segment about as long as the two preceding com-
bined. Hind coxce (except for an incurvature at trochanters)
almost parallel-sided from inner to outer margins; tarsi com-
pressed, fourth joint small and feebly produced on lower surface,
claws each with an obtuse basal swelling.
In Blackburn's Table, this genus would be associated with
Microrhagus and Entomophthalmus, but the propleural triangles
(instead of parallelograms) and prosternal sutures not sulcate,
make it certain that the genus is not even close to these. The
tricarinated clypeus of two of the species is suggestive of affinity
with Ainsus, but that genus has prosternal lateral sulci, and
metasternal sulci; the general outlines, however, and especially
the head, are much the same. The propleural triangles are
without the least traces of longitudinal sulci, but are gently
concave, or flat throughout; the carina marking the prosternal
suture touches the front margin inwards of the point where the
marginal carina touches it, instead of meeting it there as on most
BT A. M. LEA. 733
genera of the family; in consequence, the triangles are not acutely
pointed. The hind coxae are very distinctive, being slightly
wider at their outer than their inner margins; the tarsi, when
viewed from the sides, appear to be moderately wide, but very
thin from above or below. In A. basalts, the fifth segment of
the abdomen is somewhat shorter than in the other species, but
its tip is somewhat produced (although much less conspicuously
so than in Neinatodes and IVigotiopleur^ts). The size and general
appearance of all the species are suggestive of Cardiophorus of
the Elateridcc. Type-species, A.Jiavipes.
Second joint of antenni^ (viewed from above) distinctly shorter
than tliird flavipes.
Second joint distinctly longer than third.
Elytra entirely reddish rufipemiis.
Elytra reddish only about base hasalis.
Arisocephalus flavipes, n.sp.
Black; extreme apex and base of prothorax, apex of scutellum,
and antennae (basal joint darker) reddish, elytra (base, suture,
and sides excepted) and undersurface (parts of abdomen paler)
of a dingy reddish-brown, legs (hind cox£e excepted) flavous.
Densely clothed with short, more or less upright pubescence,
sooty on the head and prothorax, mostly paler elsewhere.
Head with crowded and rather small punctures; with a con-
spicuous median carina from near base almost to apex; antennary
sockets bounded by a curved carina, the same joined in front to
the side of the clypeal margining carina. Antennae extending
to about hind coxse, second joint of antennae (as viewed from
above) somewhat shorter than third, third distinctly shorter
than fourth, its apex somewhat produced to one side, fourth-
eighth strongly serrate (almost pectinate), ninth-tenth less
strongly so, eleventh distinctly longer than tenth. Prothorax
not much wider than long, sides rather strongly rounded in
fi-ont, hind angles feebly directed outwards, the carina on each
acute and about once and one-half the length of the scutellum;
with a rather feeble median line, altering at the base to a feeble
carina; punctures crowded and small, becoming smaller and still
more crowded on sides. Scutellum with moderately dense punc-
734 NEW SPEClKS OF AUSTRALIAN COLEOPTERA, xiv ,
tures, and a feeble median carina. Elytra parallel-sided to about
the middle, thence gently decreasing in width to apex; densely
granulate-punctate throughout, l)ut more densely about base
than elsewhere; striation well-defined throughout. Metasterninn
with a rather wide, shining, median line, lightly impressed along
its middle. Tip of abdomen rather densely granulate-punctate.
Length, 5 mm.
Hah. — N.S.W.: Sydney (A. M. Lea); unique.
From some directions, the pubescence of the elytra appears to
be as dark as that of the prothorax, but, from others, most of it
is seen to Vje paler: the front, produced portion of each of the
fifth-eighth joints of antennae is not much shorter than the
joint itself.
Arisocephalus rufipennis, n.sp.
Black; extreme apex and base of prothorax, elytra, abdomen
(in places feebly infuscated), legs (hind coxae and femora ex-
cepted), and antennae red or reddish. Densely clothed with
short pubescence.
Head with crowded and rather small punctures; with a con-
spicuous median carina traceable to near base, but not on to
clypeus; antennary sockets each with a narrow margining carina
from eye to side at apex of clypeus. Antennae moderately long,
second joint distinctly longer than third, and almost as long as
fourth, fourth-tenth equal in length, but fourth-seventh wider
and more strongly serrated than the ninth and tenth, eleventh
almost as long as the ninth and tenth combined. Prothorax
slightly more transverse, but otherwise much as in preceding
species. Elytra parallel-sided to beyond middle, with dense
punctures about base, becoming less crowded (but quite sharply
defined) elsewhere; striation lightly impressed and, in places,
scarcely traceable. Length, 5 mm.
Hah. — Tasmania: Southport (John O. Dawson); unique.
In general appearance close to the preceding species, but elytra
more brightly coloured, and with different punctures and striae,
second and third joints of antennae difierently proportioned, the
following ones much less conspicuously serrated, and the median
line on the metasternum less conspicuous. The clothing on the
BY A. M. LEA. 735
pale parts is somewhat stramineous; on the dark parts, it is darker.
In this and the following species, the hind coxa; at the sides are
a trifle longer than the second abdominal segment; in the pre-
ceding species, they are of exactly the same length.
ArISOCEPHALUS BASALIS, n.sp.
Black; basal fifth of elytra reddish, tibia? reddish, femora and
coxie darker, tarsi paler, antennae dull reddish-brown, second and
third joints and the tips paler. Clothed with very short and
mostly stramineous pubescence.
Head with crowded and rather small punctures; with a con-
spicuous median carina from base to apex, an oblique carina on
each side of clypeus from apex to base, where they almost touch
the median line between the antennary sockets. Antennae not
very long, second joint short but distinctly longer than third,
the two combined slightly loijger than fourth, fourth-tenth equal
in length, but decreasing in width from the sixth, and rather
feebly serrated, eleventh about once and one-half the length of
tenth. Prothorax much as in A. Jfavipes except that the punc-
tures are smaller. Elytra parallel-sided to about the middle;
with rather dense, well-defined punctures, becoming crowded at
base; striation fairly well defined. Abdomen with fifth segment
somewhat produced at apex, and densely granulate-punctate
there. Length, 3J mm.
^a6.— N.S.W.: Galston (A. M. Lea); unique.
The third joint of the antenna?, although very short, is not
" excessively minute " as in Entomophthahnus; and the fourth
joint, although somewhat larger than the fifth, is scarcely longer.
The carination of the head is more pronounced than in the other
species of the genus, and is much as on Arisus carinatice2ys.
Fornax nicer, n.sp.
Black. Clothed with short, depressed pubescence, paler about
base of prothorax and of elytra than elsewhere.
Head with crowded subasperate punctures; with a feeble longi-
tudinal carina, and a conspicuous interocular one. Antennae
moderately long, first joint about as long as three following com-
bined, second shorter than third, and third than fourth, fourth-
736 NEW SPECIES OF AUSTKALIAN COLEOPTERA, xiv.,
tenth equal in length, eleventh somewhat longer. Prothorax
with front angles somewhat rounded, sides thence parallel to
base ; with punctures as on head ; median line well defined
tow^ards base, but feeble in front. Elytra with crowded, asperate
punctures about base, becoming less crowded and more sharply
defined posteriori}^; striation distinct throughout. Hind cooi:(e
evenly and strongly narrowed to sides, greatest length about
equal to that of second abdominal segment; basal joint of hind
tarsi about as long as the rest combined. Length, 4| mm.
Hab. — W.A. : Mount Barker (R. Helms); unique.
In general appearance, strikingly close to F. siUuralis, but
readily distinguished by the comparative lengths of the third
and fourth joints of antennae. The pubescence on the under-
surface appears ashen or blackish according to the point of view;
on the upper surface, it is almost entirely dark.
Fornax castaneus, n.sp.
Castaneous, tarsi somewhat paler. Densely clothed with
short, stramineous pubescence.
Head strongly convex; with crowded but rather small punc-
tures; inter-anteni)ary carina not continued beyond antennary
sockets. Antennae not very long, second joint slightly longer
than fourth, third almost as long as fourth and fifth combined,
fourth-tenth subquadrate, fourth and fifth short, but combined
considerably longer than sixth, sixth to tenth subequal, eleventh
distinctly longer. Prothorax with sides rounded in front, thence
almost parallel-sided to base; with dense and sharply-defined but
rather small punctures, becoming crowded on sides. Elytra
parallel-sided to near apex; base with crowded subasperate punc-
tures, becoming smaller and more sharply defined posteriorly;
striation well-defined. Hind coxte strongly and evenly narrowed
to sides, which are very short, greatest length slightly more than
that of second abdominal segment; basal joint of hind tarsi dis-
tinctly shorter than the rest combined, fourth slightly narrower
than third, and scarcely produced on lower surface. Length,
6 mm.
i^a6. — N.S.W.: Sydney (A. J. Coates).
feV A. M LEA. 737
In general appearance, strikingly close to some of the larger
specimens, that I have referred, with doubt, to F. parvulus, but
the fourth and fifth joints of antennae, although short, are (com-
bined) distinctly longer than the sixth: the hind tarsi are some-
what aberrant for the genus.
A smaller (4| mm.) specimen from Queensland (Dalby, Mrs.
F. }{. Hobler) appears to belong to the species, but is more
lightly coloured, with a vague remnant of a median line on the
prothorax (completely absent from the type) and with slightly
shorter antennae and legs.
Fornax majorinus, n.sp.
Dark castaneous brown, antennse and legs paler. Very densely
clothed with rather short, stramineous pubescence.
Head with dense but not very large punctures; inter-antennary
carina widely interrupted in middle. Antennae moderately long,
second joint slightly longer than fourth, third almost as long as
fourth and fifth combined, fourth slightly shorter than fifth, and
fifth than sixth, sixth-tenth subequal in length, eleventh almost
as long as ninth and tenth combined. Prothorax with sides
strongly rounded in front, thence parallel-sided to base; punc-
tures dense, rather small and subasperate, becoming crowded on
sides. Elytra parallel-sided to beyond the middle; with dense
asperate punctures about base, becoming smaller posteriorly;
striation well defined. Hind coxce produced to points at the
sides, greatest length distinctly more than that of second ab-
dominal segment; hind tarsi with basal joint as long as the rest
combined. Length, ^\ mm.
Hah. — N.S.W.: Byron Bay (C. Watson); unique.
A comparatively large, robust species.
Dystrigonisthis laticollis, n.sp.
Of a rusty-castaneous, appendages somewhat paler. Densely
clothed with short, rusty pubescence.
Head with rather coarse, crowded punctures; clypeus shallowly
concave. Antennte with second joint short, third about as long
as fourth and fifth combined, fourth-eighth subequal in length
and with rounded sides, ninth almost as long as three preceding
738 NEW SPECIBS OP AUSTRALIAN COLEOPTERA, xiv.,
combined, slightly shorter than eleventh and slightly longer than
tenth. Prothorax with sides strongly rounded in front, and then
increasing in width to base, extreme base distinctly wider than
elytra, and about twice the width of apex, with a rather feeble
median line disappearing before apex; with very dense, and
moderately large, round punctures; with a small, round fovea on
each side, close to middle of base. Elytra parallel-sided to
beyond the middle, base densely granulate-punctate, elsewhere
with fairly dense but smaller and more sharply defined punctures;
striation w^ell-defined, first and second striae on each elytron
opening out into a short, deep, oblique sulcus close to apex.
Hind coxcfi with greatest length about equal to that of second
abdominal segment, oVjliquely decreasing to each side, which is
about one-third the greatest length. Length, 15 mm.
/ya6.— N.S.W.: Mount Irvine (Dr. E. W. Ferguson); unique.
The prothoracic punctures, although dense, are quite sharply-
defined on the disc; on the sides, they are more crowded and
irregular. From some directions, the basal fifth of the elytra
appears to be closely covered with fine, transverse corrugations.
The three, long, terminal joints of antennae are suggestive of (the
Australian species of) Dyscolocer^is, but the deep, lateral channels
of the prosternum at once exclude it from that genus.
It is with some doubt that I refer this and the following species
to Dyst7'iy()7iislhis, with w^hich, however, they would certainly be
associated in Blackburn's Table; the antenme of the two species
dififer considerably from each other, and also from those of B.
paiadoxus, and, in other families of beetles, these differences
would almost certainly be regarded as of generic importance;
but as IJonvouloir, and other workers at the family, have allowed
ail even greater range of variation in the antennae, it does not
appear desirable to propose a new genus (or new genera) for
them at present. The lateral channel on each side of the pro-
sternum is deep and conspicuously closed posteriorly (and receives
the antenna throughout its length); on the basal third of the
inner side, it is distinctly carinated, but, from the basal third to
its front margin, the side is gently rounded, without the least
trace of a carina. Although Blackburn separated Dystriyoiiisthis
BY A. M. LKA. 739
from Phceaocerus by tlie former having " Prosieinal sulcus mar-
gined within by an elevated line only in its hinder part'' as
against " a continuous elevated line," it is to be noted that
Bonvouloir says of PJuenoc-erus ^^Sillon-viargiiial . . . etunt borde
en arriere inter ieurenwnt par une ligne eleve'e.^^ Quite possibly
the following species should have been referred to Phca^iocerus,
but it is evidently distinct from P. siibclavatus by its larger size,
and different antennte and hind cox?e.
Dystrigonisthis ferrugineus, n.bp.
Dark rusty-castaneous, appendages somewhat paler. Densely
clothed with short, rusty-red pubescence.
Head with crowded punctures of moderate size; with a short
and rather wide median line; clypeus shallowly concave. An-
tenna3 rather stout, second joint short, third cylindrical, almost
as long as the three following combined, fourth-eighth short,
subequal and distinctly transverse, ninth and tenth somewhat
longer and wider (and with small fovese at apex), eleventh about
as long as the three preceding combined. Prothorax with sides
rather strongly rounded in front, and then obliquely increasing
in width to near base, which is somewhat wider than elytra, and
almost twice the width of apex; with dense punctures of moder-
ate size, smaller in middle than elsewhere, and becoming crowded
on sides. Elytra feebly decreasing in width from base; densely
granulate-punctate about base, punctures becoming smaller and
sparser posteriorly; striation as in preceding species. Hind coxie
with greatest length about equal to that of second abdominal
segment, curvilinearly decreasing to sides, which are very short;
basal joint of hind tarsi somewhat shorter than the rest com-
bined; second, third, and fourth regularly decreasing in length
and width, fourth not bilobed and scarcely produced on under
surface. Length, 11-12^ mm.
Uab. — Tasmania : Hobart (A. M. Lea).
The antennie so regularly increase in width, that the three
apical joints can scarcely be regarded as forming a club; from
above, the second joint appears to be slightly shorter than the
fourth; but, from below, it is seen to be slightly longer. The
740 NEW SPECIES OF AUSTRALIAN COLEOPTERA, xiv.,
general outlines of the prothoi-cax and elytra are much as figured
for those of Phcenocerus subdavatus (Bonv., Mon., PI. xiii., fig.l).
On one specimen, the eighth and ninth joints of each antenna
appear to be combined to form but one (with the suture com-
pletely obliterated in places); but, as the antennae of the type
are normal, this would appear to be accidental. On this speci-
men, also, there is a short, shining, median line, and two, small,
medio-discal fovese on the pronotum (quite absent from the type).
Ph^nocekus clavicornis, n sp.
Black, antennae and legs somewhat obscurely diluted with red,
tarsi paler. Rather densely clothed with short, ashen pubescence.
Head with small, crowded, partially concealed punctures, inter-
antennary carina not continued across middle. Antennae stout
and not very long, first joint as long as the three following com-
bined, second short, third slightly longer than fourth and fifth
combined, fourth slightly longer than fifth, fifth-eighth short and
transverse, ninth-eleventh forming a conspicuous club, ninth and
tenth each conspicuously wider and longer than eighth, eleventh
as wide as tenth at ba^se, but rapidly narrowing to apex. Pro-
thorax with sides strongly rounded in front, thence parallel-sided
to base, with a slight but almost continuous median line; with
dense and rather small, but sharply defined punctures, becoming
crowded on sides. Elytra feebly diminishing in width from near
base ; densely granulate punctate about base, elsewhere with
small but sharply de6ned punctures; striation well-defined
throughout. Hind coo:(e at sides about one- third their greatest
length, this slightly more than that of second abdominal seg-
ment; basal joint of hind tarsi about as long as the two apical
joints combined. Length, 1\ mm.
Hab. — Tasmania: Hobart (A. M. Lea); unique.
The antennae have a distinctly three-jointed club, a character
which excludes the species from all the genera noted by Bon-
vouloir, except Phceuocerus; but the club is even more distinct
than as figured for P. subdavatus; from the description of that
species, also, it differs in being somewhat smaller, much darker,
and prothorax with a conspicuous median line. In Blackburn's
BY A. M. LKA. 741
Table, Phcenocerus is placed with three other genera distinguished
by having " Prosternal sulcus margined within by a continuous
elevated line." Tliis, however, is not the case with the present
species, whose lateral channel is carinated on the posterior half,
but gently rounded in front; its posterior end is also open.
Galea* Australia, n.sp.
Black, appendages reddish. Densely clothed with golden
pubescence, becoming golden-red in places; on the undersurface
somewhat ashen.
Head with crowded punctures mostly concealed, but more dis-
tinct on clypeus than elsewhere; with a very thin, median carina
from base almost to apex. Antennae rather short, second joint
short, curvilinearly triangular, third-tenth each with a long
ramus, that of third somewhat shorter and thicker .than the
others; eleventh joint slightly longer and thicker than the ramus
of the tenth. Prothorax gibbous, not much wider than long,
sides rounded in front, thence almost parallel to base; disc with
irregularly granulate (in places vermiculate) elevations; the sides
with irregular, more or less concealed punctures. Elytra nar-
rowed from base to apex, tips obliquely carinated and produced;
with series of fairly large punctures, becoming smaller posteriorly,
but close to apex becoming larger. Hind coxcb with posterior
edge somewhat sinuous, outer edge quite as long as elsewhere;
tarsi with second, third, and fourth joints each with a wide and
conspicuous lamella. Length, ll-14mm.
//a6.— Queensland : Coen River (H. Hacker).
The first of its genusf to be recorded by name from Australia,
although Bonvouloir (Mon., p. 807) stated that the genus occurs
there; Blackburn, however, thought! that this may have been
due to confusion with Galbodema, there recorded as a synonym
of Gcdha^ but elsewhere (Mon., p. 442) treated as distinct. The
beautiful pubescence on the whole of the upper surface has a
*Gu6r.-Men., Voy. Coq., Entom., p.68; Bonv., Men., p.806.
+ Readily recognisable by the large size and robust form of its species,
with three tarsal joints conspicuously lamellate.
% Manuscript note.
742 NEW SPECIES OF AUSTRALIAN COLEOPTERA, xiv.,
curious waved or mottled appearance, due to the varying direc-
tions in which it is applied to the derm; thus, the elytra, when
viewed from behind, appear to have two golden fasciae alternated
with brownish ones; on altering the point of view, the brownish
ones become golden, and vice-versd, whilst, from other jdirections,
the clothing appears to consist of irregularly mixed golden and
brownish spots. The third joint of the antennae is about as long
as the two following combined, and its ramus is slightly shorter
than the first joint, the rami of the other joints being slightly
longer and subequal inter se. The elevations on the prothorax
may be regarded as forming an irregularly lobed mass, of which
the median lobe is the most conspicuous, appearing as an abrupt,
obtusely pointed elevation near the scutellum (from the sides,
its posterior end appears vertical), then, towards the middle, it
bifurcates, each arm being irregularly continued to near the
apex; on each side of it there are two other elevations feebly
connected with it, the front one somewhat oval, the back one
concave inwards and almost right-angled outwards; of the species
figured by Bonvouloir, the nearest approach to this structure is
that of G. wallacei (PI. xL, fig.l), but the elevations are all some-
what different; they are clothed with more ruddy pubescence
than on the adjacent parts. The rows of elytral punctures are
in very feeble striae, but, about the tip, the striae become deep;
the derm is densely covered with minute punctures, but these
become visible only after abrasion; the sides of the elytra, from
the base of the fifth abdominal segment, are obliquely flattened,
so as to appear as enlargements of the (elsewhere extremely
narrow) epipleurae, with the upper edge acutely ridged, and, at
the suture, rather acutely produced.
PYTHID^.
NOTOSALPINGUS BRUNNEUS, n.Sp.
Obscure piceous-brown, basal half of antennae and legs some-
what paler. Clothed with short and sparse but fairly distinct
puljescence, more noticeable about apex of elytra than elsewhere.
Head wide and gently convex, with two shallow depressions in
front, separated by a short, shining space; with crowded, sharply
BV A. M. LEA. 743
defined punctures. Antennae rather thin, extending to base of
prothorax, slightly dilated to apex. Prothorax lightl}' trans-
verse, sides regularly increasing in width from base to near apex,
and then rather abruptly narrowed, on apical half with several
very feeble denticulations; with dense and moderately coarse
punctures, more crowded on sides than on middle. Elytra at
V)ase slightly wider than head across eyes, shoulders square, sides
subparallel or feebly dilated to near apex, and then widely
rounded: with distinct rows of fairly large punctures, larger and
more irregular about base than elsewhere, and becoming rather
small posteriorly. Length, 1^-2 mm.
Hah. — Tasmania: Hobart, Launceston, Bruni Island. — AVestern
Australia: Swan River, Newcastle (A. M. Lea).
A depressed, densely punctate species, readily distinguished
from all others of the genus by the uniformly dingy-brown elytra.
The apical joints of the antennae are gradually enlarged, not
abruptly clavate as in Neosalpinyns. The denticulations on the
sides of the prothorax are so very feeble that, from most direc-
tions, they are quite invisible; one specimen has a shining median
line on the basal half of the pronotum; and, on several, there
are two vague basal depressions; the elytra are without striae,
although their punctures are in very evident rows.
Tasmosalpingus, n.g.
Head wide, obtusely produced in front. Eyes small, lateral,
prominent and coarsely faceted . Antennae inserted considerably
in front of eyes; with a conspicuous, three-jointed club. Pro-
thorax wide, sides acutely margined. Scutellum small and
strongly transverse. Elytra short. Metasternum elongate.
Leys not very long; front coxae rather widely, the others moder-
ately separated, front coxal cavities open behind; tibiae dilated
towards, and minutely spurred at apex; tarsi moderately long,
two basal joints of hind pair and three of the others moderately
wide and close together, penultimate small and simple, claw-joint
rather stout, almost as long as the rest combined; claws swollen
towards base but not dentate.
The acutely carinated margins of prothorax, each separated by
744 NEW SPECIES OP AUSTRALIAN COLEOPTERA, xiv.,
a narrow gutter from the disc, and rather widely separated front
cox^e, read il}" distinguish the genus from Notosalpinyua, to which,
at first glance, the two species appear to belong. Type of genus,
T. quadrispilotais.
Tasmosalpingus quadrispilotus, n.sp.
Brassy-black; undersurface, legs, and antennae (club excepted)
paler, elytra piceous-brown, with four, large, pale spots. Head
and prothorax with distinct but somewhat straggling pubescence,
elytra glabrous.
Head obliquely flattened, as wide across eyes as length of
antennae, with a shallow depression on each side in front; punc-
tures dense and sharply defined but not very large, becoming
smaller in front. Antennae with second joint almost as stout as
first but distinctly shorter, third-eighth small. Prothorax at
widest (which is near the apex) slightly wider than head, apex
distinctly wider than base, each side of base distinctly impressed,
margins rather acutely carinated throughout but incurved at
base; punctures much as on head. Elytra at base about the
width of prothorax at its widest, feebly dilated to beyond the
middle and then widely rounded; with rather coarse punctures
in distinct but irregular series near base, becoming smaller and
irregularly disposed elsewhere. Length, 1^-1 1 mm.
Hab. — Tasmania: Mount Wellington, in moss; Launceston (A.
M. Lea).
A short, dumpy species, with prothorax rather conspicuously
clothed, and elytra glabrous. On the type, the first spot on each
elytron is a large, irregular one, commencing on the shoulder
and obliquely directed towards the suture, near which it termin-
ates at the basal third; the second one commences just beyond
the middle, and is obliquely subtriangular. On one specimen,
the pale markings are considerably enlarged, so that they appear
to be the ground-colour, with the infuscate portions marginal,
sutural, and forming three, large, median spots; the smaller one,
on the suture, narrowly connected with the others, and these
connected with the marginal infuscation.
BY A. M. LEA. 74.7
Tasmosalpingus promiscuus, n.sp.
Dark piceous-brown, mitennse (club excepted) and legs slightly
paler, elytra with obscurely flavous markings. Head and pro-
thorax with rather conspicuous pale pubescence, elytra glabrous.
Length, 1^-1 1 mm.
Hab. -Tasmania: Hobart, Swansea (A. M. Lea).
Structurally, close to the preceding species, but elytral punc-
tures distinctly smaller and nowhere seriate in arrangement.
The elytral markings, although conspicuous, are not sharply
defined, and consist, on each elytron, of a humeral spot, lightly
or not at all connected with a small subsutural one at the basal
third; and a larger, post-median spot, fairly wide near the suture,
and strongly narrowed obliquely upwards and outwards.
Neosalpingus brevis, n.sp.
Black, appendages (club excepted) obscurely paler; head and
prothorax with a slight bluish or greenish gloss.
Hedd wide, gently convex between eyes, and somewhat
flattened in front, with very minute punctures. Eyes very
widely separated, and almost basal. Antennae moderately long,
third-eighth joints small, the three following forming an abrupt
and conspicuous club. Prothorax widely transverse, rather
strongly convex; base and apex of subequal width, sides rounded,
and each with four, small, unisetose denticulations, each side of
base with a small depression; with numerous, small but rather
sharply defined punctures, becoming somewhat crowded and
larger on sides. Elytra short, at base slightly wider than pro-
thorax, sides distinctly dilated to about the middle, and then
widely rounded to apex: smooth and apparently impunctate.
Length, 1-1 1 mm.
Hah. — Queensland: Cairns (Macleay Museum, and E. Allen).
In general appearance, like small, dumpy specimens of N.
politus, but prothoracic punctures considerably smaller, basal
impressions smaller and oblique, head smoother, and elytra con-
siderably more dilated. The upper surface is entirely glabrous,
the undersurface almost so. The head, when seen from the front,
appears to be of subtriangular shape, but with sinuous sides; the
58
74 6 NEW SPECIKS OF AUSTRAI>IAN COLEOPTERA, xiv.
elytra might fairly be regarded as impunctate. as tlie punctures
are so extremely small that it is only from certain directions, and
in a good light, that they may be seen under a ^'oddington lens.
CURCULIONID^.
Cyrotyphus variegatus, n Sp.
Black, parts of tarsi obscurely diluted with. j>'d. Densely
clothed with variegated, depressed pubescence or setfie, the elytra
with numerous, small fascicles.
Head and base of rostrum with dense, normally concealed
punctures of moderate size, apical half of rostrum with small,
dense punctures. Antennte moderately long, third joint twice
the length of second, and distinctly longer than fourth, eleventh
about once and one-half the length of tenth. Profhorax about
as long as basal width, which is considerably more than that of
apex, sides bisinuate, with an irregular ridge on each side of
middle, rising into a small tubercle near base, each side with a
feeble, granulate elevation: with dense, more or less concealed
punctures, and a few scattered granules. Elytra much wider
than prothorax, almost parallel-sided to near apex, each elytron
with two obtuse ridges near base, the inner one moderately
elevated about summit of apical slope ; with small, distinct
granules about base, but other granules, and dense punctures,
normally concealed. FHmora stout, rather lightly but distinctly
dentate. Length, 13 mm.
Hah. — Tasmania (J. E. Philp); unique.
Readily distinguished from C fasc'Cidaius, by the prothorax
being without a conspicuous, transverse series of four tubercles;
the elytra also are much more conspicuously variegated. The
apical half of the rostrum is glabrous, but all other parts are
more or less densely clothed, the pubescence being mostly stra-
mineous, but varving to white (there is a conspicuous, bisinuate,
white mark traversing the suture slightly beyond the middle)
and black; on the elytra, there are numerous, small fascicles
(especially along the suture), all of which are black: the abdomen
has a vaguely striped appearance.
74
OX A NEW SPECIES OR FORM OF EUCALYPTUS.
By Cuthbert Hall, M.D.. Ch.M., Parramatta.
(Plate Ixxv.)
Eucalyptus Marsdeni, f. vel sp.nov.
Arbor, 30'-50' altitudine, eortice fibrose inferne superne kievi,
foliis petiolatis, lanceolatis, acuminatis, falcatis, obliquis, fere
membraneis; cyiiiis axillaribus; pediiiiculis 4'" loiigis, pedicellis
Ih" ', operculo hemisplia?ric'o, uniboiiato; fructibus hemisphiiericis,
valvis par u 111 exsertis.
A tree, 30 feet high in specimen observed, and probably would
attain a height of 60-80 feet when fully grown.
Seedling. — Cotyledons ver}^ small, orbicular-reniform, entire,
purplish on undersurface, glabrous. Leaves oj^posite, decussate,
obtuse, shortly petiolate, lanceolate, venation pinnate, rather
oblique, edges sinuate. Stem reddish, and both it and the leaves
covered with fine, stellate hairs.
Jdvenile leaves of a more advanced stage than in the small
seedling are alternate, petiolate, narrow-lanceolate, acuminate,
glabrous. Mature leaves alternate, petiolate, lanceolate, falcate,
acuminate, obli(jue, greyish on dryiiig, almost membranous, occa-
sionally shiny, and having a pleasant, aromatic scent. Lamina?
6"-8" long by j" broad, petiole slender, .V" long. Lateral veins
oblicjue, alternately fine, intramarginal vein fairly distant from
€dge.
Inflorescence axillary, peduncles h' long, with rather few
flowers in head, 6-9; buds turbinate, 5'" long, operculum hemi-
spherical, shortly acuminate. Stamens all fertile, anthers kidney-
shaped. Fruits hemispherical, 3'" in diameter, rim domed, valves
small, slightly exserted.
Jiark of an unusual character for a Eucalypt. While it falls
in the group of the stringy-barks, yet it is laminated, with a sort
of ochreous deposit on the outer surface of each layer. Inner
748 NEW SPECIES OR FORM OF EUCALYPTUS,
bark very hard and compact. But while the trunk and lower
branches have such bark, that of the upper branches and branch-
lets is smooth and greyish, so that the tree is really a half Vjark.
Timber light brown in colour, fairly heavy, close, straight in
the grain, annual rings prominent in the young stage, planes
and dresses well, and should be useful for technical purposes;
gum-veins few.
Locality. — Toongabbie, New 8outh Wales, at the rear of the
Public School, on the Wianamatta clay.
Remarks. — As seen from the description, this form of Euca-
lypt, on a cortical classification, seems intermediate between the
smooth-barks and the stringy-barks. The timber has not the
texture of that of the stringy-barks, but more nearly resembles
that of E. viminalis in physical characters. The early buds
resemble those of E. obliqua, but there is no resemblance in the
mature stage. The mature leaves are generally markedly oblique.
The fruit resembles that of E. eugenioides, but it tapers more
into the pedicel, and is not so flat; nor are the fruits so clustered
on the peduncle. The seedling is intermediate between those of
E. eugenioides and E. Jloorei; and, in its hair}' seedling-leaves
and reniform cotyledons, approximates strongly to the string-
barks. The reniform anthers also place it in that category, but
the bark, timber, and oil, are quite distinct from those of this
class. As, so far, only a single tree is known, one is strongly
inclined to conclude that it is either a hybrid or a sport. Strong-^
colour is lent to the hybrid theory by the fact of its possessing
so many of the characters of the stringj^-barks, especially in the
seedling-stage; yet differing from them in others in the mature
stage, as for instance in the bark, oil, and timber. Since the
only known tree has, unfortunately, lately been cut down, further
comparison is at present impossible. Now that a description
has been published, search may reveal further specimens, and
more definitely establish its status. The tree was a young one,
about 12-15 years old, and growing on land that had been mostly
cleared, but with a few well grown trees of E. hcemastom'f, E.
resinijh'a, and E. siderophloia in proximity. Other trees near l)y
BY CUTFIRKRT HALL. 749
were E. crehra, E. eugenioides, E. hemlphloia, E. i->nnctata^ and
E. tereticornis . I have named this form or species, tentatively,
E. Marsdeni, after the Rev. Samuel Marsden, the first incumbent
of St. John's Church, Parramatta.
Oil. — Mr. H. G. Smith reports tliat the oil of this Eucalyptus
was obtained by steam-distillation, in the ordinary way, from the
leaves and terminal branches. It was somewhat thick and vis-
cous, having almost the viscosity of castor oil. It has little
resemblance, either in appearance or constitution, to the majority
of ordinary Eucalyptus oils, but is more closely allied to the oils
distilled from E. nova-anylica and a few others. It has no re-
semblance to the oils of the members of the E. tereticm^nis-gvow^.
Cineol was practically absent, and phellandrene could not be
detected. The amount of oil available did not permit of the
active terpene being isolated in a pure condition, but it is pro-
bable that dextro-rotatory pinene occurs in small amount.
Only 38 pounds of material were received for distillation, from
which 4 ounces of oil were collected, equal to 0'66 percent. The
oil was but little coloured, and had a fairly pleasant odour. It
gave the following results: —
Specific gravity at 15°C. = 0"9469.
Rotation aD = +4-8°.
Refractive index at •20°C. = 1-4989.
Insoluble in 10 volumes 80 per cent alcohol at 16°C.
Saponification number of esters and free acid = 2, so that
esters were practically absent. The small amount of free acid
was removed, and this gave a strong odour of butyric acid.
The phenol gave the reactions for Tasmanol, but it is probable
that both phenols occur. The indications are that the oil con-
sists largely of sesquiterpenes. Probably some high-boiling
alcohols are also present, as indicated by the odour.
I have to record my sincere thanks to Messrs. R. T. Baker
and H. G. Smith for much help in preparing this paper.
EXPLANATION OF PLATE LXXV.
Portion of branchlet of Emalyptns Marsdeni, f. \el sp.n.
59
750
STUDIES IN AUSTRALIAN NEUROPTERA.
No. 6. The Family Psycho psid.f., with Descriptions of new
Genera and Species.
By R. J. TiLLYARD, M.A., D.Sc, F.L.S., F.E.S., Linnean
Macleay Fellow of the Society in Zoology.
(Plates Ixxvi.-lxxviii.; and four Text-figures.)
In No.4 of this series of Studies (15, pp.271, 289), I gave my
reasons for considering the Psycho2)sid(e to be a separate family,
and stated that it was my intention to deal with these insects in
a monograph to follow later.
In a paper read in October, 1915(13), but unfortunatel}^ not
bearing upon it any indication of the actual date of publication,
Navas published, in Spain, a monograph of these insects, and
classified them as a separate family. This paper has only recently
reached me, and I cannot tell whether it should take priority
over mine or not; however, in view of the earlier date of reading
of Navas' paper, I feel that I must give him the credit of having
first raised these insects to family rank.
The conditions arising from the War have made it impossible
for me to receive, for study, the material that I had hoped for,
in the non- Australian species of this family. For this reason,
as well as because of the publication of Navas' paper, I have
decided to abandon my projected monograph, and to confine
myself mainly to a study of the Australian species. Only in so
far as this study aflfects the classification of the family, will it be
necessary to discuss the other genera and species.
During the past three years, a large amount of work has been
carried out upon this family. Of special importance has been
the working-out in detail of the life-history of one species,
Psychopsis flegans (Guerin), and the dissection and photographing
BY R. J. TILLYARD. 751
of the wings of the pupa, in order to study the tracheation.
Including a considerable number of bred specimens of this species,
I have had through my hands for study, at one time or another,
about one hundred examples of these rare insects, and have thus
been enabled to gain a first-hand knowledge of all the Australian
species except one. That exception is Psi/chopsis meyricki
McLachlan, of which the only known specimens are in the
McLachlan Collection, and are, therefore, not available for study.
In the present paper, I propose to deal fully with the wing-
venation of the family, firstly by a study of the pupal wing-
tracheation, and secondly by applying the results so obtained to
elucidating the best method of founding genera in the family.
This matter is one of considerable difficulty, owing to the essen-
tial variability of the characters selected for this purpose by
Navas, viz., the amount of cross- venation present. Descriptions
of new genera and species will also be included, together with a
general discussion of the relationships of the family. The full
account of the life-history of Psychopsis eleyans (Guerin) will
form a separate paper, No. 7 of this series of Studies.
My thanks are due to Dr. R. Hamlyn Harris, Mr. H. Long-
man, and Mr. H. Hacker, for the three fine photomicrographs
reproduced in Plate Ixxvi.
The Venation of Psy chop sis eleyans (Guerin).
In Text-fig. 1, the complete venation of Psychojjsis eleyans
(Guerin), is shown. Text-figs. 2, 3 A, give, for comparison, portions
of the precedent tracheation of the pupal wing, drawn by means
of the camera lucida from freshly dissected pupal wings.
It will be seen at once that, broadly speaking, the venation
and the precedent tracheation agree remarkably closely, as is the
case with all the more archaic members of the Order Planipennia.
Tracheae precede every main vein, right down to its tiniest
branchlets; but no tracheae precede the cross-veins, which are
only lightly indicated on the pupal wing. It is, therefore, very
necessary that we should be careful to distinguish between the
true cross-veins in the imaginal wintj, and those veinlets (or
752
STUDIES IN AUSTRALIAN NEUROPTERA, VI.
Text-fig. 1.
Psychopsis elegans (Guerin). Wing-venation; ( x 5). lA, first analis; 2A,
second analis or basilar; 3 A, third analis or axillary; ap, apex; C,
costa; cov, costal gradate series of cross- veins; cr, recurrent costal or
humeral veinlet; Cu, cubitus; Cuj, Cus, its branches; gi, discal gra-
date series of cross-veins; ^2» internal gradate series of cross-veins;
M, media; Mi, M.^, its branches; M^-Cui, anastomosed portion of
M2 withCui; R, radius; Rs, radial sector; Si -Si 3, its branches; tji,
tornus; fr, terminal gradate series of cross- veins.
BY R. j. TILLYARb.
753
braiichlets of main veins) which arc preceded by trache.T. For
example, the veinlets passing from 8c to the anterior border are
true veinlets or branches of Sc; but, in families where they are
not so prominently developed as in the l\sijchoj)sid(e, they are
frequently spoken of as cross-veins. The small veins which join
these veinlets together, forming a longitudinal series of gradate
Text-fig.2.
P-sf/rhopsIs eJegans (Giierin). Tracheation of basal half of pupal wing;
( X 18). Lettering as in Text-fig. 1.
veins, are true cross-veins, and are to be regarded as a com-
paratively late acquisition, correlated with the broadening out
of the costal area in this family. So, also, the three other series
of gradate veins are true cross-veins, not being preceded by
tracheie. A further point of distinction between veinlets and
cross- veins, in this and all other archaic families of Planipennia,
is that the veinlets, like the main veins of which they are part,
are hairy, whereas the cross-veins are not.
754
STUDIES IN AUSTRALIAN NEUROPTERA, vi.,
The most stiiking specialisation in tlie venation of the Psych-
ojysidce, apart from the great enlargement (jf the costal area, is
the manner in which the three parallel main veins, 8c, R, and
Rs, have become strengthened, for from two-thirds to three-
quarters of their lengths, to form a kind of midrib, from which
the rest of the wing, apart from the much reduced median,
cubital and anal areas, is supplied with numerous, delicate,
radiating veins. I propose to term this strengthened portion of
these three veins the vena trip/ lea; its distal end is already known
as the anastomosis. In order to understand this structure, we
must consult the precedent tracheation. There (Text-fig. 3, A) we
Text-fig. 3.
P.^f/cJiopsIs ehf/ans (Guerin). The radial anastomosis in the forewing. A,
pupal tracheation; E, imaginal venation; .Vj, .I'a, the two cross-veins
which join the distal ends of the vena triplica.
shall find that the trachete run normally through the anastomosis,
without any thickening basad to it, and without any true anasto-
mosing between them. The strengthening of these veins, from
the base to the anastomosis, to form the vena triplica, their ap-
parent unions at the anastomosis, and the strong differentiation
of their distal portions so as to appear exactly the same as the
branch-veins above and below them, are purely imaginal special-
isations, peculiar to this family, and not found elsewhere in the
Planipennia. The development of the irregular series of cross-
BY K. J. TILLYARD. 755
veins between So, R, and Rs may also be correlated with the
development of the vena triplica, as an additional strengthening
for the midrib.
Let us now study the so-called anastomosis more fully. In
Text-fig. 3, A and B, I have contrasted the pupal tracheation
with the imaginal venation in this region. It will be seen at
once that the actual termination of the vena triplica is not
accomplished by means of an anastomosis between any of the
main veins, but by the interpolation of the two cross-veins, x^ and
.x'2, together with the bending-down of the portions of Sc and
Ri preceding them. We must contrast this strongly with the
condition to be seen in the Osmylidce, where Sc and Rj approach
one another distally at a very slight angle, and then fuse together,
without the iiitervention of an auxiliary cross-vein, for a consider-
able distance. There is then, in the Osmylidcn, a true anasto-
mosis between these two veins; whereas, in the Psychopsidce,
l-i
M;
■Ml
Text-fig. 4.
Types of anastomosis between IVI and Cu in Psychojjxis. a, usual type in
Ps. mimica Newman, forewing; h, usual type in Ps. elegans (Guerin),
forewing. In Ps. i)iso/e)is McLach., there is usuallj^ no anastomosis in
this region at all. Lettering as in Text-fig. 1.
there is no true anastomosis, and it would appear to be somewhat
misleading to term it so, as Navas has done. However, the term
has already been applied, in Perlaria and Trichoptera, to junc-
tions between main veins, brought about by means of cross- veins
in line with one another; so that we may allow it to remain,
provided that we understand that we are speaking of a junction
of this type, and not a true anastomosis between two or more
main veins.
756 STUDIES IN AUSTRALIAN NEUROPTERA, vi,,
Turning next to the region of the media and cubitus, on the
character of wliich Navas attempts to divide the Australian
species from the African, as two separate tribes, on the ground
that the former have no anastomosis in this region, the condition
to be seen in PsychopsU eletjans is shown in Text-fig. 1 and Text-
fig. 4, b. In the tracheation, M._, approaches Cu^ very closely for
a considerable distance, and then diverges from it again. In the
corresponding venation, M2 fuses with Cu^ for this same distance,
and the parts of Mg just before the fusion, and of Cuj just after
it, are bent so as to appear like oblique cross-veins. It is cleai'
that here we have a true anastomosis, comparable with that
between Sc and R already mentioned in Osmylidce. But this
anastomosis is not a constant character, either for the species
elegans or for the genus Psychopsis, as I have noted more fully
on p. 761. The type of anastomosis in this region in Psychojysis
inimica, in which, usually, Mj just anastomoses with Mo at a
point, and Mg in its turn with Cuj, is shown in Text-fig. 4, a. In
other species, such as Ps. insolens, there may be, in the majority
of specimens, no anastomosis whatever in this region.
The arrangement of the cross-veins, which are not represented
in the precedent tracheation, is of great interest. In Ps. elegans
(Text-fig. 1), we note, first of all, the presence of a gradate series
closing the disc distally (f/i). This may be termed the diseal
gradate series. Internally to this, and crossing the disc not far
from its middle, there is another series {g^, which may be called
the iiiternal gradate series. These two series will be at once
recognised as the homologues of the two gradate series normally
present in the wings of (Jhrysopid(e. Besides these, there will be
seen a long series of cross-veins connecting the veinlets of the
costal space, and extending right from near the base to the apex.
This may be termed the costal gradate series (cov). Below the
apex, this series is continuous with a gradate series roughly
parallel to the discal series, but lying between it and the termen
of the wing. This may be called the terminal gradate series {tv).
Except for a slight development of costal cross- veins in such
genera as Drepanepteryx, the costal gradate series does not appear
bV R. .t. TiLLYARD. ' /0<
in any of the lower Plaiiipeiinia; the terniiiial gradate series is
absolutely peculiar to the Faychojjsidw.
Of these series, only the discal is invariably present. The
internal series is usually present also, but not always. The costal
and terminal series are later developments, confined to the larger
species, and evidently correlated with the greater expansion of
the areas which they help to support. Further, there is great
\ ariability in the condition of these series in almost all species.
As an example of this, T would point out, in Text-fig. 1, the
presence of a single cross-vein in the forewing between the
anterior ends of the internal and discal series. This is frequently
absent; but, in some specimens, it may be replaced by either
half of an extra series, in others by a complete series of as many
as thirteen cross-veins, crossing the disc between the internal
and the discal series ! In one case, I noted the presence of this
extra series upon one forewing of a specimen, while it was absent
upon the other I It will be at once obvious that attempts to
define genera on such variable characters as these, are foredoomed
to failure.
In conclusion, it is interesting to note that, regarded as a
whole, the wing of Psychopsis presents a very close analogy in
structure to the feather of the wing of a bird — an analogy which
is also suggested by such wings as that of Drepant'ptery.n^ but by
no means so closely. The vena triplica corresponds with the
rachis, forming the main support or midrib in both cases; the
position of this support is the same in both cases, viz., nearer to
the anterior than to the posterior margin. The numerous veins
radiating out from both sides of the vena triplica are the
analogues of the barbs of the bird's feather; and the series of
connecting cross-veins, or gradate series, are analogues of the
barbules, which serve to connect the barbs together.
Classification of the Family.
We are now in a better position to consider the true value of
the various venational structures which have so far been used in
subdividing it into tribes, and in defining genera.
758 STUDIES IN AUSTRALIAN NEUROPTERA, Vl.,
Our study of the venation shows us that the cross-veins are
not preceded by trachejt?, and that they are to be considered as
a later addition to the main sclieme of venation. This appUes
more especially to the costal series of cross-veins and its continu-
ation, the terminal gradate series, which are not fully developed
in any other family, and are evidently comparatively recent
acquirements, correlated w itli the great enlargement of the costal
area. With regard to the discal and internal gradate series, it
must be evident that vaiiations in the amount and position of
the cross-veins formijig them will depend partly on the size of
the insect, and partly upon the number of branches of the radial
sector possessed by it; we should, therefore, be prepared to find
a great deal of variation in these characters. If, then, the
present system of classification should fail to stand the test that
we propose to apply to it, we shall be faced with the problem of
finding more reliable characters, on which to rebuild a sounder
arrangement of the species.
Though it is not the purpose of this paper to deal exhaustively
with the non- Australian species, it will be first of all necessary
to call attention to certain errors and omissions in Navas' most
recent work. A paper that claims to be a monograph of a family
should surely contain a complete bibliography of all the more
important papers concerning it, and should also at least list, if
not describe, all the known species. But I find that Navas
omits from his work all reference to two of McLachlan's most
important papers (9, 10), in which that fine entomologist described
a new South African species on the one hand, and the only
known Burmese species on the other (Fsi/chopsis mar shall i^oLi.,
and Ps. binnana McL., respectively). Consequently, Navas
gives only four species instead of five for South Africa (unless,
indeed, one of his own species is synonymous with McLachlan's I),
and is quite unaware that a Burmese species exists at all I As
there are only fifteen species known altogether, it will be seen
that these omissions are veiy serious for a monograph.
We must now note some further errors of equal gravity. In
1910, Navas described specimens of Pst/chopsis elegans (Guerin),
fiV R. J. TtLLYARb. 759
surely one of the best known and most typical species of the
family, as a new genus and species of Jlenierob'ddai^ under the
name ZygoiMebius rrrnaitxlniis Navas{ll). The description
clearly shows that, at the time, Navas was quite unaware that
there was such a genus as Fsychopsis, though it had been pro-
posed nearly seventy years before, and its type-species was one
of the most striking of all known Planipennia. Later in the
same year, N. Banks(l) pointed out that Zijijophldnus was clearly
synonymous with FsychopsU', whereupon, in his next publication,
Navas suppressed his generic name in favour of Fsychopsis. If
he had stopped there, all would have been well. But, in his
monograph (13), he proceeded to resurrect Guerin's old generic
name Arteriopteryx (given by that author in ignorance of New-
man's then quite recently published name, Psychopsis), the type
of which is eleymis Guerin, synonymous with vei-reauxinus Navas.
He then, quite illegally, further resurrects his generic name
Zygophlehius, of which the original type was verreauxinus Navas
( = elegans Guerin), and proceeds to redefine it, and to give it a
new genotype, Psychopsis zebra Brauer, from South Africa. At
the same time, he forms the tribe Zyyophlebini to receive the
whole of the South African species, leaving eleyans Guerin,
together with all the other Australian species, in the tribe
Psychopsini !
Now Zyyophlebius was proposed by Navas as a monotypic
genus, and hence there can be no question as to which species
was its type. Hence, whether Navas was correct in placing that
species in Psychopsis, as he did in 191 2, or in Arteriopteryx, as
he did in 1916, is immaterial to the question. In either case,
Zyyo2)hlebius sinks as a synonym, either of Psychopsis Newman,
or of Arterio2)teryx Guerin. As a nomen niidain, it may not
again be resurrected in the form of a new genus, to receive, as
its type, another species of Psychopsis, viz., Ps. zebra Brauer.
Further, the tribal name Zygophlebhii must sink with it. If
Ps. zebra Brauer, is really generically distinct from Ps. niiniica
Newman, it now requires a new name. I therefore propose the
name Notopsychop>s, n.n., to replace Zygophlebias Navas, 1916,
760 STUDIES IN AUSTRALIAN NEUROPTERA, vi.,
(type zebra Br.), invalidated by ZyyophlpMns Navas, 1910, (type
verranxinus Navas = eZe^mii? Guerin) itself a synonym for Arteri-
opfery.v Guerin, synonymous with P^ychopsis Newman.
Having thus cleared the ground to some extent, let us now
pass to the consideration of the division of the family into tribes,
as proposed by Navas in 1916 (13, p. 188). I give herew^ith his
key, translated from the original Spanish : —
(1)
(2)
Costal area of both wangs divided, at least partly, by a median
longitudinal series of gradate veinlets {reniUas en esculinata) 2.
Costal area simple or totally undivided in both wings. Asiatic
species 3. Balmesini, nov.
Two anastomoses in both wings; the anterior formed by the
subcostal, the radius and its sector, which are united in the
region of the stigma, the posterior by the procubitus { = M),
the cubitus and its sector, which are connected in a similar
manner. African species 2. Zygophhhini Navas, restr.
One evident anastomosis, viz., the anterior or radial, formed by
the three veins mentioned; the others run parallel without
bending manifestly in an anastomosis. Australian species...
1 . Psychojjsmi, nov.
In his description of Psychopsis hirmana{2), which Navas has
evidently not seen, McLachlan says: — "It differs from all Aus-
tralian species except Ps. 7ney?'icki in the absence of a line of
oblique connecting nervules in the costal area" ; and, lower down,
he divides the known species of Psychopsis into two groups, of
which the first, consisting of Ps. meyricki and Ps. hirmana, is
defined by the words "No line of oblique nervules in costal area,
only two series of transverse gradate veins."
The facts of the case are that Ps. meyrirkl normally lacks the
costal series of cross-veins, as McLachlan here states. Navas,
however, who admits that he has never seen this species, is com-
pelled to place it in his geiuis Magallanes, together wdth Ps.
hisohns McLach., though he defines this genus as having the
costal space of the forewing subdivided by the presence of the
series of costal cross-veins, which Ps. meyricki lacks. For, if he
had anywhere mentioned this character, which McLachlan
emphasises, his tribe Balmesini at once breaks down !
BV R, J. TILLYARD.
61
Here, then, we have an instance of an entirely artificial sub-
division into two tribes, Balmesini and Psychopsini. We may
well ask, what is the essential point in the subdivision"? Is it
to divide the Asiatic from the Australian species'? If so, then
the division on the character of the costal space breaks down,
and Ps. meyricki, agreeing with the Asiatic species in this
respect, must nevei'theless stand apart from them in the tribal
division. But if it is the character of the costal cross-veins that
is the essential matter, the case becomes even worse; for then
the Australian species Ps. meyricki must at once be removed
from the tribe Psychopsini to the tribe Balmesini (all the rest of
which are Asiatic species), and must also be removed from the
genus J/agallanes, of which its close ally, Ps. insoJens McL., is
the genotype, and find its proper place in the Asiatic genus
Bahnes !
There is only one escape from this dilemma, and that is, to
admit that the tribal characters here under discussion are of no
value, and to suppress the tribe Balmesini altogether.
Turning next to the characters of the tribe Zygophlehini Navas,
as here defined, it is only necessary to draw attention to my
remarks on the condition of the venation of Psychopsis elegans
in the region of M and Cu (p. 756). Navas' definition for the
tribe Zygophlehini indicates the presence of an anastomosis in
the region of these two veins for this tribe, but the absence of
the same anastomosis in the tribe Psychojysini. One would have
Mttle hesitation in deciding that both Psychopsis mimica New-
man, and Ps. elega7is Guerin, must go into the tribe Zygoj^hlehini,
on this definition. Whether the form of anastomosis is the same
in these species as it is in the African ones, I am not able to
say; the point is, that the amount of anastomosis in this region
is very variable, not only for diff'erent species, but also for dif-
ferent individuals of the same species. There is only one species
known to me, viz., Ps. insolens McL., in which it can truly be
said that the great majority of the known individuals have no
anastomosis in this region; hence, strictly speaking, this is the
only species that can be rightly admitted into the tribe
762 STUDIES IN AUSTRALIAN NEUROPTERA, vi.,
Psychopsini, as defined by Navas. Thus we come to a rpductio
ad ahsurditm, which it is not necessary to pursue any further.
The only remedy is to admit, at once, that the tribal characters
under discussion are of no value, and to suppress the tribe Zygo-
phlebini altogether. We are then left with only the tribal name
Psychopsini, which ceases to have any value as restricted by
Navas' definition, and may be disregarded; leaving us with a
single family of closely united species, the Psychopsidce, within
which no legitimate tribal divisions can so far be recognised.
Having settled these questions, we may now proceed to define
the family Psychopsidce afresh, and to deal more fully with the
Australian genera and species. This will necessitate a discussion
of the validity of the genera proposed by Navas for the Austra-
lian species, a re-grouping of the species, and descriptions of two
new genera and species.
Family PSYCHOPSID^.
Head: size small or medium; eyes large and rounded, well
separated, projecting laterally ; ucelli absent or vestigial ;
antenncB short, set fairly close together, many-jointed. Mandihles
small, with only a single internal tooth below apex; ma.vilke with
five-jointed palpi; labial palpi three-jointed.
Thorax: broad and strongly built; ])7'olho7'ax short and com-
paratively narrow; mesothorax very broad and large; metathorax
also broad, much shorter than mesothoi-ax, and not fused with
it. Legs short, the tibiae with a pair of terminal spurs, the tarsi
five-jointed, with small terminal claws and a broad empodium.
Wings very broad, with broadly rounded apices. Costal
space excessively widened from base to apex without any dif-
ferentiated pterostigma; the costal veinlets numerous and fre-
quently branched, generally connected by a more or less complete
series of longitudinal cross-veins (absent or reduced in the smaller
species); a recurrent vein at the base of the costal space. 8c, R,
and Rs are strong veins, and run closely parallel, forming a strong-
midrib to the wing, the cena tripHca, supported by numerous
cross-veins; this formation ends at a point between one-quarter
BY R. J. TILLYARD. 763
and one-third of the length of the wing from the apex, where So,
R, and Rs become joined by strong cross- veins, forming the
anastomosis; from this point onwards, these three veins continue
only as weak branching veins, indistinguishable from those
around them. Branches of Rs very numerous, closely parallel
to one another, their direction almost parallel with that of the
dorsal border of the wing. Central area of wing marked off as a
disc, bounded anteriorly by Rs, posteriorly by Cu, and distally
by a series of gradate cross-veins, the discal series. Other series
of gradate cross-veins may also be developed; in particular, an
internal series crossing the disc near its middle, and a terminal
series, lying between the discal and the termen, and continuing
the line of the costal cross- veins; apart from these, there is little
or no development of cross-veins upon the wing. M variable,
but always with its branches closely parallel, and occupying only
a little of the wing-space. Cu either two-branched or simple,
with or without a variable amount of anastomosis with M. Anal
veins short, three in number, mostly much branched.
Abdomen short, moderately stout, completely covered by
the wings in the position of rest. Ap])e7idages variable, usually
either forcipate or valve-like in the males.
Eggs oval, with rounded ends; laid separately and attached
by one side.
Larva very distinct on account of its large, squarish head,
from which the enormous jaws project like a pair of calipers;
mandibles without internal teeth; the body somewhat flattened,
of moderate width, fairly long, tapering posteriorly to the anal
sucker ; the legs short and formed for running backwards.
Colouration greyish-brown, with pruinescence. Inhabits crevices
in the bark of large forest-trees; fiercely carnivorous.
Cocoon spherical, or nearly so, resembling a large pearl;
spun from the anus, of fine cream-coloured silk, in crevices of the
bark.
Pupa short and stout, of normal Planipennian form. Active
at metamorphosis, when it cuts open the cocoon and ascends the
tree for some distance before disclosing the imago.
764 STUDIES IN AUSTRALIAN NEUROPTERA, vi.,
(A full account of the egg, larva, and pupa will be given in
dealing with the Life-History of Psychojwis elegans Guer., which
will form the next of this series of Studies).
Distribution: Australia (but not Tasmania), South Africa,
Burma, Thibet, and China; suggesting a Gondwanaland origin
for the family Close relatives of the family were present in the
Trias of Ipswich, Q.
The Fsychopsidft" are at once distinguished from all other Plani-
pennia by their shortened antennae, the joints of w^hich remain
distinct and similar, and by their remarkable wing-structure, in
which the most striking characters are the great enlargement of
the costal area, with formation of the connecting costal cross-
veins; the strengthening of Sc, R, and Rs to form the i'>ena
triplica, supported by cross-veins between them; the strong dif-
ferentiation betw^een the vena triplica and the parts of Sc, R, and
Rs lying beyond the anastomosis; the formation of the anasto-
mosis by interpolation of two small cross-veins; and the great
abundance and closeness of the branches of Rs.
As regards the definitions of the four Australian genera, I give
herewith Navas' key to them (13, p. 188), translated from the
original Spanish: —
'Forewing with four transverse series of gradate veinlets, three
in the disc and one the external continuation of the costal
series; hindwing with only three such series, two discal and
the external one which continues that of the costal field par-
allel to the external margin of the wing. ... 1. Psychopsis Newman.
Forewing with only three series of transverse veinlets 2.
'Both wings with three series of gradate veinlets, two in the disc
between the radial sector and procubitus ( = M), and the ex-
ternal one which is continuous in a curve with that of the
costal field 2. Arteriopteryx Gu6r.
.Hindwing with two or one series of gradate veinlets 3.
I ''Three transverse series of gradate veinlets, very distinct and
complete, in forewing; only two in hindwing... 3. Wernzia Navas.
^.i; - Disc of forewing with two complete series of gradate veinlets,
at times a trace of a third in continuation of the costal series;
\ hindwing with only one 4. Maoallanes Nav^as.
According to Navas, the arrangement of the gradate series
(1)
(2)
BY R. J. TILLYARD. 765
shown ill Text fig. 1, determines the insect as belonging to tlie
genus Arteriopteryx Guer. If, however, the forewing has an
extra series interpolated in the disc, between the internal and
discal series, then the insect belongs to Ptii/chopsis Newman. If
the arrangement for the forewing remains as in Text-fig. 1, but
the hind wing has one series missing, then the insect goes into
Wernzia Navas. If, in addition to this, the terminal series in
the forewing is absent, or only partially formed, then the insect
belongs to Magcdlanes Navas.
A complete study of the whole of the cross-veins in the speci-
mens in my collection has been carefully carried out. The total
number of specimens studied was sixty-four; but many of these
have since been given away or exchanged. They represent eight
species, two of which are new. The following very interesting
and instructive results are here given : —
Pt^ychopsis illidyei Froggatt. — The male in my collection has
three complete series in all four wings. In the forewings, there
are numerous scattered cross-veins in the upper distal portion of
the disc (Plate Ixxvii., fig. 4), but no sign whatever of a fourth
series between </i and g^. Hence, according to Navas, this
insect belongs to Arteriopteryx Guer.
In a large female of the same species, which I examined in the
Queensland Museum, and of which photographs are given in
Plate Ixxvi., figs. 1-3, there are only three series in the forewings,
and no irregularly placed cross-veins elsewhere in the disc. This
specimen, then, also- goes into Arteriopteryx^ according to Navas.
Of two specimens in Mr. Froggatt's Collection, one has three
complete series of gradate cross-veins in all four wings. The
other has four complete series on the left forewing, and only three
on the right. Hence, according to Navas, the first specimen
belongs to Arteriopteryx^ the second partly to that genus and
partly to Psychopsisl
Psychopsis mimica Newman. — I have examined seven speci-
mens altogether. The forewing shows also three complete series;
in addition, there are a variable number of cross-veins in the
60
766 STUDIES IN AUSTRALIAN NEUROPTERA, vi.
disc, between (/, and g.^, numbering from four to thirteen. In
only one specimen do these form a complete fourth series; in the
rest, they form either an incomplete series, or are just scattered
irregularly. In the hindwing, the terminal and discal series are
complete, but the internal series is very variable, being some-
times complete, and sometimes very incomplete, with as few as
seven veins in it.
This is the type-species of the genus Psychopsis. Of the seven
specimens examined by me, Navas would place only one definitely
within Psi/chopsis; the rest are either intermediate between
Psychopsis and Arteriopteryx, or between Psychoj^sis and Wernzia.
Psychopsis gracilis, n.sp., (Plate Ixxviii., fig. 10). — -The only
known specimen shows the three series of the forevving complete,
together with a very irregular, incomplete fourth series in the
disc between ^j and (/._,. In the hindwing, the discal and terminal
series are practically complete, but tlie internal series consists of
only four cross- veins, not forming a continuous series, and not
close together in line. One would like to know where Navas
would place this specimen.
Psychopsis elegans (Guer.), { = Ps. newmani Froggatt). —
Twenty-eight specimens of this species have been examined, most
of them bred specimens. The cross-venation is very variable.
In the forewing, the three series are always complete; but there
may or may not be a fourth. In some specimens, this fourth
series is complete, consisting of as many as thirteen veins in line;
in others, there is only half a series; in others again, only three
or four irregular veins; while, in the majority of specimens, there
is only one intermediate cross-vein (as in Text-fig. 1), or none at
all. I have seen one specimen having four complete series in one
wing, and only three in the other I In the hindwing, the ter-
minal and discal series are always complete, but the internal
series is very variable, being sometimes a complete series of from
eight to ten veins in line, and sometimes merely from three to
five irregularly placed cross-veins. Thus, different specimens of
this species might almost be placed in three of Navas' genera,
and we are not surprised to find this author himself placing
RY R. J. TILLYARD. 767
pjpgan^ in A7'ferioptery:i\ and neumiani in Psychopsis, altliough
they are one and the same species ! I
Piii/chopt^i:^ ccelivagus (Walker). — Five specimens have been
examined. The forewing has the terminal and discal series com-
plete, the internal series eithei' complete or incomplete. In the
hind wing, the internal series is absent, the terminal series is
complete, and the discal series is very variable. It may he com-
plete and in normal position, or it may be remo\'ed inwards to a
position midway between the normal positions of the discal and
internal series, or it may be entirely replaced by irregular cross-
veins; in one specimen, the disc is covered nearly all over by
ii'regular cross-veins, not forming any gradate series at all. This
xariability should be contrasted with Navas' definition of the
genus Wernzia, of which this species is the genotype.
Psyrhopsis inmlens Walker. — Twenty-one specimens have been
examined. In this species, the forewing has the discal series
complete, the internal either complete or nearly so. The terminal
series is usually quite absent; but, in some cases, there are from
one to five cross-veins belonging to the upper end of this series.
In the hindvving, the discal series is complete, the internal series
very variable. Normally either absent or represented by only
one cross- vein, it is nevertheless present in a fair percentage of
specimens, either as an incomplete series of two or three veins, or
as a complete series of four consecutive veins; further, right and
left wings do not always show the same condition. The terminal
series is absent in the hindwings; and the costal series, which is
complete in both wings of all the preceding species, is here
reduced to a shorter series, running from the base outwards for
a variable distance, from one-third to two-thirds the entire length
of the costal area. Thus we see that Navas definition of the
genus Mayalhtnes, of which this species is the genotype, will not
include the whole of the specimens examined.
Finally, in a single specimen of a new species (Plate Ixxviii.,
fig. 11), which will form the type of a new genus, we find the
discal and internal series present on both fore- and hindwings,
the discal series of the hindwings being not quite complete, and
768 STUDIES IN AUSTRALIAN NEUROPTERA, vi.,
removed somewhat inwards from the normal position. The mar-
ginal series is absent from both wings, and there is only a mere
vestige of the costal series, at the base of the wing.
(Psychopsis meyricki McLachlan, which I have not seen, is
stated by its describer to have both discal and internal series
present in the forewing; but nothing is said as to the condition
of the hindwing; the costal series is absent).
Reviewing the above evidence, it is clear that: —
(1) The genus Psycho^ysis Newman, being founded upon Ps.
mimica Newman, a species which itself shows great variability
in the number of its series of cross-veins, cannot be restricted by
any definition which attempts to limit this variability.
(2) The genus Artej'iopteryx Guer., founded upon P.s\ elegans
Guer., a species very closely allied to P><. mimica, and with almost
equally variable cross-venation, must be suppressed entirety, and
its type-species retained within Psychopsis Newman.
(3) The genus Wernzia, as defined by Navas, does not even
include all the specimens of its type-species. If, however, the
definition of the genus be widened, then it at once forms a part
of the Psychopsi^-sei-ies of species, and should be included in that
genus.
(4) The genus MagaUanes is founded upon a type-species which
grades into Wernzia in the characters of some of its individual
members. This should also, therefore, be removed back to
Psychopsis.
If these conclusions be accepted, the genus Psychop)sis will
include forms showing a gradual transition, from four series to
two in the forewing, and from three series to one in the hind,
with scarcely a break anywhere in the series. Bearing in mind
that the gradations are not specific constants, but essentially of
individual value only, it must be evident that any attempt to
subdivide this genus, upon the basis of these variable characters,
is bound to fail.
We have, therefore, to ask whether we should be content to
leave all the Australian species in the genus Psychoj^sis, or
whether we should seek for some new division.
BY K. .1. TILLYAKb. 769
I believe that a natural division t)f the Australian species is
possible, without attempting to make use of characters so essen-
tially variable as the cross-venation. Anyone who has studied
these insects at all fully will be at once struck with the very
great difference exhibited between Psychojyds illidyei Froggatt,
and all the other species. This may be seen at once by com-
paring Plate Ixxvii., fig.i, with Text-fig. 1 and Plate Ixxviii., fig.
10. Apart from its much greater size, Ps. illidgei differs from
all the other species by the shape of the wings, in which (but
more especially in the forewing), the dorsal margin is strongly
exca\ated, so that the torn us projects very prominently beyond
the rest of the wing-area. This species is also unique in possess-
ing raised or embossed areas upon the forewing. Further, there
is a gj-eat difference in the venation, in that the branches of the
radial sector are far more numerous and closely set in this species
than in any other, giving the wings the appearance of having
been made from spun silk or some other fine material. Finally,
if we examine the condition of the median vein, we shall find
that it has from four to six branches within the disc; whereas,
in all the other species, it is only two-branched. These differ-
ences are so striking, that I have no hesitation in separating out
Fd. iUirhjei from the other species of the genus, to form the type
of a new genus Megajysychops, whose full definition will be found
below.
All the Australian species left over in the genus Psychopsis
agree in possessing a moderate number of branches of the radial
sector, in having a hindwing about two-thirds as wide as the
fore, and with a very characteristic, dark, round spot upon it,
towards the apex. But I have, in my collection, a small i\e,\w
species, in which the hindwing is much narrower, being only
about half as wide as the forewing, and the dark spot is absent.
Though these characters do not separate this species so distinctly
from the rest as do those of Ps. illidyei, yet they have the merit
of being obviously constants. Hence I propose to describe this
new species under a new generic name, P sychopsella, of which it
will form the type. The full definition of this genus is given
below.
770 STUOIKS IN AUSTRALIAX NEUROPTERA, vi.,
There now remain within the genus Psi/chopsis the six Austra-
lian species Dihuica Newman, (jracilis, n.sp., eleyans Guer.,
cwlwof/iis Walker, insohns Walkei', and meyricki McLach. Of
these, the first three are distinguished by their usually larger
size, the somewliat angulated tornus of the wings, and the pink
or red colouration of the markings above the tornus of the fore-
wing. Contrasted with these, tlie other three are usually of
smaller size, with more rounded wings, the tornus not prominent,
and the colouration either fawn, brown, or black. It would
appear at first sight that these two groups suggest a natural
division of the genus into two. But, unfortunately, tlie species
elegans Guer., is so variable that it forms a connecting link
between the two groups. I have a number of specimens that
are no larger than well-sized examples of y*-. insolens', moreover,
the smaller the specimen, the more rounded does the wing
become; so that my series of elegans offers almost a complete
transition from the angulated tornus, seen in mhnica, to the
rounded toriuis of iusoleus. There remains, therefore, nothing
but the colour to go upon. The only conclusion I can legiti-
mately come to is, that these six species form a natural assem-
blage, which is best expressed as a single genus. But, having
said that, it should also be remarked that all six species are very
distinct forms, and there is no difficulty whatever in separating
them at sight. A key to these species will be found below.
Key to the x\ u s t r a 1 i a n Genera.
Ocelli vestigial. Fore and hindwings of almost equal width,
with the branches of Rs excessively numerous (27 to 32
in forewing, 18 to 24 in hind), exceedinglj' close together, so
as to give tlie wing the appearance of the close texture of spun
silk; dorsal margin strongly excavated before the tornus, the
latter very prominent, especially in the forewing. M at least
four-branched. Forewings with raised or embossed areas,...
ME<;ArsYCHOFS, n.g. (Type, Psychopsis illidgei Yvogga,tt).
Ocelli absent. Hindwing distinctly narrower than fore.
Branches of Rs moderately numerous (from 7 to 16), not ex-
cessively close together; Al only two-branched; dorsal margin
straight or only slightly excavated; no i-aised or embossed
areas on forewing 2.
(1]
(2)
BY R. J. TtLLYAKb. 7*71
Hind wing about two-thirds as broad as fore wing; and with a
distinct dark rounded spot towards apex or ternien
PsYCHOPSis Newman.
Hindwing only just over one-half as wide as forewing, and
without any such spot
PsYCHOPSELLA, n.g. (Type, Pi^i/chojjsef/a gallardi, n.g. et sp.).
Me G APS Y CHOPS, n.g. (Plates Ixxvi., Ixxvii.).
Characters as given above. Large insects, expanse over
50 mm.
G e n o t y p e, Fsychopsis illidgei Froggatt.
Megapsychops illidgei (Froggatt). (Plates Ixxvi., Ixxvii.).
Psychoftsis iUidyei Froggatt, These Proceedings, 1903, xxviii.,
p. 455, PI. xxi., figs. 7, 9. Kavas, Congreso de Yalladolid, 1916,
p.191, fig.2.
There is no need to add very much to the excellent description
given by Froggatt. The insect is so distinct and remarkable,
that it cannot fail to be recognised at sight.
Hah. — Mount Tambourine, Soutli Queensland ; 1,900 feet.
Exceedingly rare, only five specimens having l)een taken during
twenty years, all attracted to light.
Plate Ixxvi., fig.2, is a very fine photograph of the magnificent
specimen in the galleries of the Queensland Museum. This
specimen is an exceptionally large female. Plate Ixxvi., figs.l, 3,
show the venation of the fore- and hindwings of the same speci-
men.
Plate Ixxvii., fig. 4, shows the venation of the small male in
my collection. This is apparently the only known male of this
species. A short description of the points in which it differs
from the female is here given: —
Total letugth, \3'b mni^; forefcitig, 26 mm.; hindwitig, 22 mm.;
expanse, 51 '5 mm. Colouration not quite as dark as in female;
the spots near tornus of hindwings pale and semi-transparent.
A b d o m e n very short, only 7 '5 mm. long, narrowed at base,
but much swollen anally. A2:)pendages consisting of a pair of
superior and a pair of inferior, broadly triangular, valve-like
772 STUDIES IN AUSTRALIAN NEUROPTERA, vi.,
lamina?, forniiiig a pyramid, which ahiiost completely hides the
end of the abdomen from view (Plate Ixxa ii., fig.5).
Type, in Coll. Froggatt.
As the extinction of this fine insect would seem to be only a
matter of a few years, owing to the gradual clearing of the
mountain which is its only known home, the following account
of its habits and resting position, taken from notes supplied by
Miss M. Davidson, of Cedar Cottage, Mount Tambourine, will
be of considerable interest to all entomologists. The figures
given on Plate Ixxvii., figs. 6-7, were drawn from pencil-sketches
of the living insect made by Miss Davidson.
At rest, the position of the wings depends upon the kind of
surface on which the insect is placed. When, as usual, it rests
upon the bark of a tree, or some fairly wide and flat surface, the
wings are held roof-wise over the body, but inclined wide apart,
at an obtuse angle, as is shown in Plate Ixxvii., fig. 6. In this
position, the resemblance to a Reptilian head is very striking, as
is also said to be the case with the gigantic Hepialid, Leto staceyi
Scott. Both insects have the embossed area in much the same
position, resembling a pair of wide-open eyes. But, in Meyapsy-
chops, there is the additional suggestion of the forwardly project-
ing snout, formed by the coalescence of two dorsal bands I'ight
up to the head.
The local name for this insect on Mount Tambourine is, as I
am informed by Mr. W. H. Davidson, the "Cross-bones." This
would appear to have been suggested by the resemblance of the
transverse band on the forewing, with its somewhat narrow
stalk, terminated at either end by an enlarged embossment, to
the crossed femoral bones in the traditional pirate's emblem of
the "skull and cross-bones."
When resting on a tw^ig, the w ings are brought closer together,
so as to form a steep roof o\er the body, as may be seen in Plate
Ixxvii., fig.7. The head is almost completely retracted, and the
antennae folded beneath it out of sight. At night-time, when
the insect becomes active, or when feeding, the head is protruded
and raised up, w^hile the antennae are held almost vertically
upwards.
BY n. .t. TiLLYAtlt). / io
111 either position of rest, Miss Davidson states that tlie re-
semblance of the insect to a moderate-sized fungus is very marked.
Certainly, the white colour of the wings, with their yellowish-
in-own markiny:s, would sujxirest this; but I do not know whether
there is actually any fungus growing on the mountain for which
the insect might be readily mistaken. Unless there is, we can-
not say that the appearance of the insect is such as to aftbrd it
readily any protection from birds. From the fact that its wings
have been seen several times lying upon the ground, the insect
having evidently been seized and eaten by a bird, and also that
Mr. Davidson himself lost a fine female from his verandah,
through a bird seizing it, there would seem to be prwia facie
evidence that its appearance affords it no protection whatever.
Birds may, therefore, have been one of the main factors in the
reduction of the Psychojysidm to a mere remnant since Jurassic
times, and also in the total extinction of their Jurassic allies, the
Proheynerobiidai and the gigantic KaWgrammatidce.
Miss Davidson's sketches were made from the specimen in my
collection, which was captured on Dec. 7th, 1916, and was kept
alive for about a fortnight on sugar and water, in the hope that
it might lay some eggs. Unfortunately, it proved to be a male.
A fine female, which was apparently attracted to it one night,
was seen in the early morning resting upon the verandah; but,
as already recorded, was seized and carried off by a bird, before
Mr. Davidson could secure it.
PsYCHOPSis Newman.
Newman, Entomologist, 1842, p. 4 15. Hagen, 8tett. ent. Zeit.
1867, p.375.
Arteriopteryx Gueriii, Iconographie du Regiie Animal, 1845^
iii., p.389.
Arttrr'wptery.K (partim) Navas, Coiigr. Valladolid, 1916, p. 189.
Zyyophlebias Navas, Broteria Braga, 1910, p. 82.
Wernzia (partim) Navas, Mem. R. Acad. Ci. Art., Barcelona,
1912, X., p.l95.
Maycdlanes (partim), Navas, Mem. R. Acad. Ci. Art., Barce-
lona, 1912, X., p.l97.
774 STUDIES IN AUSTRALIAN NEUROPTEKA, vi.,
Characters as given in the key on p. 771, with the following
additions: — Moderate-sized insects, expanse varying from 25 to
50 mm. ; with or without anastomosis between M and Cu in
fore wing.
Genotype, Psychopsis mimica Newman.
With rejxard to the variable character of the amount of anasto-
mosis between M and Cu in forewing, Text-fig. 4 shows two of
the variable conditions to be found in the genus. In the type-
species, Ps. mimica, the most frequent condition is that shown
in Text-fig. 4rt; but a condition resembling that shown in h may
also occur in one or both wings. In Ps. elegans, the commonest
condition is that shown in h, but there are many irregularities
in one or both wings. In Ps coelivarjiis, either of the two con-
ditions shown in a and b may be present, or the anastomosis may
be complicated by the presence of numerous cross-veins, or there
may be no anastomosis at all; this last condition is the most
usual one for Ps. insolens.
Not having seen any of the African species, I cannot say
whether the type of anastomosis on which Navas bases his separa-
tion of these species, as a tribe separate from the Australian ones,
is of the type here shown or not: but, as he qualifies his statement
that M, Cu and its sector (by which, I presume, he means Cu,.)
are anastomosed, by the phrase "magis vel minus sensibili'" (13,
p. 199), I anticipate that here, as almost everywhere else, he has
failed to select a constant character for his subdivision, and that
a thorough revision of the African species will result in the com
plete collapse of his tribal distinctions.
Key to the ►Specie s o f P s y c h o p s i s .
Larger species, expanse 83 to oUnini., with detiiiite, transverse
fasciie in forewing, and pink or red markings near toriuis;
forewing with more or less prominent tornus. Dark spot on
hind wing placed well helotr the anastomosis of 8c, R, and Rs. 2.
Smaller species, expanse 25 to 36 mm., with tesselated, fawn-
coloured, grey or black markings on forewing; tornus well
rounded. Dark spot of hindwing placed either upon the
anastomosis itself, oi- between it a'ld the apex 4.
BY R. J. TILLYARD. 775
TFasciie of forewing very distinct; no blood-red patch situated
about one-fourth of the wing length from the base 3.
(2) -11- no
^ ' ' T^Vscia? less distinct; a blood-red patch on forewing, at about
1 Fascij
1^ one
(3)
(4)
one-fourth of the wing-length from base Ps. elKjaui^ (Guer. ).
^Two very distinct and well separated black spots, of moderate
size, situated near tornusof forewing; at least five transverse
fascia?, of which the tirst ends in a dark blotch overlying Sc,
R, and Rs at about one-fourth of the wing-length fioni tlie
base, while the last three converge towards the tornus
Ps. mrm/ca Newman.
Two very minute black spots close together at tornus of fore-
wing; only four transverse fasciw, of which the first com-
pletely crosses the wing to the dorsum, while the third and
fourth converge to meet above the tornus Ps. (/racifis, n.sp.
iSpot of hind wing covering the anastomosis 5.
8pot of hindwing placed about half-way between anastomosis
and apex; colouration of forewing yellowish-brown or fawn-
colour Ps. t nsolens Mohach.
TForewing heavily marked with black; spot of hindwing black...
_ j Ps. Cii>lira<ins { Walker) ,
I Forewing tesselated with pale grey; spot of hindwing pale
I fuscous Ps. wjcyr/'f^-/ McLach.
PsYCHOPSis MiMiCA Newmaii.
Newman, Entomologist, 1842, p. 4 la.
Froggatt, These Proceedings, 1903, xxviii., p. 454, PI. xxi., figs.
3-0 (nee Psychopsis inimica Froggatt, These Proceedings, 1902,
xxvii., p 367).
Navas, Congr. Valladolid, 1915, p. 190, fig.l.
Ilemerobins mimicns Walker, Cat. Neuroptera Brit. Museum,
1853, p. 27 9.
Type, in British Museum.
Tliis is tlie most widely distributed of all the known species,
but by no means the commonest. I have seen specimens from
several localities in the south-west of Western Australia, fronj
South Australia (including several taken at light in Adelaide),
from Victoria, and from the western parts of New South Wales.
Along the coastal strip of New South Wales, it is replaced by
Ps. eleyaiis; but it reaches the coast again around Brisbane, Q.,
776 STUDIES IN AUSTRALIAN NEUROPTERA, VI.,
where a number have been taken at electric lights. I know of
no record furtlier north than this.
PsYCHOPSis GRACILIS, n.sp. (Plate Ixxviii., fig. 10).
(J Total length, 11 mm.; abdomen, 5-5mm.; /oi'ewhig, 20 5mni :
hiadwing, 17*5 mm.: expanse, 42-5 mm.
Head pale brownish in colour, 1*8 nnn. wide, eyes black,
antenme pale ochreous, 4 mm. long.
T h o r a X : prothorax pale ochreous, 06 mm. long by 1*2 unn.
wide. Pterothorax pale ochreous anteriorly, shading to dark
greyish-brown behind, irregularly marked with brown blotches:
greatest width 2 mm. Legs very short, very pale ochreous in
colour, slightly darker on tibiae and tarsi.
Wings with a general resemblance to those of Ps miinica,
but much more transparent, more delicately built, and less hairy.
If placed upon a dark background, the wings of Ps. mhnica
appear white, with their pattern of dark fasciae and other mark-
ings showing up very distinctly; those of Ps. gracilis, on the
other hand, are very transparent, so that neither tlie venation
nor the colour-pattern shows up well under these conditions.
Forewings more triangular in shape than those of Ps. miinica,
more narrowed towards the base, and slightly wider at the level
of the tornus. Only four distinct transverse fasciae, of which the
first two are only lightly indicated in the costal area (those of
Ps. mimica are most strongly marked in this area); the most
basal fascia runs as a slight arc from Sc to the dorsum, at about
one-fourth of the wing-length from the base; the second is incom-
plete, and passes from Sc transversely only half-way across the
disc, a little basad of the middle of the wing; the third is well
marked on the costal area, slightly broken as it approaches Sc,
then well marked as it passes downwards, somewhat obliquely,
to meet the fourth above the tornus; the fourth fascia is also
well marked in the costal area, and passes just outside the
anastomosis, transversely downwards, to meet the third at an
angle of about 50°. On this fourth fascia, just before it meets
the third, are two distinct blackish spots; below the union of the
fascia, and extending to the tornus, is a broad pinkish band,
BY H. .1. TILLYAKD. / / t
carrying two small black spots at the tornus itself. All the
above fasciie are formed, as in Ps. mimica, of two more or less
parallel dark lines placed fairly close together, and enclosing a
paler area. The only other markings are an irregular brownish
area close to the base, between R and Cu, a series of irregular
markings along the dorsum, and indications of an extra fascia
lying below the apex of the wing, parallel and close to the ternien
(as also in Ps. mimica). Hind wing quite differently shaped
from that of Ps. mimica, the base being much narrower, the
terminal border cut off straight, and the tornus approaching a
right angle; hind border or dorsum slightly excavated. Dark
spot somewhat larger than is usual in Ps, mimica, less definite in
shape, and brownish instead of black.
Abdomen pale brownish, with darker markings, somewhat
narrowed basally (apparently much shrunken), enlarged anally.
Appendages (Plate Ixxvii., fig.8) consisting of a pair of superior
and a pair of inferior, triangular, valve-like laminae, partly em-
bracing the terminal segment, but with their apices not meeting
together in the middle line.
Type, (J, in Coll. Tillyard.
i^a6. — Booyong, N.S.W , Nov. 1904 (S. W. Jackson). There
is also a very damaged specimen in the Queensland Museum,
Brisbane, from Mount I'ambourine.
This very delicate and graceful species is easily distinguished
from all the known species except Ps. m-imica; the differences
which separate it from this latter species have been emphasised
in the key and in the description given above.
PsYCHOPSis ELEGANS ((iuerin). (Text-figs. 1-3).
Arteriopteryx eleyans Guerin, Iconographie du Regne Animal,
1845, iii., p 389. Navas, Congr. Yalladolid, 1916, p. 193.
Zyyo2>hlehins verrauxinus Navas, Broteria Braga, J 910, p. 84,
fig.23.
Psychopsis Tuimica Froggatt, These Proceedings, 1902, xxvii.,
p.367.
Psychopsis netvmani Froggatt, These Proceedings, 1903, p. 454,
PI. xxi., tigs. 1-2. Navas, Congr. Valladolid, 1915, p. 191,
778 STUDIES IN AUSTRALIAN NEUROPTERA, vi.,
Type, in Paris Museum.
Hah — Eastern coast of Australia, from Sydney to North
Queensland (Cairns, Kuranda, Herberton).
This species is occasionally taken at light, or beaten from
bushes, but was considered quite a rarity until Mr. Luke Gallard,
of Epping, N.S.W., discovered the larva. Since then, he has suc-
ceeded in bleeding a number from larvie found around Epping,
and I have bred out nearly two dozen from larviB taken in various
localities, including Hornsby and Berowra, in the Sydney dis-
trict, Wauchope on the North Coast of New South Wales, Bris-
bane, Stradbroke Is., Caloundra, Caboolture, Landsborough and
Mount Tambourine, in Queensland Mr. F. P. Dodd has taken
the insect at Kuraiida and Herberton, N. Queensland.
From larvae taken on One-tree Hill, Brisbane, under the bark
of an Ironbark tree (Eucalyptus leucoxylon F.v.M.), I bred both
Ps. eleyans and Ps. cfelivagus. There is so little difference
between the two larvae, except in size, that I was not able to
distinguish which was which. This seems to me to be a further
argument against removing Ps. cctlivagus from the genus
Psychopsis.
PsYCHOPSis ccELiVAGus (Walker).
Hemerobius aelivayus Walker, Cat. Neuroptera Brit. Mus.,
1853, p.279.
Psychopsis ccelivag\is Froggatt, These Proceedings, 1903, xxviii.,
p. 455, PI. xxi., fig. 8 (nee Froggatt, /.c, 1902, xxvii., p. 368).
Wernzia coelivaya Navas, Mem. R. Acad. Ci. Art , Barcelona,
1912, X., p.l95. Navas, Congr. Valladolid, 1916, p.l94.
Type, in British Museum.
JJab. — Queensland coast, from Brisbane to Kuranda and Her-
berton. A rare species.
This exceedingly striking species is at once distinguished by
the remarkable metallic black markings of the forewings, as well
as by the pure whiteness of the venation, which gives it a most
striking appearance. Some years ago, it was not uncommon on
One-tree Hill, Brisbane: but recent severe bush-fires seem almost
to have extei^minated it.
BY R. J. TILLYARD. 779
PsYCHOPSis INSOLENS McLachlan.
McLachlan, Journ. Entomology, 1866, ii., p. 114.
Pst/chopsis aelu-aga Froggatt, These Proceedings, 1902, xxvii ,
p.367.
Fsychopsis riisolens Froggatt, I.e., 1902, xxvii., p.367; Froggatt,
I.e., 1903, xxvii., p.454, f.6.
Mayallanes insolens Navas, Mem R. Acad. Ci. Art., Barcelona,
1912, X., p.l97. Navas, Congr. Valladolid, 1916, p.l96.
Type, in British Museum.
Hab. — New South Wales and Queensland; not uncommon
It occurs only sparingly in the Sydney district, but is commoner
on the tablelands, particularly on the Queensland border. I
have good series from Killarney, also from Mount Tambourine.
It has been taken as far north as Herberton (F. P. Dodd). Most
of the specimens appear to have been taken at light. Nothing
is known of its life-history.
This species is less beautiful than any of the others, but freshly
emerged specimens are of a very effective fawn colour, which
fades considerably with age.
PsYCHOPSis MEYRiCKi McLaclilan.
McLachlan, Ent. Mo. Mag., 1887, xxiv., p.30.
Magallanes meyrieki Navas, Congr. Valladolid, 1915, p. 197.
T y p e, in Coll. McLachlan.
^a6. — Jindabyne, N.S.W., 2,800 feet.
I have not seen this species, of which the unique series of seven
specimens taken by Mr. Meyrick on Jan. 20th, 1885, is in the
McLachlan Collection. McLachlan gave the habitat as "Kos-
ciusko, 2,800 feet,'' but this is evidently an error. Mr. Meyrick
informs me {in litt.) that the specimens were actually taken near
Mr. Body's Station at Jindabyne. This latter township is on
the main route to Mount Kosciusko, but more than thirty miles
from its summit; the Kosciusko Range does not begin to rise
from the Monaro Plateau until the junction of the Snowy and
Thredbo Rivers is reached, and this is some miles beyond Jinda-
byne. It seems desirable to point this out, since the type of
country in which the insect was taken dififers very greatly from
that found at higher elevations on the Kosciusko Range.
780 STUDIES IN AUSTRALIAN NEUROPTERA, vi.,
PsYCHOPSELLA, n.g. (Plate Ixxviii., fig.l 1 ).
Characters as given in the key on p. 771. Small insects, with
an expanse of about an inch; the hind wings very much shorter
than the fore.
Genotype, PsychopseUa gallardi, n.sp.
PsYCHOPSELLA GALLARDI, n.sp. (Plate Ixxviii., fig. 1 1 ).
^. Unique. Total length, 7*8 mm.; abdomen, 5mm.; forewivci,
12"5 mm.; hindwing, 95 mm.; expanse, 245 mm.
Head 1-3 mm. wide, pale ochreous: eyes dark grey; antennae
dull brownish, very short, 18 mm.
Thorax: prothorax pale ochreous. Pterothorax dull brown-
ish, with darker markings. Legs very short, with pale ochreous
femora, slightly darker tibise and tarsi.
Wings: foreiving with discal and internal gradate series
complete, the former consisting of fifteen cross-veins, the latter
of nine. Costal series of cross- veins rudimentary, consisting of
only about six cross-veins near base, and a single one about half-
way along the costal area; marginal series absent. Colouration
a very pale ochreous, tesselated all over with slightly darker
markings, and here and there small dark brown patches; of
these, the most prominent are three sets along the vena triplica,
another just beyond the anastomosis, one near the middle of the
discal series, two in the marginal area, and some irregular dark
markings along Cu. Hindivhig with short discal and internal
gradate series, not complete, the former of six or seven cross-
veins, the latter of only four; no marginal series, and only one
or two costal cross- veins near base. iVo markings at all upoot
the hindtving, which is dull whitish and quite transparent.
Abdomen pale ochreous, prettily marked with a mid-dorsal
series of dark spots and smaller lateral spots on each segment.
Appendages', superior, a pair of very conspicuous, depressed, sub-
triangular processes, laterally flattened, and appearing somewhat
forcipate from above; these project well beyond the tip of the
abdomen; each one has its apex rounded, and carries near its
base, on the outer side, a raised embossment; the whole of the
appendage above the embossment, and outwards to the tip,
BY R. J. TILLYARD. 781
carries long, ciu-ved, wliitisli hairs. Veiitrally, these appendages
are separated by a broad, sliort, tiapezoidal inferior apj^endaye,
wliitish in colour (Plate Ixxvii., fig. 9;.
Type, in Coll. Tiliyard.
Ilah. — ^The specimen carries no label; but Mr. Gallard, from
whom I obtained it, informs me that he bred it from a larva
found near Gosford, N.S.W.
At first sight, this species appears to resemble a small speci-
men of Ps. insolens. It may readily be separated from the latter
species by the much more delicate build, the much shorter and
narrower hindwings, with any markings on them, and the more
delicate, paler, narrower, and less strongly marked forewings.
Relationships of the P s y c h o ]> s i d fH .
There can be little doubt that the nearest relatives of the
Psychopsidce are to be found in the Mesozoic Fossil families Pro-
hemerobiidm and Kalliyraminatidce. The formei', which verv
probable represent the original type of the Order Planipennia,
are known from the Upper Trias of Ipswich, Queensland, and
from the Lias and Upper Jurassic of Europe. The genus Proto-
psychopsia Tiliyard, appears to connect the Prohnmerobiidcp. with
the Psychojfsidce very closely. This relationship has since been
further emphasised by the discovery of a second fossil from these
beds, differing very little from Megapsychops illidyei itself. This
fossil will shortly be described by me in Part 5 of my "Mesozoic
Insects of Queensland." The differences between the Psychopsidce
and the Prohemerobiidce are simply those in which the formei-
family shows specialisation, viz., the enlargement of the costal
area, with development of a series of connecting cross-veins, the
formation of the vena triplica and anastomosis, and, very pro-
bably, the shortening of the antennae. In all known Prohemero-
biidce, the costal space is either quite narrow, or only moderately
wide, for its whole width, or only widened at the base; costal
veins are never developed: Sc, R, and Rs run close together, but
quite separate, to the apex of the wing, without being partially
strengthened and separated oft" to form a vena triplica, as in
Psychopsidce: cross-veins apj ear to be entirely absent from most
61
782 STUDIES IN AUSTRALIAN NKUROPTERA, vi.,
species, but may be very weakly indicated between Sc, R, and
Rs, and also in the position of the gradate series (these, though
shown in my figure of P^^otopsychopsis, are so faintly indicated,
that I have been doubtful how far they really existed). The
oldest forms of Prohemerohiidce were only of small to moderate
size, with only moderately numerous branches of Rs; specialisa-
tions in this family appear to have led to the development of
forms with an immense number of closely-set branches of Rs.
The Upper Jurassic forms are, on the whole, considerably larger
than the Liassic forms.
As regards the Kalligrammatidcp, these gigantic insects differ
from the Prohemerobiidce only in the development of a close
series of connecting cross- veins all over the disc; in the beginning
of an anastomosis, by connection of Sc with R distally by means
of a short cross-vein, very much as in Psychopsis; and in the
development of the large eye-spots on the wings, recalling those
of Alegapsychops illidgei. 'J'he rise and fall of this wonderful
family was probably very quick; for. if their larvae, as is pro-
bable, were at all like those of Psychopsin, they could scarcely
have attained to dominance before the rise of the Birds must
have quickly exterminated them.
Here let it be noted that Megapsychops illidgei shows, in the
possession of a media with more than two branches, a character
that was evidently possessed by the Kalligrainmatidce (the true
M of this family is labelled Cu by Handlirsch), and by certain
of the Liassic Prohemerohiidce, if not by all. Combining this
fact with another obvious one, viz., that the development of
numerous branches of Rs, occupying a very large area of the
wing, is a specialisation confined to the Planipennia, and must,
of necessity, have caused a squeezing out and consequent reduc-
tion of the media, we are bound to conclude that the ancestors
of the Planipennia approached more closely to the Panorpoid
type, as regards the structure of Rs and M, than they do to-day,
and that the many-branched media of Alegapsychops illidgei is an
archaic survival that stands to remind us of this fact.
I regard the Psychopaidm as a Mesozoic remnant, considerably
isolated, at the present day, from all other existing families. Ii^i
BY R. J. TILLYARD. 783
order to understand their position, it must be emphasised that
they have closest affinity witli those groups in which the cross-
venation is limited to gradate series, and Sc is not united with
R in a true anastomosis (i.«., by actual fusion of the two main
veins). Thus they show affinity, on the one hand, with the
Hemerobiidce (s.str.) and Dilaridce, but remain more primitive
than either of these, on account of the single radial sector of the
forewing; while, on the other hand, they are related to the Tri-
chomatidxfi^ Apochrysidce, and Chrysopidcf. The 7richoviatidce,
like the Psychopsidce, appear to be a direct offshoot of the old
Prohemerobiidcp; but their tendency has been towards reduction
and narrowing of the wings; whereas that of the Psychoj^sidm
has proceeded in the opposite direction. In the Apochrysidce,
we recognise a group of insects with certain cliaracters of the
wings strongly suggestive of Psychopsid affinities, viz., the widen-
ing of the costal area and the extreme differentiation of the disc.
Bearing in mind, however, the slender build and elongated
antennae, and their evident close relationship with the Chryso-
pidce, I am inclined to regard these venational characters as the
result of convergence, rather than of true relationship. Both
the ApochrysidcE and the Chrysojndce differ radically from the
Psychopsidce in the manner of development of the branches of
Rs, which tend to cross the wing transversely, and have thus
brought about new tendencies in the development of the gradate
series, and finally the unique formation of the pseudomedia and
pseudocubitus.
The larval form of Psychopsis is such that, from it, there can
be equally easily derived the small-jawed series of larva? found
in the Hemerobiidce, Chrysopidw, etc., or the long-jawed series of
the Bei'othidce and Osmylidce, or the large-jawed series of the
Nymphidce, Myrmeleoiitidce, etc. In the last series, there has
been a gradual addition of internal teeth to the mandible,
beginning with one only in Nymphidce, and passing up to three
or more in MyrmeleoididcB. Hence we shall probably be on safe
ground, if we regard the larval type found in Psychopsia as
closely similar to that of the extinct Prohemerobiidce.
From the Berothidct and Osmylidce, the Psychopsidce are at
784 STUDIES IN AUSTRALIAN NEUROPTERA, vi.,
once distinguished by the lack of a true anastomosis between Sc
and R. The Berothidcv. are clearly a specialisation from the
older Trichomatidce, the Osmylidce from a type resembling the
still existing PoJystcechofes, which, to my mind, will go well
enough into the BerotJiidcF as at present defined. It should be
noted that both palseontological and morphological evidence
tends to show that the groups with numerous cross-veins in the
disc are of more recent origin than those without; it is only by
stressing this point that we can be sure of obtaining a correct
view of the Phylogeny of the Planipennia, which are an essential
portion of the Panorpoid Complex, and originated from more
open- veined ancestors.
BIBLIOGRAPHY.
1. Banks, N., 1910. — " Synonymical Notes on Neuroptera." Ent. News,
Philadelphia, 1910, p. 389. {ZyyophJehia and Bahnes Navas con-
sidered to be synonyms of Psychopsis Newman).
2. , 1913. — "Synopses and descriptions of Exotic Neuroptera.""
Trans. Anier. Ent. Soc, 1913, xxxix., pp.201 -242. (Subfamily
Fsychopsinie established on account of the union of Sc, R, and Rs).
3. Froggatt, 1902. — " Notes on Australian Neuroptera and their Life-
Histories."" Proc. Linn. Soc. N, S. Wales, 1902, xxvii., p.358,
4. , 1903. — " Notes on the genus Psychopsis Newman, with
Descriptions of new Species.'" Ihid., 1903, xxviii., p. 453, PI. xxi.
5. (iUERiN, 1845. — Iconographie du Regne Animal, 1829-1844, iii., p. 389.
(Description of Arteriopteryx elegans).
6. Hagen, 1867. — " Hemerobidarum Synopsis synonymica " Stett. ent.
Zeit., 1867, p. 375. (Characters of the genus Psychopsis Newman).
7. McLachlan, 1866. — "On some new Species of Neuropterous Insects
from Australia and New Zealand, belonging to the Family Hemero-
biidce,'' Journ. Entom., 1866, ii., p.lll. (Description of Ps.
insohns).
8. , 1887, — Psychopjsis meyrichi, n.sp. Ent. Mo. Mag., 1887,
xxiv., p. 30.
9. , 1891. — "An Asiatic Psydiopsis" {Ps. /j//v?ia«a, n.sp.). Ent.
Mo. Mag., 1891, xxvii., p,320.
10. , 1902. — " A second African Species of Psychopsis" (Ps.
marshani :SlcLa.c\\.). Ent. Mo. Mag., 1902, xxxviii., p. 234.
BY R. J. TILLYARD. 785
11. Navas, 1910. — " Hemerobidos mievos, eon la clave de los tribus y
generos de la familia." Bioteria Braga, 1910, ix., pp. 69-90. (Char-
acters of genera Z>/;/ojjhfthiK'< and /ia//nt-^-, description of Z. rer-
reauxinns).
12. , 1912. — '*Insectos Neuropteros nuevo« o poco conocidos."'
Mem. R. Acad. Cienc. Art. Barcelona, J 912, x., No.9, pp. 135-202.
(P. 62, tribe P,^yrhopshu defined: p. 63, Wernzla; and p. 65, MaijaU-
ane-<, n.g., characterised).
13. . 1916(?) — " Essaj^o nionogratico de la familia de los .Sicopsidos
(Ins. Near.).'" Asoc. Espanola para el Prog, de las Ciencias : Go)i-
greso de Valladolid, Oct. 1915, pp. IS] -210.
14. Newman, 1842. — " Entomological Notes." Entomologist, 1842, pp.
413-415. (Genus P.^i/chopsi^: defined, and type-species, Ps. miniica,
described).
15. TiLLYARi), 1916. — " Studies in Australian Neuroptera," No. iv. Proc.
Linn. Soc. N. S. Wales, 1916, xli., pp. 269-332, PL xii.-xix. (See
pp.271 and 289).
16. , 1917. — "Mesozoic Insects of Queensland," No. 1. Ihid.,
1917, xlii., p. 178, PI. viii., fig.3. (Gemis Protopsf/chopsis defined,
and Ps. venosa described.
17. Walker, 1853. — Catalogue of the Neuroptera in the British Museum.
1853, p,279.
Xote. — This Bibliography^ is not intended to be a complete one, but onh-
to list the papers actually' mentioned in this work. The references to
Australian species are, however, complete.
EXPLANATION OF PLATES LXXVL-LXXVIII.
Plate Ixxvi.
Me(japsy chops i/lidgti (Froggatt).
Fig.l. — Venation of forewing of large ? in Queensland Museum; ( x about
2i).
Fig.2. — Large 9 in Queensland Museum; ( x about H).
Fig.3. — Venation of hindwing of same specimen; ( x al)Out 2j).
(Photomicrographs of figs. 1 and 3 taken by Mr. H. Hacker, Entomolo-
gist to the Queensland Museum, and sent by Mr. H. Longman, F.L.S.,
Director of the Queensland Museum. That of fig.2 sent by Dr. R. Handyn
Harris, late Director of the same Institution).
1S6 STUDIES IN AUSTRALIAN NEUROPTEKA, vi.
Plate Ixxvii.
Mtga-'^ychops iUidgei (Frot^gatt).
Fit;. 4. — Complete venation of both wings of small 6 in Coll. Till3'aicl;
(x4). (Compare with Plate Ixxvi., and conti'ast with Text.fig.I :
tn, tornus; venational notation as in Text-fig. 1).
Fig,5. — Appendages of the same 6 , postero-doisal view; ( x IH).
Fig.G — The same insect, alive, resting upon a flat surface. Dorsal view;
(;<U).
Fig.7. — The same insect, alive, resting upon a twig. Lateral view; ( x H).
Psychopsis gracilis, n.sp.
Fig. 8. —Appendages of 6 . Postero-dorsal view; ( x IH).
P)<ycho}jsdla (jallardi, n.g. et sp.
Fig.9. —Appendages of 6 . Postero-dorsal view; ( x 27).
(Figs. 6-7 drawn from pencil-sketches sent by Miss M. Davidson, of
Cedar Cottage, Mount Tambourine, Q. The dark colouration is a lich
brown, and resembles a fairly thick layer of A'arnish. In fig. 4, the dotted
double-arc indicates the external raised edge of the lai'ge embossment in
the forewing; the dotted oval represents the area covered by the dark
spot in the hindwing. In fig.7, the abdomen of the insect is slightly
visible through the semitransparent wings).
Plate Ixxviii.
Fig. 10. — Pi^ychopsis (jracilis, n.sjj., 6 , in Coll. Till3^ard; ( x 4).
Fig. 11. — Psychopsella ycdlardi, n.g. et sp. Unique J in Coll. Till^'ard;
(x6).
781
STUDIES IN AUSTRALIAN iVEUROPTERA.
No. 7. The Life-History of Psycuopsis elegans (Guerin).
By R. J. TiLLYARD, M.A., D.Sc, F.L.S., F.E.S., Linnean
Macleay Fellow of the Society ix Zoology.
(Plate Ixxix., and twelve Text-figures).
The only account of the life-history of any species of the family
Psyckopsid(e is the short, popular account of the life-history of
PsychopsU eleyans Guerin, given by Mr. Luke Gallard in the
"Australian Naturalist" of 1914.* From this we learn that
Mr. Gallard captured a female of Ps. elegans (he uses the syno-
nym newmani Froggatt, throughout, for this species) at Kent-
hurst, N.S.W., in 1904. This insect laid over fifty eggs, some
of which Mr. Gallard raised to about one- third of the full larval
size. Mr. Gallard informs me that it was one of these specimens,
newly hatched, which is figured on p. 62 of Mr. Froggatt's "Aus-
tralian Insects" as the larva of Pfiychopsis nii'niica Newman.
Mr. Gallard did not succeed in rearing the imago until 1911.
A larva taken in March, 1911, by Mr. J. Blake, of Narara, near
Gosford, N.S.W., was kept alive by Mr. Gallard until December
11th of the same year, when it spun a cocoon in the box, the
imago emerging on January 16th, 1912. Since then, Mr. Gallard
has discovered the larvae in many localities round Sydney, and
has bred a number of specimens.
In 1915, when I had begun the study of the Neuroptera
Planipennia, and was very anxious to study the Psychopsidce in
particular, Mr. Gallard very generously invited me to accompany
him in the field, and showed me the ingenious ways by which he
found these larvae. I shall never forget the skill and energy
that he displayed in this work. As the whole credit for the
discovery of this larva rests with him, and is simply due to his
* "Notes on Psychopsis newmani.'''' By Luke (lallard, Australian
Naturalist, iii., Part 3, 1914, pp.29-32.
788 STUDIES I\ AUSTRALIAX NEUROPTEUA, vii.,
persistence and keenness in following up clue after clue fof many
years, I should like to take this opportunity of congratulating
him upon th^ fine results of his work, and of thanking him very
heartily for tli '. help offered to me, without which I feel quite
certain that neither T nor anyone else could possibly have hit
upon the track of such a remarkable larval form as this.
Since Mr. Gallard first taught me how to find this larva, I
have discovered it in many localities around Sydney, also near
Wauchope on the North Coast of N.S.W., and in many places
in S. Queensland, including Brisbane (One-Tree Hill), Strad-
broke Island, Mount Tambourine, Caboolture, Caloundra, and
Landsborough. It probably occurs all along the Eastern Coast-
line of Xew South Wales and Queensland, wherever there are
suitable rough-barked, Myrtaceous trees for the larva to hide in.
In the present paper, 1 propose to give a full description of
the egg, larva, and pupa, together with an account of the habits
of the larva, the spinning of the cocoon, the emergence of the
imago, and some details about the latter that have not yet been
carefully investigated.
My thanks are due to my wife for the execution of Plate Ixxix.,
from the living larva and pupa, in collaboration with myself.
The Life-cycle of Psychopsis elegans (Guerin).
The complete life-cycle of this species occupies about two
years. The larva, like almost all others of this Order, has only
three instars, during each of which an enormous increase occurs
in the size of the body, the size of the head remaining constant
in the meanwhile. The following Table exhibits the duration of
each period : —
Period. !
Duration.
Eyg ... ■■■\
i About twelve day.s.
First far ml insfar ..
About ei^lit months, including hibeination. (Feb -
Sept.).
SeroiKl larra/ iiif<tar ...
From four to tive months, without hiljernation.
(Sept. -Feb. or March).
Third kirral instar ...
About nine months, including a second hibernation,
and a fortnight to three Aveeks within the
cocoon before pupation. ( March -Nov. ).
Pupa
About three weeks.
Imago
About two months. (Dec. -Feb.)
6Y R. J. TILLYARD. 789
The Egg. (Plate Ixxix., H-. 1 ; Text-fi^^l).
The eggs are laid separately, or only two or three together, at
considerable intervals of time; probably, in the natural state,
upon the bark of Myrtaceous trees, especially Eucalypts. In
captivity, most of the eggs were laid upon cottonwool. Thev
are not stalked, and are laid upon one side, which is attached to
some object by a slight secretion of gelatinous matter.
The egg itself is oval, about 1 mm. long by 04 5 mm. wide in
the middle, and is of a semi-opaque creamy
colour, tinged with pale green. At its anterior
end there is a distinct micropylar projection,
as shown in Text-fig. 1. The egg is quite
smooth, without any pattern or sculpture.
The eggs are usually laid in January or Feb-
ruary. At the end of about twelve days, thev
hatch. The young larvae, hitherto curled up ^ ' '
inside, crawl straight out of the broken shell, and at once make
for some small crack or crevice in the bark, where they may
escape the numerous enemies that would otherwise speedily
compass their destruction.
First Larval Instar. (Plate Ixxix., tigs. 2-8; Text-H^^2).
The newly-hatched larva is somewhat more than three times
as long as the egg in which it was confined, the measurement
being taken from the tip of the mandible to the anal papilla.
As with all Planipennia, such a comparatively large larva can
only be contained within tiie egg by considerable folding of
parts; the head being tucked down below the breast, and the
posterior half of the abdomen being again folded forwards under
the head.
At first, before the larva has taken any food, the head is very
large in comparison with the rest of the body. P)Ut, after one
or two large meals, the abdomen begins to be distended with
food, and the general appearance of the larva undergoes a great
alteration, as can be seen by compai-ing Plate Ixxix., tigs. 2 and 3.
This change takes place in every instar of all Neuropterous larvae
* Two eggs of Psydiopais dtyans vGuer. ); ( x 2(J).
790
STUDIES IN AtJSTRAMAN NEUROPTERA, vii.,
known to ine. The reasons for it are two; firstly, that there are
onl}^ three or four larval instars in this Order; and, secondly,
that no food at all is excreted, the waste-products being stored
in a special sac or cham-
ber of the alimentary
canal, which is not cast
out until the emergence
of the imago. Hence it
is clear that a large in-
crease in the size of the
abdomen is inevitable
during each larval in-
star, whereas the size of
the head can only be in-
creased at ecdysis.
The head of the young
larva is somewhat
flattened, trapezium-
shaped, and wider in
front than behind ;
length about 0'6mm.,
breadth in front about
the same. At the two
anterior angles, which
are somewhat rounded,
are the two groups of
simple eyes, occupying vv\- — - — 7
the area from which the
compound eyes of the . ^'"^ " ^'~"
pupa and imago are latf^r developed. There are five ocelli in
each group. The anterior border of the head projects in the
middle to form the prominent triangular labrum, on either side
of which lie the long, slender antennm, which are eight-jointed.
The mouth-parts are very extraordinary, consisting of a pair of
* Newly-hatched larva of Ps. elegant (Guer.); ( x 30): />, part of hind leg
of same, to show tarsal claws and enipodiuni, tlie latter in the form of an
elongated process terminating in a sucking-disc; ( x 87)-
a
BY R. J. TILLYARD. 791
enormous caliper-like mandibles, 0-7 mm. long, a similar but
slenderer pair of maxillae without palpi, and a suiall triangular
labium carrying a pair of four-jointed palpi. There are no teeth
on the mandibles.
The segments of the thorax are slightly narrower than the
head, and subequal, the prothorax being the longer, but some-
what the narrower. The legs are short, the femora being fairly
stout, the tibi?e shorter and slenderer, and the tarsi veiy short
and unjointed. Distally, the tibiae are armed with a pair of
strong spurs. The tarsi end in a pair of short, strong claws,
between which there projects a long, slender empodium, ending
in an enlarged suction-disc, and closely resembling the same
structure in the larva of Chrysopidce. The larva is able to crawl
forward slowly, but prefers to walk backwards, which it can do
with considerable speed, using its anal papilla as well as its legs.
The segments of the abdomen are nine in number, together
with a terminal anal papilla, which represents the reduced tenth
segment and the anal appendages. These segments are narrow,
and taper from before backwards. The whole of the abdomen,
as well as the head and thorax, is clothed with short, stiff hairs.
The spiracles in the newly-hatched larva are eighteen in num-
ber, there being a pair upon the prothorax, and also upon each
of the first eight abdominal segments. They are, however, very
difficult to make out, the spiracular openings being small and
devoid of armature.
When first hatched, the larva is semi-transparent, with very
little colour-pattern. Later it darkens to a greyish-brown, which
tends to become overlaid with a whitish pruinescence. These
changes become more marked in the following instars.
At the end of the first instar, the larva has about doubled its
length, and the segments of the thorax and abdomen have
increased greatly in width and length, so that the head now
appears very small in comparison.
As soon as the cold weather sets in, in May or June, the larva
ceases to feed, and remains motionless, hiding away in a crevice
of the bark, until the warmer weather begins in September. It
then becomes active again; but, after one or two good meals, it
792 STUDIES IN AUSTRALIAN NFUROPTEKA, vii.,
again rests for some days, and tlien undergoes its first ecdysis.
This appears to bn fatal to many of tlie larvye, cbietiy owing to
the great difficulty experienced in removing the head from the
very hard shell of the cuticle enclosing it.
Second Larval Instar. (Plate Ixxix., fig.4; Text-ti<5,3).
At the tirst ecdysis, there is a great increase in the size of the
head of the larva, which now becomes about 14 mm. long in an
average-sized larva. (There is considerable variation in the sizes
of individual specimens, and this is not rectified in the imagines,
which also vary greatly in size). The head becomes more de-
finitely trapezium-shaped than before, owing to the increased
definiteness of the four angles; its colour is a rich brown. An
extra joint is added to the antennse, which are now nine-jointed,
and to the labial palps, which become five-jointed. The man-
dibles and maxill?e are almost as long as the head, shaped as in
the first instar, but of stronger build. The ocelli become more
plainly marked, each set of five being placed upon an irregular
darkened area just behind the base of the antenna.
Text-fig. 3 shows the cast skin of the head of the larva at the
end of this instar. The armature of the head is very remarkable,
consisting of numerous raised papillae, from each of which a tiny
hooked hair projects: these are especially conspicuous upon the
sides of the head, and upon the projecting triangular labrum.
The figure shows very plainly the mid-dorsal and lateral splits
which take place in the cuticle at ecdysis.
As at hatching, so also at the beginning of the second instar,
the three thoracic segments are approximately equal in size, the
prothorax being somewhat longer and narrower than the other
two. After a meal, the meso- and metathorax swell up, like the
abdomen. But the prothorax can only swell up posteriorly, the
neciv-constriction remaining unchanged; so that this segment
soon becomes much narrower than the other two. The legs
remain small, and formed as in the first instar, with un jointed
tarsi.
The abdomen is at first fairly slender and tapering towards
the anus. As the larva feeds, it swells up rapidly, and assumes
BY R. J. TILLYARD.
793
the somewliat broad, flattened shape shown in Plate Ixxix., fig.4.
A slight pattern becomes noticeable upon the abdomen and
thorax, more markedly in some individuals than in others. This
is mainly due to the onset of pruinescence, which leaves the
Text-fig. 3.
Cast skin of head of larva of Ps. elegans (Guer.), at second eedysis; ( x 30).
original dull brownish colour of the body more definitely marked
mid-dorsally and in the sutures. A pair of darkish spots can be
made out on the meso- and metathorax, marking the positions of
the pinacula of these segments (see p. 800).
At the end of the second instar, the larva has increased to
794 STUDIES IN AUSTRALIAN NEUROPTERA, vii.,
about 8 mm. in length, and is very stout. As this instar lias
been passed entirely in the summer-months, with an abundance
of food and warmth, growth is comparatively rapid, and the
second ecdysis usually takes place during February or March of
the second year of larval life. This ecdysis does not appear to
be such a crisis in the life of the larva as was the first one: pos-
sibly because the larvae are more active in the warm weather,
and make greater efforts to free themselves from the hard cuticle
of the head, which is again the principal cause of any mortality
that occurs at this period.
Third Larval Instar. (Plate Ixxix., fig. 5; Text-figs.4-9),
As before, this second ecdysis results in a great increase of
the size of the head, which broadens considerably, becoming
squarish, as shown in Plate Ixxix., fig.5. The increase in length
is only a moderate amount, but in breadth it is more than 50
per cent, of the width at the end of the second instar. The
labrum broadens with the head, and loses its triangular shape,
as may be seen by comparing Text-figs. 3 and 6. The antennae
sometimes become ten-jointed, but I have only been able to
count nine in several specimens. The labial palps remain five-
jointed, and there is no change in the shape of mandibles or
maxillae.
Thorax, legs, and abdomen remain of the same shape as in the
second instar. The pattern varies greatly for different indi-
viduals, some being brown all over, others brown with grey
pruinescence, and others entirely grey, or almost white, and
strongly pruinescent all over. The head usually remains a rich
dark brown; but I have seen specimens with strong pruinescence
upon the head also.
During March and April of its second year of existence, the
larva feeds up rapidly, and many specimens become apparently
full-fed by the time winter sets in. None, however, attempt to
spin up, but remain dormant in crevices of the bark until the
warmer weather of the Spring returns. During this second
hibernation, great mortality occurs, many larvae being attacked
by some obscure fungoid disease, and others apparently dying of
BY R. J. TILLYARD.
795
cold. Out of thirty larvae brought from Wauchope, N.S.W., in
February, 1917, I succeeded in rearing only six to maturity.
Before describing the spinning of the cocoon, we may profit-
ably study, somewhat more minutely, certain of the lai-val struc-
tures, as they are to be seen at the end of the third instar.
The Mandibles (Te.Kt-fig.4, a, h) are a pair of huge curved jaws,
immensely strong, and of a
rich dark brown colour.
They are smooth in outline,
and without visible teeth;
but an examination under a
higher power will show the
presence of about six small,
backwardly directed serra-
tions on the inner edge, close
up to the tip, as in Text-
fig. 4, 6. These undoubtedly
help in preventing the escape
of the prey, when once it has
been pierced by these power-
ful jaws. On the ventral
side of each mandible, there
is a very definite groove {gr)
which receives the maxilla,
lying directly below it. The
length of the mandible is
2-7 mm.
The J/axiZ^fc (Text-fig. 4, cj
are not unlike the mandibles
in size and shape, but are
slenderer, and slightly more
curved towards the tip. They are entirely devoid of armature,
except for their very sharp points. Each maxilla carries a very
faint groove upon its dorsal surface, a little wider than the groove
Text-fig. 4.
* a, Mandible of larva of Pst/chopsis ekgam {Guer.), third instar; ( x 30).
b, Tip of same, to show the series of fine serrations; ( x 87). c, Maxilla of
same; c, cavdo; p, palpiger; st, stipes; ( x 30).
796 STUDIES IN AUSTRALIAN NEUROPTERA, vii.,
of tlie inaudible. In the living larva, the maxilUe ma}^ be seen
at times playing freely forwards and backwards beneath the
mandibles; so that it is evident that tliey have considerable free-
dom of movement, in spite of being designed to act as a pair of
complete sucking-tubes in conjunction with tliese latter.
Whereas the groove in the mandible can be followed with ease
right up to tlie tip, that of the maxilla cannot be so followed,
and is evidently of a much more imperfect nature. Hence the
complete sucking-tube must be formed at least partly by pressure
of the two paiis of jaws, one upon the other, and not by a close
interlocking of parts. I have frequently seen the tips of the
maxillae pushed a considerable distance forward beyond the tips
of the mandibles; this could scarcely happen unless the fit of the
maxilla upon the ventral surface of the mandible were a some-
what loose one, seeing that the calibre of the groove varies
greatly from base to tip.
When the maxilla is dissected out, it is seen to be considerably
swollen near the base, and to carry, below the swollen part, three
flat, chitinised plates, two of which ai'e broadly triangular, the
third very narrow. These are placed in the positions of cardo^
sti^jes, and palpiger of a normal maxilla, and appear to represent
those three sclerites. As regards the elongated spear that forms
the principal part of the maxilla, this would appear to be the
galea. But, unless the transformation of these parts, from their
abnormal form in the larva to their normal form in the pupa,
can be followed out at metamorphosis, in such a way that there
can be no doubt upon the matter, I do not think that we can be
sure of their homologies.
7'Ae Lahritm and Clypens (Text-fig. 5). These together form
one piece in the larva, indistinctly divided transversely, as shown
in the figure. The clypeus is covered with the usual hooked
hairs, set upon raised papillae The labrum is a strongly pro-
jecting curved lobe, carrying only three rather large hooked
hairs on either side.
The Labium (Text-fig.6). This consists of an undivided tri-
angular basal portion, from the two anterior angles of which
project the five-jointed palpi. The basal joint of each palp
BY R. J. TILLYARD,
797
carries a single hooked liair on its inner side, and there is a
larger one situated just below it. Five or six smaller hooked
hairs are situated on either side of the middle line, upon the
basal portion of the labium.
This basal portion of the labium remains undivided through-
out all stages in Psychopsis, and probably represents both
inentum and subinentum combined.
Text-fis.'").^
Text-fi.i^.O.t
7'he Body- Armature (Text-figs. 7, 8). We have already de-
scribed the peculiar hooked hairs, set upon raised papillae, which
are found upon the head of the larva. They also occur upon the
hard tergal plate of the prothorax, which is formed of tough
dark brown chitin, like that of the head. Upon the rest of the
thorax, and upon the abdomen, these hairs are almost entirely
replaced by more highly specialised structures, which I shall call
dolichasters (Greek SoA-iyo?, long; and da-ri'ip, a star). The doli-
chaster is a more or less elongated, hollow, chitinous structure,
set upon a definite papilla; it is very narrow at the base, but
expands distad in narrow pyramidal form, and ends distally in
from four to nine sharply projecting points, which pass beyond
the general periphery of the enclosing chitinous membrane.
Labium and clypeus of larva of Ps, eleyans (Guer,), third instar; ( x 30).
t liabium of larva of P.s. e/egaiis (dui'-r. ), third instar; ( x 30i.
b'2
798
STUDIES IN AUSTRALIAN NEUROPTERA, Vll.,
Viewed end on, it is more or less star shaped, being in the form
of a fairly regular polygon with concave sides. Typical doli-
chasters are shown in Text-fig. 7, a, b, and c. These structures
are very abundant all over the abdomen and thorax, excepting
upon the hard tergal plate of the pro thorax, on which only an
occasional one can be seen.
The dolichaster is clearly a specialisation from a normal hollow
sensory hair or macrotrichion. This may be proved by a study
of the eighth and ninth segments of the larva. On the ninth
segment, the macrotrichia are of normal form. On the eighth,
yv there are a few of normal form,
but most of them are slightly
"^ thickened and blunted at the tips,
and a few can be seen having the
projecting distal points or angles
of the true dolichaster-form. On
the seventh, the dolichaster-form
is fully established, but remains
very elongated, for the most part.
On the rest of the abdomen, all stages from a very narrow,
elongated dolichaster with only four or five rays, to a shorter
and stouter dolichaster with from seven to nine rays, can be
easily followed.
Dolichasters vary from 40 to over 100/x in length, and from
about 20 to 30/x in extreme width distal ly.
We may compare the dolichaster with the bulla, such as is
found in the larva of Micropteryx (Order Lepidoptera). In the
bulla, the macrotrichial chitin becomes very soft, and the internal
cavity is enlarged so that the hair expands into the form of a
soft, swollen bulb. Weak longitudinal supporting rays are, how-
ever, present, as may be seen by a careful examination of the
bulla under a high power. Moreover, when the bulla is viewed
end on, it, too, like the dolichaster, exhibits the star-shaped
* Dolichaster.s from integument of larva of Ps. elegans (Gii^r. ), third
instar; ( x 330). a. Long, five-ra3'ecl form; h, four-rayed form; c, shorter,
seven-raj'ed form. Above h and c are shown four examples of micrasters,
also X 330.
Text-fig. 7."
BY R. J. TILLYARD. 799
distal end, thougli tlie peripliet-yof the soft cliitinous bulb, in this
case, projects out well beyond the outline of the star. Tlius the
bulla is a higher specialisation from the original macrotrichion
than is the dolicliaster, and can be developed from the latter by
further softening of the chitin forming it, and further swelling
up of the internal cavity.
The dolichaster may also be compared ver}'^ closely with a
typical scale, such as is found upon the bodies of Collembola, or
upon the wings of Lepidoptera. If the dolichaster were to
become flattened down upon itself symmetrically from the side,
it would differ ver}^ little from a scale; the longitudinal edges or
rays would become the longitudinal stride of the scale, and the
sharp distal points of the dolichaster Avould remain as a series of
distally projecting angles, such as are very commonly found in a
large number of Lepidopterous scales, particularly amongst the
Heteroneura.
Besides the dolichasters, the body of the larva in Psychopsis
carries, in a number of places where the chitin is very soft, an
immense number of minute, star-shaped structures, which I shall
call micrasters (Greek iJ.iKp6<i, small; and wrri'jp). Where they
occur, they are developed from every single hypoderm-cell in the
neighbourhood, and are thus many times more numerous than
the dolichasters of the same region. The micraster is a minute,
sessile, flattened stai-, not set upon any definite papilla, and
raised but little above the general surface of the integument.
Text-flg.7 shows a set of four micrasters with four, five, six, and
seven rays or points respectively.
The micrasters vary from 1 to 3/x in height, and from 8 to 12/x
in extreme width. Thus they are always smaller than the basal
papillse of the dolichasters, which range from 12 to over 20/x in
width.
It can be easily seen that the micraster is simply a specialisa-
tion of the minute microtrichia which occur normally upon many
parts of the body of an insect, especially in the sutures and
other places where the chitin is soft, in the form of tiny hooked
hairs. A search over the larval integument of Psychopsis shows
that, for the most part, these njicrotrichia are only feebly de-
800
STUDIES IN AUSTRALIAN NEUROPTERA, Vll.
velopecl, as miiiute processes flattened down close to tlie integu-
ment itself. In other places, they can be seen to be divided
into two short processes at an obtuse angle to one another.
From this form, the transition to a four-ra\'ed micraster is quite
a simple one. As microtrichia are developed from every single
unspecialised hypoderm-cell, in the regions in which they occur,
it follows that the same will be true of the micrasters in any
given region, as we find to be the case here.
Piiiacida, or small plates of hardened brown chitin, carrying
one or more hairs or setie, occur upon the meso- and metathorax,
and also upon the sixth to eighth ab-
dominal segments. Each segment men-
tioned has a single pair of pinacula placed
more or less dorso-laterally.
The simplest pinacula are those of the
abdomen, of which one is shown in Text-
fig. 8, together with its corresponding-
spiracle It is a somewhat irregular oval
patcli, which carries a single excessively
elongated and slender hair, quite unlike
any other hair to be seen in this larva,
and about one-third of a millimetre long.
The two pinacula of the eighth segment
are placed rather close together, on either
side of the mid-dorsal line. Those of the
J "p*^ seventh segment are placed further apart.
Those of the sixth are much smaller, and
• Sp still further apart. Sometimes a vestige
of a similar pinaculum may be observed
Text-fig 8.* upon the fifth segment also, in good
chitin-preparations.
The pinacula of the meso- and metathorax are much larger
and darker areas, whose position has been already indicated in
' Details from the integument of seventh abdominal segment of larva
of Ps. eJ e ff cms {Gwer.), third instar. d, A dolichaster; pn, pinaculum with
long, slender macrotrichion; s]!, spiracle, in situ; ( x 167). Three other
dolichasters surrounding the spiracle are omitted.
BY R. J. TILLYARD. 801
f] -scribing the colour pattern on p.793, since they occupy the
darkisli spots tliere mentioned, and shown in Plate Ixxix., figs. 4,
5. Each pinaculum carries from two to four dolichasters, and a
single small seta upon a raised papilla. In specially good pre-
parations, I have seen the blind ending of a small trachea upon
the integument close to the pinaculum; this would appear to
indicate the original position of the meso- or metathoracic
spiracle, which has evidently become suppressed.
The Sjnracles. — In all three larval instars, the number of
functional spiracles is eighteen, viz.. a pair upon the prothorax,
and a pair upon each of the first eight abdominal segments. They
are all very small, simple openings, the rim of which is slightly
strengthened and darkened. As far as I can see, the}^ are quite
devoid of armature or ornamentation of any sort. A single
trachea, whose calibre is approximately equal to that of the
spiracle itself, passes inwards from it to the main tracheal trunk
on each side. The prothoracic spiracles are the largest pair:
next in size are those of the eighth segment, with those of the
seventh, sixth, etc., in descending order of magnitude up to the
first. Text-tig. 8, .<?/;., shows one of the spiracles of the seventh
segment in situ, with its corresponding pinaculum (all but one
of the surrounding dolichasters removed). Owing to the great
number of these latter, it is always difficult to locate the spiracles
in this larva.
The Anal Papilla (Text-fig.9).— The last two segments of the
abdomen, viz., the ninth and tenth, are much narrower than the
rest, and are more or less retractile within the eighth. They do
not carry dolichasters, but only very delicate, slender macro-
trichia of reduced size. The tenth segment forms the anal
papilla or sucker, by means of which the larva is enabled to move
rapidly backwards, or to hold on tight when attacking its prey.
This segment is furnished with a pair of dorso-lateral curved
processes, probably representing the original cerci, and a pair of
ventral processes, somewhat cornute in shape, and curving out-
wards to meet the incurving tips of the dorsal processes. These
latter are armed with two series of closely set, short, sharp setaj,
one set directed outwards and the other inwards. I have found
802
STUDIES IN AUSTRALIAN NEUROPTERA, Vll.
it impossible to observe how these organs are used by the larva;
but the result is certainly to give it a very powerful grasp upon
anything that they grip, and the whole papilla appears to act as
forcefully as a true sucker. Whether there is, besides the actual
grasping effected by these appendages, any sucking action per-
formed by the tenth segment, I cannot say. Remembering that
the silk for the cocoon is spun from the anus, it seems possible
that an exudation of the same nature may help in the action of
the anal papilla as a grasping or sucking organ.
Text-fig. 9.
Anal papilla of larva of Z^-, c/egans (Guer.), third instar; ventral view;
( X 87).
Habits of the Larva.
In order to discover the larva in its natural haunts, it is
necessary to provide oneself with a stout, wide-bladed chisel, or
some other instrument that will act as a lever for removing the
bark of Myrtaceous trees. The ordinary methods of bark-collect-
ing, as practised by ColeopUrists, are useless in this case. All
old trees, in which the bark is hanging in shreds or long strips,
or is dry and attacked by white ants, are of no use in searching
for Pttychopsis larvie. Instead, it is necessary to select healthy,
medium-sized trees, in which the sap is running freely, and in
6y r. j. tillyard. 803
which, consequently, tlie l>ark,*\vhen torn away, is found to be
slightly moist inside. The best trees are those in which the
bark is thick and rough, with many crevices, as in the case of
the Ironbark {Eucalyptus siderophloia Bentli.) and Blood wood
{E. corymbosd 8ni.). Probably the Ironbark is one of the best
trees of all for this larva; but it is so difficult to work, on account
of the extreme toughness of its bark, that it is best to leave it
alone, unless one sees a piece of bark that can be levered off
without damage to one's tools. Around Sydney, one of the very
best trees is the Bloodwood; but in Queensland, where the bark
of this tree is thinnei" and more scaly, I met with little success
in exploring it. Another excellent tree is the Forest Apple
(Aitgophora inter meaia DC). Trees in which the bark comes
away in long strips of considerable thickness, such as the Tallow-
wood (E. microcorys F. v.M.), Turpentine {Syncarpia laurifolia
Ten.), and Stringy-bark {E. oapitellata Sm.), generally provide
very good hunting also: but those with thin bark are useless.
In searching for the larva, of which, probably, not more than
five or six at the most would be found on any given tree, even
if all the bark were to be stripped from it, much time and trouble
may be saved if one watches for likely places where the larva
may be expected to hide. Such are, for instance, particularly
thick or richly creviced portions of bark; and, above all, the
cracks and crannies surrounding a gum-flow. Many insects
come at night to taste this gum; and it is to be presumed that
the Psychopsis lar\a takes up his station near by, with the special
object of attacking these insects. On One Tree Hill, Brisbane,
I noticed a rather small Ironbark-tree, in which there was a
good gum-flow, caused by a small cut with an axe. By levering
up the two tough projecting ends of the bark, above and below
this cut, r secured four larvte of Ps. elegans and two of Ps. atdi-
vagus — th^ best haul I ever obtained from a single tree.
When a piece of bark is removed, the Psychopsis larva will
almost always be found upon the trunk of the tree, with its body
flattened down, and quite motionless. Thus, the trunk and its
exposed crevices should be examined first; then, if nothing is
seen there, the piece of bark may be likewise examined, and
804 STUDIES IN AUSTRALIAN NEUROPTERA, vii.,
sometimc'tj vields a larva. Veiy*soon the larva will begin to
move rapidly backwards, feeling all the time, with its mobile
anal papilla, for a suitable crevice in which to hide. Jf once it
succeeds in reaching one, it will not be easy to secure it. Hence
it should be picked up at once with the forceps, and transferred
to a glass-tube, whose open end should be stopped up with cotton-
wool. Not more than one larva must be put into one tube,
unless each is separated from the next by a plug of cotton-wool;
for one is sure to attack the other and suck him dry.
The larva may be reared right through, in a single SRiall glass-
tube. The best food for the earlier stages is white-ants. These
should be given alive, but with their heads crushed; otherwise
there will be a fight between the larva and its intended victim,
in which the tables may be turned. In the second and third
instars, the larvae of the Oodlin Moth make excellent food; but
these also must be offered with their heads crushed, unless one
is willing to take the risk of damage to the Psychopsis larva from
the strong mandibles of the Codlin grub. Under natural con-
ditions, the Psychopsis larva guards himself from attacks of this
kind by retreating into a crevice, after he has seized his prey.
With his body thus covered, and only his large jaws projecting,
he is perfectly safe. But it is different in a glass-tube, where
the whole of the larva's soft body is exposed to attack from the
jaws of his writhing victim.
The larvae feed but seldom; each meal, however, is a very sub-
stantial one. A larva supplied with a Codlin grub will usually
attack it at once, advancing cautiously with its jaws wide apart,
until they are well placed on each side of its victim. Then, with
a sudden vicious snap, the jaws are driven home, and the victim
is secured. If the victim struggles furiously, the Pyychopsis
larva will frequently let go its hold, and retreat into a corner,
with every sign of fear; nor will it be induced to attack the same
victim a second time. Generally, however, the larva is able to
hold on, until the loss of blood occasioned by the wound weakens
the victim sufficiently to prevent its struggling further. Then
the jaws are driven far in, and the play of the maxillae to and
fro beneath the grooved mandibles shows that the larva is suck-
BY R. .1. TILI.YAKD. 805
iiig vigorously at its victim. It may take some hours before the
latter is completely emptied of its juices. By this time, the
Psychopsis larva will have swollen out to a very great extent,
and will frequently be quite torpid, like a snake. After such a
meal, no food should again be offered for two or three weeks.
Larv?e that are overfed usually die from the effects of it; so that
it is necessary to keep a strict watch on the feeding, and only to
offer food to those that are really in need of it.
The Cocoon. (Plate Ixxix., fic^ji).
When the larva is full fed, it remains torpid for a considerable
period, usually two or three weeks. It then sets about con-
structing its cocoon. First of all, a number of loose and irregular
threads are spun from the anus, making an irregular mesh work,
the size and appearance of which depends entirely upon the place
selected for the cocoon. Under natural conditions, the cocoon
is spun in a crevice of the bark, which is usually barely wide
enough to contain it. In such a case, the preliminary meshwork
is reduced to a minimum, and only consists of a slight scaffolding
or anchorage for the cocoon. But, in such an artificial condition
as in a glass-tube, the larva may expend a considerable amount
of skill upon this meshwork, before it is satisfied that conditions
are satisfactory for the actual building of the cocoon.
Having selected the exact position for the cocoon, the larva
next spins a small platform within the meshwork already men-
tioned. Upon this it lies, back downwards, while it weaves
around itself, with mar\ellous dexterity, the beautiful spherical
cocoon. The outer sheath of the cocoon is mostly composed of
fairly loose and irregular threads; as these are spun in larger
and larger number, the spherical shape of the cocoon begins to
take shape. Within tliis loose outer covering, the threads are
spun with wonderful accuracy and closeness, so that the larva is
finally enclosed in a dense white or cream-coloured ball of silk,
which, when cleared of its outer and looser threads, closely re-
sembles a pearl. As the spinning proceeds, the larva shrinks
more and more in size, and becomes doubled right over upon
itself. When the cocoon is finished, it becomes almost impossible
806 STUDIES IN AUSTRALIAN NEUHOPTERA, vii.,
to conceive how so large a larva could have managed to encase
itself in it.
As regards the actual mode of spinning the thread, the mobility
of the anal papilla is very marvellous. Lying upon its back, the
larva can protrude this papilla to a great distance, and spin silk-
all round itself, even behind its head and thorax, with very little
effort. Most of the threads, however, ai'e spun transversely
across the larva, beginning with a small arc at the anal end,
then swinging out into a longer arc across the middle of the
abdomen, and finally making shorter arcs behind the back of the
thorax. All these transverse strands appear to be carefully con-
nected, on both sides, with the small platform already mentioned,
which itself is made to form a part of the outermost coat of the
cocoon. When the first complete layer of the sphere is finished,
the larva has its head bent over so as almost to touch the tip of
its abdomen: but the mobility of the anal papilla appears to be
but little decreased in this position. In spinning tlie inner layers
of the cocoon, the larva changes its position every now and
then, so that the anal papilla may cover all parts of the sphere
with an equally thick layer of silk.
As found under natural conditions, the cocoon is an oblate
spheroid, with the two poles in contact with the sides of the
crevice in which it is placed, and the larva lying so that its
sagittal section is in the equatorial plane of the spheroid. When
spun, however, in an open tube, the cocoon is practically a sphere.
It closely resembles the cocoon of a Chrysopid, particularly that
of the genus Nothochrysa, from which it would not be possible
to distinguish it for certain. But, unlike most of the Chrysnpidoi,
Psi/chojjsis does not attempt to spin any external matter into
the outer mesh of the cocoon. Such a proceeding is, of course,
unnecessary, considering the position in which it is placed. The
diameter of the cocoon varies with the size of the larva, but is
usually about 5 mm. (Plate Ixxix., fig. 6).
The Pupa. (Plate Ixxix., %. 7; Text-tig. 10).
As far as the external form is concerned, the true metamor-
phosis is that between larva and pupa. The latter is a pnpa
BY R. J. TILLTARD. 807
libera, closely reseiiibliug the imago in everything except its
uuexpanded wings (Plate Ixxix., fig. 7). At the metamorphosis,
the pupa experiences a great difficulty in casting oflP the hard
larval head, and a number perish from inability to do so. The
rest of the larval skin is soft, and shrivels up into a minute mass
within the cocoon; the head may be found sometimes almost
intact, sometimes with the mandibles and maxilhv broken off
from it. The splitting of the head is mid-dorsal ly and postero-
transversely, as in the other larval ecdyses (Text-fig. 3).
The change from larva to pupa does not take place until a
considerable time after the spinning of the cocoon, generally
from a fortnight to three weeks. At first, the fresh pupa is
almost colourless, the body having a slight greenish tinge, the
head and wings cream-coloured. The compound eyes are large,
and soon become dark brown and functional. If the cocoon be
opened, the pupa watches every movement with anxiety, and
can be made to turn round and round, merely by the movement
of a pin held at a short distance in front of the head.
The antennce, are fairly short (Plate Ixxix., fig. 7), but are com-
posed of a large number of joints, there being thirty-five in the
pupa which I dissected. Each joint is less than half as long as
wide, except only the first three; of these, the basal joint is
stouter than the rest. Unlike those of the imago, the antennte
in the pupa are devoid of hairs.
The mouth-parts are of considerable interest (Text-fig. 10;. The
lahruin is distinctly bifid, and carries numerous hairs on small
raised bases. Below the labrum, on either side, are the large
and very strongly chitinised mandibles. Each of these consists
definitely of two lobes, separated by a narrow slit. The outer
or distal lobe has a sharply pointed, tooth-like apex, and, below
it, a more or less broad and fiat cutting-area, forming a right-
angled projection. The inner or basal lobe is rounded, and much
less prominent. The two mandibles are not similar in shape.
The right mandible has the smaller basal lobe, but the broader
and flatter cutting-surface; wliereas the left mandible has the
cutting-area narrower, and the apical tooth raised above it on a
high ridge. Thus, in the action of cutting open the cocoon, for
808
STUDIES IN AUSTRALIAN NEUROPTERA, Vll.
which tliese strong mandibles are employed, it is evident that
the apex of the right mandible phiys in and out of the groove
formed between the apical ridge of the left mandible and the
Texl-ti.u-. 10.
Mouth-paits of pupa of /^. <:lc<ians ((Ua-r.). », Lal>ruiii (///r) and nvAU-
(lil)les (wcZ). /), Right maxilla and labium; cd cardo, ya galea, la
lacinia, Ip labial palp, ml median lobe of labium, mxp maxillary palp,
8t stipes; ( x 48).
rectangidai- shelf below it. This is also proved by the fact that
the cutting-edge of the distal lobe of the right mandible is very
sharp, whereas that of the left mandible is thicker and blunter,
BY R J. TILLY A RD.
809
and is evidently formed more as a pad or receptacle, against
which the I'ight mandible can work.
The maxilhe and labinni lesemble those of tlie imago fairly
closely, hut are broader and softer, as if the parts were all
swathed in whitish bandages. The principal differences are that
the galea of the maxilla is a broad rounded lobe without a small
distal joint, and the median lobe of the labium is simply bifid,
instead of being formed as in the imago.
After about a week, the imaginal colon is loegin to appear upon
the body and wings of the pupa. Before emergence, the whole
beautiful pattern of the wing-colouration becomes set fortli upon
the wings of the pupa. It was by this means that I was able to
recognise the pupa of Ps. ccelivagus, with its intensely black wings,
in spite of the fact that it failed to complete its emergence,
after escaping from its cocoon.
Emergence of the Imago. (Text-fig. 11).
The pupa emerges from the cocoon by cutting it neatly open
with its sharp mandibles, whose scissor-like action is specially
adapted to this purpose. Alwut one-fourth of the circumference
is left uncut, forming a strong hinge. The pupa pushes up the
lid of the cocoon, and climbs out.
On Dec. 1st, 1915, at 5.30 p.m., I noticed a pupa emerging
from its cocoon, which was placed upon some sand in a glass-jar.
The pupa crawled along the sand to a strip of blotting-paper,
placed nearly vertically, near by. Up this it climbed to a height
of about one and a half inches, and then took a firm grip of the
blotting-paper with its claws, and remained resting for a few
minutes. During this time, it gradually swelled up and became
very taut, as shown in Text-fig. 11, a. At 5.37 p.m., it began to
work its abdomen and wing-sheaths about; so that, in a few
seconds, the fine pupal skin split dorsally down the thorax, tlie
abdomen became straightened out, and the imago began to
emerge from the pupal skin. During emergence, the wings begin
to expand at once, the insect arching tliem strongly outwards, as
may be seen in Text fig. 1 ], c,d. The basal portions of the wings
begin to enlarge before the apical portions are freed from the
810
STUDIES IN AUSTRALIAN NEUROPTERA, Vll.,
pupal wing-sheaths. Consequently, as in the case of the Stone-
flies or Perlaria, the wings, when finally withdrawn, have the
distal portion bent at an angle to the basal portion, as sliown
in Text-fig.ll, d. This applies more to the forewing than to the
hind, which becomes freed with less difficulty.
e.
Text-fig.ll.
Emergence of the imago of Ps. ehyans {Gu^r.). a, Pupa just before meta-
morphosis, 5.36 p.m. h, Imago emerging, 5.40 p.m. c. Imago freed
from pupal skin, 5 42 p.m. d, Imago expanding its wings, 5.47 p.m.
e, The same at 5.49 p.m. /, Imago resting with wings fully expanded,
6.8 p.m. (Drawn from sketches made while watching the actual
emergence on Dec. Ist, 1915; d, dorsal view, the rest lateral).
As soon as the imago was quite free, it climbed upwards
further away from the pupal skin, the latter being left clinging
to the blotting-paper, as shown in Text-fig.ll, c. Having taken
up a position well above the pupal skin, the imago remained
stationary with its wings arched strongly outwards, the forewing
BY R. J. TILLYARD. 811
at first complete!}' hiding the liindwing. While the basal half
of the forewing expanded rapidly, the distal half remained bent
under it for some time. Meanwhile, the hindwings expanded
rapidly, and their apical portions appeared beneath the still
bent forewings, as seen in Text-fig.ll, d. Next, the distal por-
tions of the forewings became straightened out, though at first
they remained full of longitudinal rucks, as seen in Text-fig.ll, e.
The wings were held well away from the abdomen up to 5.49
p.m., by which time they were about three-fourths expanded.
Then the apical rucks became rapidly smoothed out, and the
wings were arched forwards, forming a steep roof over the body,
so that the costal margins of the forewings come almost into
contact with the blotting-paper (Text-fig.ll,/"). From the time
of leaving the cocoon to the time that the wings were fully ex-
panded, only thirty eight minutes elapsed. After resting for
some time in the position shown in Text-fig. 1 1,/j until its wings
were sufficiently hardened, the imago spread them out in the
broad, flattened, roof-like manner usually adopted in this family.
The pupal skin left behind is a flimsy, wliite structure. It is
very difficult to detach from its position. But, when once this
has been done, the slightest breath of air causes it to move
readily, so light and delicate is its construction. The thorax
and base of the abdomen are flattened, and wide open dorsally.
From the prothoracic spiracles, two long threads stand up; these
are the intima of the tracheal trunks, withdrawn through these
spiracles when the imago emerges. The back of the head is split
open, and the large dark brown pupal mandibles are left gaping
wide apart.
The Imago. (Text-fig. 12).
A few details about the imago which have not been published
may suitably be given here.
The mouth-parts are figured in Text-fig. 12. The lahrum re-
sembles that of the pupa, but is not definitely bifid, there being
only a very slight indication of the division into two distinct
lobes. It carries numerous hairs, not set upon raised bases.
The mandibles ai-e smaller and weaker than in the pupa, and
show considerg^ble modification in shape. The ri^ht mandible
812
STUDIES IN AUSTRAIJAV XEUROPTKRA, VU.,
in the imago has only a single lobe, with a less prominent apical
tooth and a broader cutting-blade. The left mandible, on the
other hand, is still bilobed, the basal lobe being rounded as in
the pupa, but the distal lobe forming a single strongly projecting
Text-fig. 12.
Mouth-parts of imago of Ps. ehfjans (Guer.), 9 . a, Labruni and mandibles.
/>, Left maxilla and labium; A/? hypopharjnix; rest of lettering as in
Text-fig. 10; (X 48).
tooth, whose internal border is definitely hollowed out. Thus,
in the imago, the cutting-edge of the right mandible works into
the groove of the left in a very simple manner. These mandibles
may possibly be used for feeding upon gum-exudations. In cap-
BY R. J. TILLYARD. 813
tivity, the insects feed willingly upon sugar moistened with water
and spread upon cottan-wool, using their mandibles to chew the
sticky crystals.
The maxiUfe are of a very generalised type, with complete and
separate galea and lacinia, and an elongated, five-jointed palp.
The stipes is long, the cardo very short. The lacinia arises from
a broad oblique base, in the form of a simple elongated lobe with
rounded apex, and carrying numerous hairs. The galea is
remarkable in being set well above the lacinia, upon a short
transverse base formed as a definite projection from the stipes,
and also in carrying a small but very definite distal joint. The
basal joint is somewhat club-shaped, and carries numerous hairs
upon its distal half. It is possible that the existence of a distal
joint is an archaic feature, and that it ma}' prove to be of import-
ance in the study of the Phylogeny of the Holometabola.
The maxillary palp is carried, in the position of rest, winh its
distal joint turned inwards almost at right angles to the other
four, as shown in Text- fig. I'J, h. The first and second joints are
shoi-t, the third twice as long as the second, the fourth shorter
than the third, and the distal joint longer than any, with a
somewhat pointed tip.
The labium is in the form of a broad!}' rounded median lobe,
on each side of which there is a narrower lobe, also rounded, but
folded over above the median lobe, between it and the hypo-
pharynx. The median lobe carries hairs on either side, distally;
the two lateral lobes carry numerous hairs upon their upper
surfaces. The palps, which arise low down towards the base of
the median lobe, are three-jointed, the basal joint being the
shortest, and the distal the longest; each joint carries a few hairs.
In the natural position, if the mouth be opened, there will be
seen abroad sub-triangular lobe projecting outwards and slightly
upwards above the labium. This is the hypopharynx (Text-fig.
\2,hp). Its distal border is well rounded, rather strongly
chitinised, and curved over; it carries no hairs.
Bearing in mind the fact that the Psychopsidce are the most
ancient of existing l^lanipennia, as far as the Palieontological
record of this Order is known, it would seem that much valuable
63
814 STUDIES IN AUSTRALIAN NEUROPTERA, vii.,
evidence should be obtainable when a careful comparison can be
made between the mouth-parts here described and those found
in other families of the Order. This research, I hope to carry
out later on.
Habits of the Imago.
During the day-time, the imagines rest concealed and motion-
less, either upon the underside of a leaf, or hidden away in debris
of dead leaves and sticks. Hence they are seldom captured,
though occasionally one may be taken by beating.
At night time, they become comparatively lively. Ps. eUyans,
kept in a large glass-jar, was watched by me for several nights
from 8 to 10 p.m. During this time, it fluttered about the jar,
visited the moist sugar supplied as food for it, and fed upon it
for periods up to half-an-hour in duration. The third night,
after all the insects had fed, I was fortunate enough to see the
process of pairing. The female took up a position upon a strip
of bark, resting with her wings in the usual position, but con-
tinually vibrating them. A male, after several short flights,
at last alighted upon the same piece of bark, lower down, and
began to climb up towards the female, also vibrating his wings.
At last he arrived alongside her, on her right side. The female
then raised her right pair of wings, and the male moved in
towards her from the side, so that the abdomens of the two
insects were almost parallel, and the right pair of wings of the
female covered the left side of the male. In this position they
remained for pairing, occasionally vibrating their wings. It
was not easy to make out exactly how copulation was effected;
but it appeared that the male bent the tip of his abdomen round
towards the female, and seized her with his anal appendages, at
the same time pulling the tip of her abdomen partly round
towards him. Thus they remained for some twenty minutes,
until a second male came and alighted close to them, and began
to flutter around and over them.
The female apparently did not lay any eggs until some time
after pairing, as I searched the jar the following morning and
found none. However, during the next few days, she deposited
eggs singly, and at long intervals of time apart, in the cotton-
BY R. J. TILLYARD. 815
wool which I had supplied for that purpose. She also explored
the bark, but found it for the most part unsuitable, laying only
one egg there, as against eleven laid on the cotton- wool. The
two eggs shown in Text-fig. 1 were laid close together as drawn,
but all the rest were quite separate. This female may, of course,
have laid other eggs which I failed to find. I should say that a
single female must contain at least fifty eggs;* but it is possible
that they would not usually all be laid after a single pairing.
The males were kept alive for periods varying from ten days
to three weeks. The females live longer, and one was kept alive
for over a month, by which time her wings were considerably
torn. The only nourishment given, during all this time, was
sugar and water, which I find sufficient for most Planipennia
and Mecoptera. A very slight degree of moisture is required in
the jar. Either excess of moisture, or exposure to a hot drying
wind, is equally injurious to these insects The same may be
said of all Planipennia, although the MyrmeleontidcH and Ascala-
phidce can stand greater heat and desiccation than any of the
others.
The Economic Value of the Psychopsidae.
There is no Order of Insects, with the exception of the
Hymenoptera, whose members are so generally beneficial to man-
kind as the Planipennia. It seems, therefore, of considerable
importance that we should now attempt to estimate the value of
the Psychopsidce in this respect.
From the account of the life-history here given, it will be seen
at once that these insects are entirely beneficial to man through-
out their whole life. The preference shown by the larva for
feeding upon such obnoxious insects as the larv?e of Codlin
Moth, and other of the smaller Lepidoptera, at once establishes
it as potentially a very useful factor in the checking of insect-
pests. Hence, if colonies of Psychopsis could be established in
our apple-, pear-, and quince-orchards, there is no doubt that
they would help very materially in checking the Codlin Moth
and other Lepidopterous pests that still do so much damage
therein.
* Mr. Gallard's original female laid over fifty eggs. See p. 787.
816 STUDIES IN AUSTRALIAN NEUROPTERA, vii.,
In attempting to estimate the practicability of sucli a sugges-
tion, we have to bear in mind certain factors that must, at the
best, very much limit tlie efficacy of the Psychopsidce : —
(1) Although a single larva of Ps. eleyans, during its life,
would most certainly consume a considerable number of Codlin
Moth larvae, yet the value of this is much discounted by the
long life of the larva, which takes up the best part of two years.
The economic value of this insect would be many times enhanced
if it were al^le to pass througli all three larval stages in a few
weeks, as is the case with the Chrysopidce and Hemerobiidce.
(2) The larva requires rough-barked trees to live in. Thus it
would seem that it could only be successfully introduced into
orchards in which the trees were of considerable age. Such trees
would probably afford the requisite amount of shelter to tlie
larva, which would certainly take heavy toll of anj" other insects
that attempted to hide away in the crannies and crevices of the
bark.
(3) As an archaic survival of a very old stock, the Psychopsidce
could scarcely be expected to show that readiness to adapt them-
selves to new conditions, that is to be found, for instance, in the
more specialised Chrysopidce and Nemerobiidce. Many of our
Australian species of these two families have readily established
themselves in our orchards and gardens, without any attempt on
the part of entomologists to place them there; and are already
acting very effectively as checks upon Aphidse and Scale Insects.
So far, no similar tendency has been noted in the Psychopsidce;
unless, indeed, the apparent increase in the numbers of Ps.
insolens in such districts as Killarney and Mount Tambourine,
in South Queensland, where orchards are rapidly replacing the
original bush, is an indication that this species is beginning to
accommodate itself to new conditions. It would be natural to
suppose that insects which, in a state of nature, are so rare as
the Psychopjsidce, would not take kindly to a change of environ-
ment which man might attempt to force upon them. Thus, if a
selected orchard of old trees were to be well stocked with Psy-
chopsis-hdYvse, one would anticipate that the resulting imagines,
after pairing, would return to the nearest pifece of untouched
BY R. J. TILLYARD.
817
l)ush, and lay their eggs, as usual, on the hai'k of Myrtaceous
trees.
(4) The apparent association of the FsycJtoiJsidre with trees
belonging to the Natural Order Myrtacene nnist be considered as
a disadvantage, if it is actually a fact. But, so far, we do not
know the life-histories of most of the VsyrJiops\dm\ and it may
well be that this supposed limitation does not hold for most of
the species. Moreover, it is noteworthy that, although Fs.
insolens is our commonest species, no larva of this insect has yet
been found under Eucalyptus-bark. If the life history of this
species can be worked out, it seems probable that it might prove
to be of considerable value as a beneficial insect in checking
orchard-pests.*
(5) The small number of eggs laid by the females is a distinct
disadvantage economically. Against this, however, we may
place the fact that the females live a long time, and deposit their
eggs singly in many different places, instead of all in a lum}>.
Thus the maximum effect is produced for the small number of
larvte hatched, and the danger of a quick reduction through
cannibalism is eliminated.
Taking all these factors into consideration, I have come to
the conclusion that, although the Psychopidce do not offer us the
pi-omise of such immediately beneiicial results as could be ob-
tained by a scientific breeding and distribution of Chrysojyidtr.
or Hemer'obiidce, yet they are a group that most certainly ought
not to be neglected by economic entomologists. A few carefully
devised experiments upon old and badly infected orchards, either
with the larvte of Fs. elegans, or with those of Fs. insolens., if
they can be obtained by pairing the imagines, should be well
worth carrying out, and might conceivably yield results much
superior to the expectations that I have here indicated. There
is also the possibility that these insects, like so many other Aus-
tralian animals, would do exceptionally well in some new region,
* Ff<ychopsid(i; occur also in Africa and Asia, but nothing is known of
their life-histories in these regions. Large, rough-barked Mj'rtaceous
trees, like the P]ucalypts, being absent from these regions, it is evident
that they must be associated with trees of other Orders.
818 STUDIES IN AUSTRALIAN NEUROPTEKA, vii.
such as California or South Africa. As they are entirely bene-
ficial, I would call the attention of entomologists to their possible
value in this respect; particularly as there would not be the
slightest difficulty in sending the larvse to any part of the world,
since they are very hardy, and will go without food for several
weeks at a time.
EXPLANATION OF PLATE LXXIX.
Psychopsis elegans (Guer.).
Ficr.L— Two eggs; ( x /).
Fig.2. — Newly hatched larva; { x 7).
Fig.3. — The same larva at end of first instar; ( x 7).
Fig. 4. — The same larva, at end of second instar; ( x 7).
Fig. 5. — The same larva, at end of third instar; full-fed; ( x 7).
Fig. 6. — Cocoon; ( x 4).
Fig. 7. — Pupa, ten days old, extracted from cocoon; ( x 7).
819
AUSTRALIAN MEGALOPTERA OR ALDER-FLIES,
With Descriptions of new Genera and Species.
By R. J. TiLLYARD, M.A., D.Sc, F.L.S., F.E.S., Linnean
Macleay Fellow of the Society in Zoology.
(With three Text-figures.)
The Megaloptera are an archaic Order of Holometabolous
Insects, which contains, at the present day, only about one
hundred known species, all of them of large or moderate size.
The Order is divisible into two very distinct Sub orders, viz.,
the Sialoidea, or true Alder-flies, and the Raphidioidea, or Snake-
flies. Of these, the aquatic Sialoidea are clearly the more archaic,
the terrestrial Raphidioidea being a highly specialised offshoot
from them.
The Raphidioidea are not represented in Australia, their place
in the economy of Nature being already occupied by the older
Fsychopsidm of the Order Planipennia. whose larvae, though
differing in the nature of their mouth-parts, live, like those of
the Snake-flies, in crevices of the bark of trees, and prey upon
the insects frequenting them.
The Sialoidea, or true Alder-flies, are represented by some
sixty species throughout the world. They may conveniently be
divided into two very distinct families, according to the follow-
ing key :—
'Large insects (expaii.se -loto 100 mm. ^, with three ocelli present;
fourth joint of tarsus not bilobed; venation regular, with
cross-veins weakly formed, Larvfe with eight pairs of lateral
gills and a pair of hooked anal prolegs, but without any ter-
minal filament CORYDALID.^;.
Much smaller insects (expanse 20 to 40 mm.), without ocelli;
fourth joint of tarsus strongly bilobed; venation less regular,
Avith strongly developed cross-veins. Larvie with only seven
pairs of lateral gills and a terminal filament, but without
any anal prolegs . . Sialid.i;.
820 AUSTRALIAN MEGALOPTEtlA OR ALDER-FLIES,
Of these two families, the Corydalidce, containing the great
majority of the genera and species of the Sub-urder, may be
again divided into two subfamiHes, as follows: —
'More than three cross- veins between R and Rs; head quad-
rangular CORYDALIN.i:.
Onh' three cross-veins between R and Rs; head triangular
Chauliodin.i;.
Up to the present, only one species of Alder-fly has ]>een re-
corded from Australia, viz., Archichauliodes guttiferus (Walker).
It is a large, dull- coloured insect, recorded occasionally from
many localities in Eastern Australia, but nowhere very common.
Both in size and markings, it is ver}^ variable, some individuals
having large black spots on the wings (especially on the hind-
wings) and others having none at all. The larva is a familiar
oljject to naturalists in the Sydney district, being found under
rocks and stones in small rocky creeks. In New Zealand, the
allied species A. dubitatus (Walker), is very abundant, and its
larva is much esteemed as bait for trout, being known as the
" Black Creeper," a name that would suit the larva of the Aus-
tralian species equally well. ,
Turning now to the smaller family Sialidce, of which no repre-
sentatives have hitherto been found in Australia or New Zealand,
we find that, of the two known genera, Sialis is Holarctic, reach-
ing from Canada through Europe and Siberia to Japan, and
down into Asia Minor; while Frulosialis is confined to the New
World, one species being found as far south as Chili.
It would thus appear that, as the SialidcG do not occur in the
Oriental region, there is no possibility of their ever having
reached Australia from the North. But, in so far as they are
admittedly archaic forms, and one species is recorded from Chili,
it has to be admitted that there is a bare possibility of their
having been able to reach Australia from the South, provided
that we admit the truth of the Antarctic Theory so ably cham-
pioned by Mr. Hedley.
That being so, I have always kept in mind the possibility of
a true Sialid being discovered in Tasmania, or on some isolated
mountain-top in Eastern Australia. I now have the pleasure of
BV R. J. TILLVAHb. 821
recording the discovery of two very distinct new species of this
family, each of which will form the type of a new genus. The
first of these was taken by Mr. G. H. Hardy, late of the Tas-
manian Museum, Hobart, during a visit to Maria Island in
December, 1915. Only a single specimen was taken; but it is
in good condition, except for the loss of the antennae. The
second is a remarkable slender form, taken at light in December,
1916, on Mount Tambourine, South Queensland (elevation 1900
feet) by Mr. W. H. Davidson. Not realising what a prize he
had got, Mr. Davidson unfortunately enclosed this specimen in
an envelope with a letter to me; with the result that, when I
received it, the insect was completely flattened out, and other-
wise damaged. However, I managed to relax and set it, though
the pin had to be inserted through the thorax from side to side,
the wings being extended at right angles to the flattened thorax.
I have delayed publishing these fine discoveries, in the hope
that further material might be obtained in better condition.
Mr. Hardy again visited Maria Island last year, but he failed to
find any further specimens of the new Sialid; likewise Mr.
Davidson, who has kept a sliarp look-out for his new species, has
met with no further success. As the new insects are of great
scientific interest, further delay is no longer justified.
The following key will enable the two new genera to be dis-
tinguished from each other and from the two genera already
described, at a glance : —
Wings about one-third a.s broad as long, tlie costal Held of the
forewing noticeabl3- broadened .. 2.
Wings much narrower, the costal field of the fore wing not
noticeablj'^ broadened 3
THindwing with M1+2 ^^d M3 f 4 both simple veins Sin/is Latr
(2) -
jHindwing with M1+2 forked, M3 , ^ simple Aas(ro^iui/,s, u.ir.
ith K2-f3 and K4 f 5 both forked Sltnosial it<, n.g.
with R2-}-3 simple, and Ri-fo forked ... Profo^ia/is Weele.
(1)
TBoth wings w
j Both wings w
AusTROsiALis, n.g. (Text figs. 1-2).
Closely related to Sialis Latr. General colouration black,
with orange-red prothorax and smoky wings. Forewing with
822
AUSTRALIAN MEGALOPTERA OR ALDER-FLlES,
about ten veinlets in costal space, hindwing with about six. In
forewing, Cu^ unites with M for a space, the short basal free
portion of Cu^ resembling an oblique cross-vein. The fused por-
tion, M + Cui, is continued by Cu^ itself as a stout vein in line
with it, whereas M departs from it anteriorly at a sharp angle.
In hindwing, Cu remains quite distinct from M. In forewing,
the full complement of branches is present for both Rs and M;
in hindwing, there is one less for M, M3_|.4 remaining unbranched.
Neither R2 J^oi' 1^3 is secondarily forked, as is the case with one
or both in Sialis.
Genotype, Aiistrosialis Ignicollis, n sp.
Hab. — Tasmania.
M3>4
Text-fig. 1.
Wings oi Anstrosicdis ignicoUis, n.g. et sp. In forewing, M + Cui is the
fused portion of M and Cu,. Rest of notation as usual in the
Comstock-Needham system. The small forking distallj' on R^ does
not occur on the left forewing. (Hindwing 12 mm. long).
The black and orange colouration is also that of P^'otosialis,
to which the Chilian species belongs. It is also interesting to
note that, in the number of its branches to Rs and M, the new
BY K. J. TlLLYARD.
823
genus offers an exact parallel to the oldest Trichoptera, in which
Ms^-i is always a simple vein in the hind wing.
AusTROSiALis iGNicoLLis, n.sp. (Text-figs. 1-2).
Total length, 8 mm.; abdomen, 4 mm.; foreiving, 14 mm.; hind-
tving, 12 mm.; exjyanse, 28-5 mm.
Head shining black, smooth anteriorly, with a shallow mid-
dorsal depression; the large occipital
region curiously sculptured, with raised
longitudinal and circular areas, ar-
ranged as shown in Text-fig. 2. Aii-
tennoi with large, stout basal joint,
black ; the rest missing. Mandibles
black, tipped with orange-red.
Thorax: prothovcix bright orange-
red, flattened cylindrical in shape, with
slightly indicated mid-dorsal groove;
less than half as long as wide, the
width being slightly less than that of
the occiput. Pterothorax jet black.
Legs black, of medium length and slenderness.
Abdomen [shrivelled] dull blackish.
Type, in Coll. Tillyard.
Hab. — MdLv'ia, Island, East Coast of Tasmania. A unique
specimen, probably a female, taken on Dec. 29th, 1915, by Mr.
G. H. Hardy.
•
Stenosialis, n.g. (Text-fig.3).
Closely related to Protosialis Weele. General colouration
brown, with pale smoky-brown wings. Forewing with narrow
costal space containing only four veinlets, the hindwing with
only two. The arrangement of the veins M and Cu is the same
as described for Austrosialis above, and the number of branches
The cross-
Text-tig.2.
for the veins Rs and M is the same as in that genus
* Head and prothorax of Austrosialis ignicoUis, n.g. et sp., to show
sculpture of the occiput; the light arranged so as to fall nearl^^ horizon-
tally; { X 15).
824
AUSTRALIAN 3IEGAL0PTERA OR ALDER FLIES,
vein between R2+3 ^^^^ ^4-1 5 is exceptionally^ oblique in direction.
Genotype, Steiwsialis aitstraliensis, n.sp.
JI(tb South Queensland.
As with Aitstrosialis, so with this genus; the arrangement of
the branches of Rs and M corresponds exactly with that of the
oldest Trichoptera, M3_|_4 being branched in the forewing, but
simple in the hind. One is tempted to ask, is this merely the
result of a chance convergence, or does it indicate a closei" rela-
tionship between the two Orders Megaloptera and Trichoptera
than has hitherto been suspected 1
Text-tig. 3.
Wings of Stenosialis aastrallensi-^, n.g. et sp. Notation as in Text-tig. 1.
(The wings of the type-specimen being considerably torn, the figure
was completed by combining portions of both right and left pairs of
wings). (Hindwing 10 mm. long).
Stenosialis australiensis, n.sp. (Text-fig. 3).
7'otal length, 8 mm.; abdomen, 4 mm. ; forewing, 11 -5 mm.;
hindiving, 10 mm.: expanse, 25 mm.
Head brownish, a darker area posteriorly on occiput, isolat-
ing several paler raised circular areas placed close together on
either side of the mid-dorsal depression. Eyes dark brown.
Antennm dark brown, the basal joint large and stout, the second
joint small; the rest missing.
BY K. J. TILLYAHD. 825
Thorax dark brown. Prothorax apparently slightly wider
than occiput. Legs brown. [Too crushed for accurate descrip-
tion].
Abdomen [shrivelled] blackish basally, shading to brown
anally.
Type, in Coll. Tillyard.
Hah. — Mount Tambourine, South Queensland (1900 feet). A
unique specimen, probabl}' a female, taken on Dec. 17th, at
light, by Mr. W. H. Davidson.
In concluding this paper, I desire to record my grateful
thanks to Mr. Hardy and Mr. Davidson for these two fine
species, and to congratulate them on their discoveries. The
knowledge that, in this Order, archaic forms of. great scientific
value are still to be found in isolated places in Australia, should
stir up other entomologists to look out for these insects, which
are easily cautjht and recognised.
826
CONTRIBUTIONS TO A KNOWLEDGE OF AUSTRA-
LIAN CULICID.E. No. iv.*
By Frank H. Taylor, F.E.S.
(From the Australian Institute of Tropical Medicine, Townsville).
(Plates Ixxx.-lxxxiii.)
This paper may be divided into two parts, synonymical and
descriptive, with notes on previously described species.
Two species described by me from the Northern Territory
belong to species recorded from extra-Australian localities;
their distribution is noted in the text. The Australian Anopheles
are also tabulated. It will be seen that very few of the Ano-
plieline genera are recognised, and those only as subgenera,
except in the case of Bironella.
The new species are distributed in the following genera : —
Pseudoskusea (one), Mimeteomyia (tw^o), Culicada (one), Lof>ho-
ceratoiyiyia (one), Uranotcenia (four), and Hodgesia (one).
The type-specimens are contained in the Institute Collection.
The following tabulation of the Australian Anopheles is given
in the hope that it may prove useful in the determination of the
species in question.
Anopheles corethroides, A. stiymatiats, and B. gracilis have
been tabulated from descriptions only, as the Institute does not
possess specimens of them. A . pu7ictulatus Donitz, is omitted
from the Table, as it is unknown to me either from specimens or
description.
It will be seen that all the species, except B. gracilis, are
placed in the genus Anopheles. This is best explained by re-
ferring the reader to a paper by A. Alcock,t which deals with
the "Classification of the Culicidse," and with which the writer
* Continued from these Proceedings, 1916, p. 574.
+ Ann. Mag. Nat. Hist., (8), viii., p. 240 (1911).
BY FRANK H. TAYLOR. 827
entirely agrees, as the genera cannot, in some cases, be regarded
even as subgenera; while, in others, they form convenient sub-
generic or group-names.
Alcock* retains the following as subgenera : Aiiopliiles (s.r.),
Myzorhynchus, Myzomnyia^ and N yssorhynchus. I would add to
these the genus Bironella, as valid on venational characters— if
it really be an Aiiopheline. Theobald, himself, throws doubt on
its systematic position.
AXOPHELES.
A. Wings spotted,
i. Legs unhanded.
a. Wings with the third, fourth, fifth, and sixth long veins
brown- and white-scaled. No costal spots afratipes Skuse.
//. Wings with one fringe-spot JtarhiroatrisY. d. Wulp.
hh. Wings with several fringe-spots.. J>a/'6//'067r/.'^ var. />a?icro/i!t Giles,
ii. Legs handed.
a. AVings with numerous light and dark spots.
Costa with six spots.
Femora and tibiae with numerous pale bands, tarsals 1-4
Avith apical and basal banding annul ipes Walker.
B. Wings unspotted,
i. Legs unhanded.
rt. Thorax with long, curved, hair-like scales... corethroidej^ Theobald.
aa. Thorax with three rows of golden-j'ellow, narrow-curved
scales .^figiuaticns 8kuse.
BiRONELLA.
A. Wings unspotted, third long vein, stem of second fork-cell,
and fifth long vein curved,
i. Legs unhanded.
a. Fi rst fork-cell very small yracilis Theobald.
Anophkles (Myzorhynchus) barbirosthis Yan d. Wulp,
var. BANCROFTi (Giles).
(Plate Ixxx., fig.l).
Theobald, Mon. Culicid., v., p. 50 (1910;; Taylor, Proc. Linn.
Soc. N. 8. Wales, 1915, xl, p.l76.
This is a common and widely distributed species, extending
from Darwin, N. Territory, to Eidsvold, 8. Queensland.
The male is evidently very retiring, as the only known speci-
mens have been bred from larvae.
* Journ. Lond. 8ch. Trop. Med., ii., p. 153 (1913).
828 AUSTRALIAN CUIJCIDiE, iv.,
Anopheles (Nyssorhynchus) annulipes Walker.
Ins. Saund., p. 133 ( 1 850); Theobald, Mon. Culicid., v., p.oT
(1910); Taylor, Proc. Linn. 8oc. N. S. Wales, 1914, xxxix., p.454;
Quarantine Service Publication, Melbourne, No. 12, 1917.
/frt^>.— N.S.W.: Hay (W. W. Froggatt). -Victoria: Melbourne
(Dr. J. H. L. Cumpston); K37abram, Bamawm, Echuca, Swan
Hill, Mildura, Cohuna, Lake Boga (F. H. Taylor).— S Australia:
Renmark, Cobdogla, Pompoota (F. H. Taylor).— W. A. : Perth,
Kalgoorlie (Dr. Atkinson).
This mosquito is probably the most widely distributed species
in Australasia, as it extends from Darwin, in the North, across
to Papua, throughout the mainland, and southward to Tasmania.
It is not improbable that this species will be found in the Pacific
Islands.
ToxORHYNCHiTES INORNATUS (Walker).
Proc. Linn. Soc Lond , viii., p.l02 (1865); Theobald, Mon.
Culicid., i., p.223 (1901); op. cit., v., p.llO (1910).
Two male specimens, one of which is quite typical; the other has
the mij^ cross- vein confluent with the posterior cross- vein instead
of not meeting it, and also has the second tarsals of the hindlegs
with a broad, white, almost basal band, which occupies more
than half tlie joint. It agrees in all other points with Theobald's
description of this .species.
//a6.— Papua: Itikinumu Plantation (F. P. Dodd).
ToxoKHYNCHiTES sPECiosus (Skuse). (Plate Ixxx., fig.2).
Proc. Linn. Soc. N. S. Wales, 1888, xiii., p. 1722 (1889); Theo-
bald, Mon. Culicid., v., p.l08 (1910).
ffab. — Q.: Townsville (Dr. A. Breinl).
MUCIDUS ALTERNANS (Westwood).
Ann. Soc. Ent. Fr., iv., p. 681; Taylor, Proc. Linn. Soc. N. S.
Wales, 1914, xxxix., p.455; op. cit., 1915, xl., p. 176.
//a6. —Victoria : Kyabram (F. H. Taylor).
This species is known to range from Darwin to Victoria.
Armigeres bkkinli Taylor; Neosquamomyia hreinli Taylor.
(Plate Ixxx., fig.3).
Trans. Ent. Soc, London, 1914, Pt. i., p.l86.
BY FRANK H. TAYLOR. 829
The genus Neosqtiamomyia was founded partly on the male
genitalia, the other characters agreeing with Armigeres.
When proposing the genus, I was unaware that similar char-
acters were to be found in species of the genus Arniiyeres^ notably
A. malayi Leicester.
Edwards* notes the similarity of A. malayi Leic, and K.
breinli Taylor, and suggests that the correct position for the
latter species is in Armiyeres, a decision with which I quite agree.
The two species are distinct, but certainly closely related.
PSEUDOSKUSEA CAIRNSENSIS, Sp.n.
9. Head clothed with black, flat, and upright-forked si'ales;
palpi dusky; antennae dark brown; proboscis black.
Thorax dark reddish-brown with brown scales, scutellum
similar; pleurae brown, with white, flat scales.
Abdomen black-scaled, unhanded, segments three to the apex
with basal, lateral, white patches; venter pale-scaled.
Legs black, unhanded; ungues equal and simple.
• Wings brown-scaled; first fork-cell longer and narrower than
the second, its base nearer the base of the wing; stem of first
fork-cell one-third the length of its cell, stem of second fork-cell
slightly more than half the length of its cell; anterior basal cro.ss-
vein longer than, and twice its own length distant from, the
anterior cross- vein. Length, 1
/Tafe. — Queensland : Cairns (F. H. Taylor).
Described from a single specimen bred from a mixed lot of
larvae. It is readily distinguished from P. multij^hx Theobald,
on venational characters, ungues, and the abdominal spots. It
differs from P. basalis Taylor, in not having a banded abdomen.
MlMETEOMYIA ATRIPES (Skuse).
Stegomyia punctolateralis Theobald.
Proc. Linn. Soc. N. S. Wales, 1888, xiii., p.lToO (1889); Theo-
bald, Mon. Culicid, iv., p.l90(1907); Taylor, Proc. Linn. 8oc.
N.S.Wales, 1914, xxxviii., p.750 (191 5) (.S'c'?t^omyut); o/y. cit.,
1915, xl., p. 177 (Steyomyia).
It is quite evident, from specimens recently collected by myself,
* Bull. Ent.Res., vii., p.207 (1917).
830 AUSTRALIAN CULICIDiE, iv.,
that Skuse's species has the apical lateral spots on all the
abdominal segments, and not only on the sixth to eighth seg-
ments, as stated in ray paper (1914). The Ingleburn specimens
show that the spots vary somewhat in size on the basal segments,
so that they were either overlooked in the type, or the abdomen
was partially abraded.
There is also no doubt at all that Stegomyia punctolateralis
Theobald, is the same as M. cUripes (Skuse). ^S'. punctuIateraHs
Theob., was unknown to me, except from description, when I
stated (1914) that the two were different species; but, since then.
Dr. Bancroft has presented a series of both sexes to the Institute,
which clearly show they are the same as Skuse's species.
Theobald states that the mid-ungues of the male are unequal
and simple; this is an error, as the larger is uniserrate.
Neveu-Lemaire* places J/, atripes in llieohaldia, and records
it from Guyane. He states that the fore- and mid ungues of the
female are uniserrate; and he gives a figure of the wing-scales of
his specimens, which proves conclusively that he was notdealino;
with Skuse's species. He also states that the palpi are four-
jointed.
There is no doubt that it belongs to the genus Mimeteomym,
as, inter alia, the apex of the abdomen is very bristly.
Hab.— Q.: Eidsvold (Dr. Bancroft), Burketown, Townsville
(F. H. Taylor).— N.S.W.: Milson Island (Dr. Ferguson), Black-
heath (W. A. Thompson), Ingleburn (F. H. Taylor). -Vic. :
Miidura (F. H. Taylor).
MiMETEOMYiA ATRA (Taylor).
Stegomyia at7'a Taylor, Trans. Ent. Soc. London, 1914, p. 190.
A re-examination of the type of this species reveals the fact
that it should, more correctly, be placed in the genus Mimeteo-
myia, on account of its bristl}^ and truncated apex of the abdo-
men, among other points of agreement with the genus.
MlMETEOMYIA PULCHERRIMA, sp.n.
^. Head covered with brown, fiat scales, with a median row
of white ones, and white ones laterally; antennae pale, nodes and
* Arch. Parasitologie, vi., p. 615 (1915).
BY FRANK H. TAYLOR. 831
plumes brown, basal lobes with white, flat scales; eyes winered;
palpi brown-scaled, first segment with two prominent white
bands, penultimate and apical segments with white, basal bands.
'J'horax with dark brown scales, and ornamented with a lyre-
shaped pattern of small, white, flat scales; prothoracic lobes with
white scales and black bristles; scutellum with white, flat scales;
pleurae brown, with patches of white, flat scales.
Abdomen brown-scaled, expanded apically, segments one to
six with small, median, white, basal patches; all segments, except
the first, with white, lateral, basal patches, prominent in the last
three segments; genitalia with numerous black bristles; venter
brown-scaled, with median, white, basal patches on the first
seven segments.
Legs brown, knees white, first and second fore- and mid-tarsals
with basal, white banding, posterior tarsi one to three with
broad, white, basal bands, fourth white, with a narrow, brown,
apical band, fifth white; ungues of fore- and mid-legs unequal,
the larger with a distinct notch, hind equal and simple.
Wings : bases of fork-cells equal, first longer and narrower
than second, stem of first fork-cell not quite half the length of
its cell, stem of second slightly more than half the length of its
cell, anterior basal cross- vein longer than, and twice its own
length from, the anterior cross-vein; vein-scales brown.
Length, 3 mm.
Hab.~Q.: Cairns (F. H. Taylor).
Described from a single specimen, bred from a collection of
larvae. It is abundantly distinct from all other Australian
species of Mimeteomyia.
MlMETEOMYIA DODDI, Sp.n.
(J. Head covered with black scales, a median row of white, flat
ones, and white, flat ones laterally; antennas brown, plumes brown,
internodes pale; palpi brown, penultimate segment basally banded
white, apical segment white-scaled; eyes silvery.
Thorax covered with dusky-brown scales; scutellum with brown
scales; pleurae brown, with patches of white, flat scales.
Abdomen covered with coppery-brown scales, first segment
832 AUSTRALIAN CULICIDiE, iv.,
brown-scaled, second with a median, basal, white-scaled spot,
third to fifth and apical segments with white, basal banding, all
segments with white, lateral, basal spots; venter brown, with
white basal banding and numerous pale hairs.
Legs dusky-brown, first tarsals of forelegs basally banded
white, the rest unhanded, first tarsals of mid-legs basally banded
white [tlie rest broken off], posterior tarsi one to three basally
banded white, fourth unhanded, fifth white: ungues of forelegs
unequal, the larger with a tooth, hind equal and simple.
Wings : first fork-cell longer and narrower than the second,
its base nearer to the base of the wing; stem of first fork-cell
one-third the length of its cell, stem of second slightly more
than half the length of its cell, anterior basal cross-vein longer
and twice its own length from the anterior cross-vein.
9. Similar to (J. Antennae brown, basal half of first joint
yellow; apical third of palpi white. Abdomen : fourth and fifth
segments with basal, white banding, apex of abdomen white-
scaled, and with numerous pale bristles, all the segments, except
the first, with lateral, white, basal spots; second segment with a
white, median, basal spot; venter pale-scaled, apical segments
black-scaled.
Legs similar to those of ^, second tarsals of mid-legs with white
basal banding; ungues equal and simple. Wings as in ^.
Length : ^, 2*5; 9, 3-5 ram.
//a6. — Papua : Itikinumu Plantation (F. P. Dodd).
A very distinct species, readily distinguished from J/, p-iil-
cherrima mihi, by its palpi, thoracic and abdominal markings.
There is little doubt that the female belongs to the same species.
It affords me mucli pleasure to name it in honour of its
discoverer.
Grabhamia theobaldi Taylor. (Plate Ixxxi., fig. 4).
Theobald, Mon. Culicid., iv., p. 304 (1907). C flavifrons
Theob., nee Skuse, Proc. Linn. 80c. N. S. Wales, 1913, xxxviii.,
p.Tol (1914).
Were it not for intergrading forms, it would be possible to
make two distinct "species" out of the series of specimens before
BY FRANK H. TAYLOR.
833
me, on abdominal markings alone, as, in some specimens, the
abdomen is quite typical, whilst, in others, it varies from speci-
mens with distinct basal banding on the first two segments of
the abdomen, and the remainder with more or less indefinitely
banded and mottled, to those with indefinite basal banding and
dense mottling of creamy scales on all the abdominal segments.
The base of the second fork-cell is slightl}^ nearer the base of
the wing than that of the first, while they are nearly level in
the type.
Specimens from Eidsvold, Queensland, show similar inter-
gradient forms to the above Victorian specimens.
Hal). — Vic: Bamawm, Kyabram, Mildura, and Echuca (F. H.
Taylor).
CuLiCADA wiLSONi, n.sp. (Plate Ixxxi., figs. 5, 6, 7).
(J. Head black, covered with yellowish, narrow-curved and
upright-forked scales, with a few yellowish hairs projecting over
the eyes, sides of head with flat ones; palpi longer than proboscis,
black, first joint pale-scaled, except the apical fourth, pubescence
black on the apical segment, and apex of first, yellowish-brown
on the penultimate segment; antennse pale, nodes dark, plumes
brownish-black, dense.
Thorax black, covered with yellowish, narrow-curved scales;
prothoracic lobes with flat ones; scutellum similar; pleura? black,
with flat scales.
Abdomen black, first segment pale-scaled, second to seventh
with broad, yellowish, basal banding; sixth, seventh, and eighth
mottled; some specimens show median pale-scaled bands on most
of the segments; genitalia with some long, black hairs, latera\
pubescence dense, yellowish; venter pale.
Legs black, femora pale beneath; femora, tibise, and tarsi
mottled; fore-ungues unequal, uniserrate, mid unequal, the larger
deeply notched, the smaller uniserrate, hind equal and simple.
Wings with brown scales; first fork-cell longer and narrower
than the second, base of latter nearer the base of the wing, stem
of the first almost as long as its cell, stem of second as long as
the cell; anterior basal cross-vein about as long as, and nearly
834 AUSTRALIAN CULlCID^, iv.,
its length from, the anterior cross-vein; second incrassation well
marked,
^. Similar to ^ ; antenna^ black, basal lobes black, with
creamy-yellow scales, basal half of second joint pale, verticillate
hairs black, pubescence pale; palpi four-jointed, black-scaled,
with intermixed, scattered, pale scales; proboscis black; fore
femora pale-scaled, mid and hind pale, with an apical black ring,
tibiae and first tarsals mottled with pale scales; fore- and mid-
ungues equal, uniserrate; penultimate abdominal segment creamy-
scaled except for a small oblique apical line of dark scales, apical
creamy-scaled. Wings similar to those of the male; but fork-
cells relatively longer, stem of first fork-cell two-thirds the length
of its cell, that of second about three-fifths of the cell.
Length: (J, 7-5; 9, 6 mm.
Hab. — Vic.-: Kyabram, Echuca, Bamawm, Swan Hill, and
Mildura (F. H. Taylor).
This appears to be a well-defined species related to C. tasrna-
niensis Strickland, differing, i')Uer alia, in the wing-venation,
and the mottled femora, tibitie, and first tarsals. C. tvHsoid was
present in enormous numbers in the Goulburn Valley district,
Kyabram and Bamawm being inundated with them. There is a
small area of cypress-pine {Callitris sp.) at Bamawm, and it was
impossible for man or beast to remain in it for even a few
minutes, owing to the abundance of this mosquito.
I have much pleasure in dedicating this species to Mr. Wilson,
of Bamawm, who rendered me much help and kindly service
while I was in that district.
OCHLEROTATUS NOTOSCKIPTUS (Skuse).
Proc. Linn. Soc. N. S. Wales, 1888, xiii., p.r738 (1889) [CWex];
Theobald, Mon. Culicid., v., p.200 (1910) [Scutomyia'\\ Edwards,
Ann. Mag. Nat. Hist., (8), ix., p.523 (1912).
Hah. N.S.W.: Ingleburn.— Vic: Swan Hill (F. H. Taylor).
CuLEX siTiENS Wiedemann.
Aussereurop. zweiflug. Ins., p. 544 (1828); Theobald, Mon. Culi-
cid., v., p.331 (1910); Taylor, Proc. Linn. Soc. N. S. Wales, 1916,
xli., p. 570.
BY FRANK H. TAYLOR. 835
Hab — Q.: Cairns, Innisfail (F. H. Taylor).— Papua (F. P.
Dodd).
This common mosquito enjoys a wide distribution^ being found
from Java through Papua and Torres Straits to Victoria.
CuLEX CONCOLOR Desvoidy.
Mem. Soc. d'Hist. Nat. Paris, iv., p.405 (1825); Edwards,
Bull. Enb. Res., ii., p.262 (1911).
Hab.-Q: Innisfail (F. H. Taylor).
A short series, bred from larvae, have been compared with
specimens from other Australian localities, and also with speci-
mens of G. concolor, C. tigri2:)es, C. consiniilis, and the form de-
scribed as C. tiginpes var. Jusca, which have been received from
the Imperial Bureau of Entomology; and there is absolutely no
doubt, as Edwards states, that the Australian form belongs to
6'. concolor.
All references dealing with C. tigrvpes, as from Australia,
should, therefore, refer to C. concolor^ and G. tigripes should be
expunged from the Australian list.
CULEX BITiENIORHYNCHUS GileS.
Journ. Bombay Nat. Hist. Soc, xiii., p.607 (1901); Edwards,
Bull. Ent. Res., iv., p. 231 (1913). G. ahdomhialis Taylor, Rep.
Aust. Inst. Trop. Med., 1911, p.53 (1913).
Edwards gives several synonyms of G. bitcKniorhyiichiis Giles,
in his paper, and mentions Guliceha abdominalis Taylor, also as
a possible synonym.
I have compared specimens of my species with six specimens
of C. bitamiorhyiichus Giles, from Hong Kong, and can see no
valid reason for treating them as a distinct species, thus con-
firming the opinion held by Edwards.
CuLE}^ viSHNUi Theobald.
Mon. (JuHcid , i., p.355 (1901) (9 only); Edwards, Bull. Ent.
Res., iv., p. 233 (1913). G. parvus Taylor, Bull. N. Territory, la,
p.27 (1912).
Edwards included G. jmrvus Taylor, as a probable synonym of
G. vishiiui, when dealing with its synonymy in his paper. I
836 AUSTRALIAN CULICIDiE, iv.,
quite agree with tliis, and place C. ^;arv?ts definitely as a synonym
of C . vishnai Theobald.
It seems to me that C. vishimi 1'heobald, is closely related to
C. silietis Wied.
CuLEX TOWNSViLLENSis, nom.nov.
Culicelsa fusca Taylor.
Trans. Ent. Soc. London, 1914, p.699.
A change of name is necessary, as fnscus is preoccupied in
Culex.
The sixth and seventh segments of the abdomen have the
sides covered with white scales, except the apex; eighth pale-
scaled, unhanded. The fore- and mid-ungues are unequal, and
not equal, as stated in the description, the mid more so than the
fore. There is also a small branched process on the undersurface
at the base of the ungues in both legs. Genitalia of male of
typical Culex -form.
9. Similar to ^. Palpi brown-scaled; apex of abdomen hairy.
First fork-cell longer and narrower than second, its base nearer
the base of the wing than that of the latter, stem of the first
about one-third the length of the cell, stem of the second slightly
more than half the length of the cell; hind-tibia? the same length
as first tarsals; ungues equal aiid simple.
Ilab. — Q.: Townsville (F. H. Taylor).
The above corrections in the description of the male are based
on fresh material, which agrees perfectly with the type. It
appears to be an uncommon species.
(JULEX FATIGANS Wied.
Aussereurop. zweiflug. Ins., p. 10 (1828); Taylor, Trans. Ent.
Soc. London, 1914, p. 197.
//rt6.— N.S.W.: Sydney (F. H. Taylor).— Vic: Melbourne (Dr.
Cumpston), Kyabram, Echuca, Mildura, Bamawm, Swan Hill
(F. H. Taylor). — S. Australia : Kenmark, Cobdogla, Overland
Corner, Pompoota, Adelaide (F. H. Taylor).
C. fatiyans is a common mosquito in Southern Australia. It
occurred abundantly at Kyabram and Echuca, where it was
BY FRANK H. TAYLOR. 837
breeding in enormous ninnhers in the irrigation-channels and
street- watertables.
LoPHocEKATOMYiA CAiKNSENSis, sp.n. (Plate Ixxxii., figs. 8, 9).
(J. Head clothed with brown, narrow-curved, and black,
upright-forked scales; antennse pale, apical segments and nodes
dark brown, plumes brown, accessory organs long on segments
five, eight, and ten, apices spoon-shaped on eighth, short on sixth
and seventh; palpi black, penultimate and apical segments with
black hairs beneath, thumb-like process dark brown; eyes black.
Thorax and scutellum with brown, narrow-curved scales ;
scutellar bristles black, long; pleurpe brownish.
Abdomen black-scaled, unhanded, apex bristl}^, segments three
to seven with faint, lateral, basal spots; venter dark.
Wings : first fork-cell longer and narrower than the second,
the former nearer the base of the wing; stem of first fork-cell
about half the length of its cell, stem of second about two-thirds;
anterior basal cross-vein longer than anterior cross-vein, and
nearly thrice its own length distant from it; scales brown.
Legs black, unhanded; ungues of forelegs unequal, the larger
uniserrate, mid-ungues unequal and simple, hind equal and simple.
9. Similar to (J. Palpi black-scaled, firsi segment with a few
black bristles; clypeus black; antennae brown; ungues all equal
and simple; abdominal spots well defined.
Length: ^, 3*5; 9, 4mm.
Hab. — Q.: Cairns (F. H. 'J'aylor).
Described from seven males and ten females, bred from larvae.
It is abundantly distinct from L. anuulata Taylor, and L. cylin-
drica Theobald.
Two males and one female have the bases of the fork-cells
almost level, but there are no other difterences from the typical
specimens.
LoPHOCERATOxMYIA ANNULATA Taylor.
Proc. Linn. Soc. N. S. Wales, 1916, xli., p.57L
Ilab. — Q.'. Cairns (F. H. Taylor).
A single specimen, bred at the same time as L. cairnsensis
mihi; it agrees with the type in all respects.
838 AUSTRALIAN CULICID^, iv.,
LoPHOCERATOMYiA CYLiNDRiCA Theobald.
Mon. Culicid., iii., p. 202 (1903) [Cidex\ PI. ix.; o}?. cit., v.,
p.361 (1910); Edwards, Bull. Ent. Res., vii., p.228 (1917).
Hab.~Q.: Eidsvold (Dr. T. L.- Bancroft).
I quite agree with Edwards in placing this species in Lopho-
ceratoniyia. It is typical in every respect. The figure of the
head of the male on Plate ix., though small, shows the plates on
the antennae.
Mansonoides uniformis (Theobald).
Theobald., Mon. Culicid., v., p.i-lS (1910); Edwards, Bull. Ent.
Res., ii., p.253 (1911).
Hab.~N. Territory: Darwin (G. F. Hill).— Q.: Eidsvold (Dr.
Bancroft), Cairns, Townsville. — N.S.W : Newcastle (Dr. Dick).
This is a very abundant and widely distributed species, being
found from Africa to the Philippine Islands, New Guinea across
to Darwin, and extending as far as Newcastle, N.S.W.
Finlaya poicilia Theobald.
Mon. Culicid., v., p. 464 (1910); Taylor, Proc. Linn. Soc. N. S.
Wales, 1914, xxxix., p.465.
Ilab. — Q.: Cairns (F. H. Taylor).
MoLPEMYiA PRiESTLKii (Tavlor). (Plate Ixxxii., fig. 10).
Calomyia priestleii Taylor.
Trans. Ent. Soc. London, 1913, Part iv., p.684 (1914).
Calomyia cannot be retained as distinct from Jlolpemyia, there
being no structural differences. The writer was in error in
describing the scales on the centre of the liead and bordering
the eyes as spindle-shaped, as they are, in reality, only large
narroiD-curved scales.
The species seems to be somewhat variable, as a specimen,
taken at a later date than the type, shows complete broad band-
ing on the abdomen, whereas, in the type, the abdomen has large,
basal spots on the segments.
Skusea pseudomediofasciata Theobald.
Mon. Culicid., v., p.489 (1910).
Ilab.—Qr. Cairns (F. H. Taylor).
Two specimens, both males, before me, agree perfectly with
BY FRANK H. TAYLOR. 839
the descriiDtion of tliis species They were bred from larvte. It
was originally described from Ceylon.
Uraxot.enia albescens Taylor.
Trans. Ent. Soc. London, 1914, Pt. i., p. 705.
Hab.-Q,.: Cairns (F. H. Taylor).
The fourth segment of the abdomen is more correctly described
as brown, with a median, apical, white-scaled area. The Cairns
specimens, bred from larvse, show the fourth segment entirely
black.
Uranot.enia cairnsensls, sp.n.
2- Head with dusky-brown, flat scales, with a fairly broad
band of bluish-white scales round the eyes; antennae dark brown,
])asal lobe and base of first segment yellowish, palpi black;
clypeus dark brown; eyes silvery; proboscis dusky-brown.
Thorax brown, with brown scales and black bristles; scutellum
with flat, black scales; pleurte with white, flat scales; there is a
short, pre-alar, white line of scales.
Abdomen with dusky-brown scales, all the segments with
lateral, white scales; venter with white scales.
Legs dark brown, femora pale beneath, ungues very small,
equal and simple.
Wings longer than abdomen; first fork-cell shorter and nar-
rower than second, latter considerably nearer the base of the
wing, stem of first more than twice the length of its cell, stem
of second slightly longer than its cell; anterior basal cross-vein
longer than the anterior cross-vein, and nearly twice its length
from it; halteres pale, with black knobs.
Length 1-5 mm. (vix).
IJab. — Q.: Cairns (F. H. Taylor; July, 1917).
Described from two specimens, bred from larva\ It is easily
separated from other Australian species by its venational and
abdominal markings.
XJRANOTiENiA TIBIALIS, sp.n. (Plate Ixxxii., fig. 11).
(J. Head clothed with white scales, except in the centre, where
they are brown; antennae brown, basal lobes black, basal half of
first segment pale; palpi dark brown; eyes black; proboscis brown.
840 AUSTRALIA?? CULICID.E, iv.,
Thorax with brown scales, and long, black bristles, apparently
in four rows; a short, pre-alar line of bluish-white scales; scu-
tellum with brown scales, pleurae brownish, with bluish-white,
flat scales.
Abdomen covered with black scales; venter brown-scaled.
Wings longer than abdomen; first fork-cell shorter and nar-
rower than second; base of latter much nearer the base of the
wing; stem of the first nearly twice the length of its cell; stem
of second about the length of its cell; anterior basal cross- vein
longer than anterior cross- vein, and once and a half its length
from it.
Legs dusky-brown, femora basally pale beneath; apex of fore-
tibiai with a tuft of long, brown, hair-like scales, second tarsal
clothed with fairly long scales, tarsi three to five pale; ungues
of forelegs simple, rectangular, mid apparently the same.
9. Similar to male Scales on the costa, subcostal and first
longitudinal veins duskv as in male. Fore tibipe normal.
Length: ^, 1-75; 9, 1-5 mm.
Hab.— Q.: Cairns (F. H. Taylor).
Described from one male and two female specimens, bred from
larvae. The black abdomen, venation, and fore-tibiaa of the male
render this species distinct from its Australian congeners. The
lengths are only approximately correct, as the specimens are
doubled up.
Uranot^nia propria Taylor.
Trans. Ent. Soc. London, 1914, Ft. iv., p.704.
Q, Similar to male. First fork-cell shorter and narrower than
second; stem of former nearly twice the length of its cell, stem
of latter slightly longer than cell, anterior basal cross-vein longer
than anterior cross-vein, and once and one-half its^ own length
from it. Legs normal.
Hab. Q.: Cairns (F. H. Taylor).
This species was, previously, only known from the male.
URANOTiENIA ANTENNALIS, Sp.n.
Q. Head with pale scales in the centre, blackish elsewhere;
eyes black; antenna? brown, basal lobes yellowish, first segment
BY FRANK H. TAYLOR. 841
very long, about twice the length of the second, its base pale:
palpi blackish.
Thorax brown, with brown scales and black bristles, and a
short, narroM% pre-alar, white line of scales; scutellnm with black
scales; pleuriB brown, with white scales.
Abdomen covered with black scales; venter brown-scaled.
Legs dusky-brown; femora pale beneath, apical tarsi pale.
Wings : first fork-cell shorter than second, stem of first fork-
cell about twice and one-half the length of the cell; stem of the
second slightly longer than the cell; anterior basal cross- vein
longer than anterior cross-vein, and twice its own length from
it; scales on the costa, subcostal, and first long vein dusky, paler
elsewhere.
Length 175 mm.
Hab. — Q.: Cairns (F. H. Taylor).
Described from two specimens, bred from larvae. The first
joint of the antennae, and the venation separate this species from
other described Australian species.
IjRANOTiENiA HiLLi, sp.n. ( Plate Ixxxiii., fig. 12;
(J, Head covered with brown, flat scales; antennae pale, nodes
brown; eyes black and silvery; palpi brown, slightly longer than
usually found in the genus.
Thorax covered with dusky-brown scales, prothoracic lobes
prominent, dark-scaled; scutellum dark, denuded of scales; pleurae
yellowish-brown.
Abdomen with black scales; venter pale-scaled.
Legs dark brown; femora pale beneath.
Wings : fork-cells the same length, base of the second nearer
the base of the wing; stem of the first fork-cell slightly shorter
than its cell, stem of second not quite one-third shorter than its
cell; anterior basal cross-vein longer than the anterior cross-vein
and about once and one-half its own length distant from it.
Length 2 mm.
Hab.—^. Territory: Darwin (G. F. Hill; No.321).
Described from two specimens. The length of the first fork-
cell is relatively longer in comparison with the second than is
842 AUSTRALIAN CULICID^, iv.,
found in Uranotceniaj but I prefer to regard this species as
belonging to that genus.
Mr. Hill notes "larvae in crab-holes in mangrove-swamps." It
affords me much pleasure to dedicate this species to its discoverer.
Paratype in Coll. Hill.
HoDGEsiA CAiRNSENSis, sp.n. (Plate Ixxxiii., figs. 13, 14).
Head with black and bluish scales. Thorax orange, witii pro-
minent black markings. Abdomen l)lack-scaled, fifth segment
apically white, the second to sixth segments laterally white.
Legs unhanded.
9. Head with black scales, with a triangular patch of bluish-
white scales on the occiput and on the sides towards th^ base;
eyes, palpi, and proboscis black; antennae black, first joint long,
ba.sal half yellowish.
Thorax orange, with a prominent black spot above the wing-
roots, and a broad, median, brown stripe from the centre to the
posterior margin of the scutellum, sparsely covered with sliort,
hair-like scales: scutellum pale on the sides, covered with small,
black scales; prothoracic lobes with pale, flat scales.
Abdomen with black scales, first segment paler, segments two
to six with lateral, white patches, fifth with a broad, white-
scaled, apical band, apex bristly; venter pale-scaled, apical seg-
ment dark.
Wings considerably longer than abdomen, black-scaled; first
fork-cell longer and narrower than second, base of the latter a
little nearer the base of tlie wing: stem of the first fork-cell
about three-quarters the length of the cell, stem of second about
two thirds of its cell; base of second long vein carried well beyond
the transverse vein; halteres pale, with black knobs.
Legs black; femora wdth the basal half above, and the under-
surface, creamy; ungues very small, equal and simple.
Length, 1 -5 mm.
Hab.-Q.: Cairns (F. H. Taylor).
Described from a long series taken in shady situations, on the
edges of swamps, mainly sheltering in tree-holes, It is readily
distinguished from //. trianynlata Taylor, by its thorax and
abdominal bandiiiii'.
BY FRANK H. TAYLOR. 843
HoDGESiA TRIANGULATA Taylor. (Plate Ixxxiii., fig. 15).
Trans. Ent. Soc. London, 1914, Pt. i., p. 204; Proc. Linn. Soc.
N. S. Wales, 1915, xl, p.l84.
Hah.-Q.: Cairns (F. H. Taylor).
Fresh specimens show that segments two, three, five, and six
have white, lateral spots, and that the venter is white-scaled.
This species is wrongly recorded as Hodyesia trlmaculatus in
the Zoological Record, li., Insecta, xii., p. 277, 1914(1916).
EXPLANATION OF PLATES LXXX. -LXXXIIL
Plate Ixxx.
Fig.l. — Anopheles {Myzorliynchu-i) harhirostri--^ var. haucrofti ((lile.s); head
of male.
Fig. 2. — Toxorhynchifes speciosus (8kiise); head of male.
Fig.3. — Armiyeres hreinli Taylor; genitalia of male.
Plate Ixxxi.
Fig. 4. — Grnhhamia theohaldi Taylor; wing.
Fig. 5. — CiiUcada icUsoni, sp.n. ; head of male.
Fig. 6. — CuUcada wilsoni, sp.n.; genitalia of male.
Fig. 7. — Cidkada wilsoni, sp.n.; wing of female.
Plate Ixxxii.
Fig. 8, — Lophoceratomyia cairnMenms, sp.n.; head of male, showing an-
tennary organs.
Fig. 9. — Lophoceratomyia cairnsensis, sp.n.; wing.
Fig. 10. — Molpemyia pi'iestJeii Taylor; wing of female.
Fig. 11. — Uranotcenia tibialis, sp.n.; wing.
Plate Ix^xiii.
Fig. 12. — Uranotoinia hilli, sp.n.; wing.
Fig. 18. — Hodgesia cairnsensis, sp.n.; wing.
Fig. 14. — Hodgesia cairnsensis, sp.n.; portion of wing under high power.
Fig. 15. — Hodgesia triangidata, sp.n.; wing.
844
N(yrE ON THE TEMPERATURE OF ECHIDNA
ACULEATA.
By H. 8. Halcro Wardlaw, D.Sc, Linnean Macleay Fellow
OF THE Society in Physiology.
(From the Physiological Laboratory of the University of Sydney.)
(Witli two Text-figures).
In a previous communication, the present writer has recorded
a number of observations of the rectal temperature of Echidna
ac.nl eata (Wardlaw, 1915). The observations were made at
different times of the day, and during different seasons of the
year. As the temperatures showed considerable variations, even
outside of th6 periods during which the animals were hibernating,
some difficulty was experienced in arriving at an estimate of
their normal waking temperature. Average values calculated
from results varying like those obtained have no precise meaning.
Yet to give the range of variation alone is hardly sufficient : a
central value is necessary as a point of departure.
These results, therefore, have been submitted to a further ex-
amination, in which the graphic method of statistical analysis,
due to Galton, has been applied in the manner described in a
former paper (Wardlaw, 1917). Tn this w^ay, it has been shown
that the observations of temperature are not distributed at
random over their whole range of variation, but occur more fre-
quently in the vicinity of certain values, and that the tempera-
ture of this animal has certain definite most probable values for
different times of the day at different seasons of the year.
In the accompanying diagrams (Text-figs. 1 and 2), the fre-
quency of occurrence of different temperatures of Echidna, outside
of the periods of hibernation, have been plotted. Temperatures
of the animals are measured along the abscissae. The relative
BY H. S. HALCRO WARDLAW.
845
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Text-lig.l. (See p. 849).
65
846
TEMPERATURE OF ECHIDNA ACULEATA,
;i C* ^» ^ r/
O H f\ rV r^
S - — ^-^y
h'^
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1
i
i
5*
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Text -fig. 2. (See p. 849).
BY H. S. HALCRO WARDLAAV. 847
frequencies of occurrence of the vari<ju8 temperatures are pro-
}»orlional to the ordinates. Autunni and winter temperatures
are given in Text-fig. 1; spring and summer temperatures are
given in Text-fig. 2. The morning and aftei-noon temperatures
are plotted separately in each diagram. The mean air-temper-
atures, corresponding to each range of I'C. of the animal's tem-
peratures, are also shown. The mnnhers of results represented
by each of the four pairs of curves aie as follows; autumn-winter
period, morning, 188, afternoon, 242 ; spring-sununer period,
morning, 98, afternoon, 102. Ah these numbers are different,
the ordinates of the different curves are not directly comparable
with each other. This does not matter, however, as only the
different ordinates of the same curve require to be compared.
Curve A, Text-fig. 1, shows the ogive of the morning temper-
atures of Echidna during the autumn-winter period. Each
ordinate is proportional to the number of observations occurring
at and below the corresponding temperature. Curve A' is the
derived or frequency-curve; each ordinate is proportional to the
number of observations occurring at the corresponding temper-
ature. It will be seen that the maximum of this frequency-curve
lies at 29'7 C; this is the most probable morning-temperature of
Echidna during the autunni-winter period. It will also be noticed
that the curve A' flattens out and becomes almost horizontal
below a t{!m})erature of about 27 "GX'. This portion of the curve
continues tlown to about 8*0 C, but has not all been shown. The
curve indicates that, below 27'6 C, Echidna allows its tenqjer-
ature to vary at random, and displays no tendency to bring it
towards a definite value. By the time its b(xly-tenqjerature has
fallen to 27 '5C., Echidna has thus become a completely poikilo
thermal animal.
The line A" joins the mean air-temperatures corresponding to
the different body- temperatures of Echidna, and is marked by
circles.
Curxe B, Textlig. 1, is the ogive of the afternoon-temperatures
of Echidna during the autumn- winter period. Curve B' is the
corresponding frequency-curve. Its maximum occurs at 32*3°C.,
84:8 TEMPERATURE OF ^:CHiD^~A ACULEATA,
and tliis is the most probable afteinoon-leinperature of Echidna
(hiring the pcnod in question. I'he line B' joins the air-teniper-
atiires corresponding to the diii'crent temperatures of Echidna,
and is indicated by dots.
Curve C, Text-fig. 2, is the ogive of the morning-temperatures
of Echidna during the spring sunnner period. C is the corres-
ponding fre([uency-curve. Its maximum lies at .'^OO C, and this
is the most piobable morning-temperature of Echidna during
spring and summer. The line 0" joins the average air-temper-
atiu'es as before, and is indicated by circles.
Curve D, Text-fig. 2, is the (jgive of the afternoon-temperatures
of Echidna during the spring-sunnner period. D' is the corres-
})onding frequency-curve. The curve, it will be noticed, does
not fall after rising to its maximum, but remains constant. The
maximal value is reached at a temperature of 32 6°C. Above
this temperature the frequency-curve is horizontal, and tlie tem-
perature of Echidna varies at random. These facts indicate that
there is an upper, as well as a lowei-, limit of temperature beyond
which the temperature-regulating mechanism breaks down, and
Echidna behaves as a poikilothermal animal. This mechanism
is only etfective while the animal's body-temperature lies between
about 27 "6 C. and -Vi'G'C, that is, over a range of variation of
about 5 C.
The line D' joins the average air-temperatures corresponding
to the dili'erent aftei'noon-tcm}»eratures of Echidna during the
spring-summer period.
The four curves show, that most probable temperatures of
Echidna are 2*6'C. lower in tlie morning than in the afternoon
during both of the periods in which the observations were
made.
The most }n-obal)le temperatures of the spring-summer period
were very slightly (0'3'C.) higher than the corresponding tem-
peratures of the autumn-winter period. The average air-temper-
atures were about 5"C. higher in the former than in the latter
period.
by h. s. halceo wardlaw. 849
Summary.
The iiiufit piubablc tcinpcraturt's of Echidna are: in the morn-
ing, 29 TX'. during autunni and winter; 300'C. during spring
and summer. In the afternoon, 32-3°C. during autunni and
winter; 32'6°C. during spring and summer.
The temperature-regulating meclianism of Echiihia is only
ertecti\e wliile the body-temperature lies betweeji •27'6°C, and
32 fi"!/. Outside of these limits, Echidna behaves as a poikilo-
ihermal animal.
KEFERENCE.S.
(iALTON. — Natural Inheritance. London. 1880.
Wakdlaw.— Pioc. Linn. Sou. N. 8. Wales, 19L3, xl., p.231,
Ibid., 1917, xlii.. p.Slo.
LEGENDS OF TEXT-FK^UKES.
Text-%.1 (p. 845).
Kolalixc frequency of (occurrence of ditFerent autunni and w inter temper-
atures of Echidna. Ahscissie, Ijodj'-teinperatiu-es of Echidna.
Ordinates of curves A and !^ are proportional to inu!d)ers of ob-
servations at and below the corresponding temperatures. Ordinates
of curves A' and B' are proportional to numbers of observations at
the corresponding temperatures. The points joined by the lines A "
and B" indicate the average air-temperatures corresponding to each
degree range of body-temperature. Letters A, A', A" refer to
morning-temperatures; letteis B, B', J5" to afternoon-temperatures.
Text-tig. 2 (p 847).
Rclalivc f'requenc,y of occurrence of different spring and summer temper-
atures of Echidna. Abscissje, body-temperatures of Echidna.
Ordinates of curves C and IJ are proportional to numbers of ob-
servations at and below corresponding temperatures. Ordinates of
curves C and D' are proportional to numbers of ol)servations oc-
curring at the corresponding temperatui-es. The points joined by
the lines C" and D" indicate the average air-temperatures corres-
ponding to each degree range of body-temperature. T-ietters C, C",
C" refer to morning-temperatures: letters 1), D', D" to afternoon-
temperatures.
850
THE OCCURRENCE OF METHYL L^VO-INO.STTOL IN
AN AUSTRALIAN POISONOUS PLANT.
Bv .Iamivs M. Pktkik, D.Sc, F.I.C, Linxkan Maclhay Fellow
OF THE Society in Biochemistky.
(With two Text-figures.)
(From the P/njsioloyicaf Lcihoralory of the University of Sydney.)
Heterodendron olecefolmm Desf., (Family Sapindace?e) is a large
shrub growing on the plains of the Western and Northern In-
terior of New South Wales, and is also found in all the other
States of Australia. It has been described as a valuable forage-
plant in the stock country because of its drought resistant
character.
Some time ago this plant was suspected to be the cause (jf
certain fatalities among cattle and horses, and a sample was
received by the writer U)v chemical in\ estigation. It was found,
when examined, to be a strongly cyanogenetic plant.
The main object of the extensive investigation cariied out on
this plant was, therefore, the attempt to isolate the cyanogenetic
principle and to study its pro[)erties. During the course of the
work there was separated a reuiarkable and intei'esting comp(jund
of hcvo-inositol, and this })aper will be confined to an account of
the method by which it was obtained, and a general description
of its characteristic features.
The material for the in\ estigation was c»^llected near Coonamble
by Stock Inspector E. W. Procter, and forwarded to the Uni-
versity through the kindness of the Chief Inspector of Stock, Mr.
S. T. D. Symons, M.R.C.V.S., to whom the author expresses his
indebtedness and thanks.
BY .lAMKS M. PETRIE. 851
Method of Extraction.
The plants were air-fhned, and then tlie leaves were separated,
crushed, and afterwaids fhu^ly powdered. Of this air-dried leaf-
powder, 2' 5 kilos, corresponding- to about 1 1 -5 kilos, of fresh
leaves, were extracted with f<()% alcohol. The extraction was
continued until almost complete; and the alcoholic fluid, measur-
ing 31 litres, was distilled under diminished pressure, and at a
temperature not exceeding .'^)0 C. The distillation was continued
till the residue was free from alcohol, and C(mcentrated to a thick
syrup. This viscous residue was treated with warm water,
filtered through calico, and the residue washed until no more \vas
dissoh ed. There was collected on the cloth-filter a black, sticky
mass, consisting mostl}' of chlorophyll and resins, and weighing,
when dried, 200gms., or 8% of the air-dried leaves. The aqueous
filtrate, aftei* standing in tall cylinders for a few days, deposited
a considerable quantity of brown, amoi-phous resin. The lattei'
was removed and washed with cold watei*.
The opaque, dark brown solution was next purified by the
addition (1) of an equal volume of 10% lead acetate solution,
and (2) of an excess of basic lead acetate, after removing the
previous bulky deposit. These very voluminous yellow precipi-
tates were separated by spinning in the centrifuge, and were
washed in the same manner with cold water, and reserved. Next,
the solution was made free from lead by saturation with hydrogen
sulphide, and the precipitate removed and well washed.
The aqueous solution, measuring 18 litres, was now concen-
trated by distillation at a low temperature, to a volume of about
2 litres. At this stage, the solution was thoroughly extracted
by shaking out with ether, and in this wa}^ the free acetic acid
was removed.
Crystallisation.
Two volumes of strong alcohol were then added to the solution,
and, on standing to settle, a dark syrupy deposit formed, from
which the solution was decanted. The latter was then concen-
trated by evaporation to a viscous mass, and on the addition of
95% alcohol to this residue, an insoluble viscous substance re-
852 METHYL L.EV0-IN0SITOL IN A POISONOUS PLANT,
mained. The alcoholic fluid was allowed to stand for some days,
when there was formed a considerable quantity of clear glassy
crystals. On evaporation at 35°C., and again treating with
alcohol, a further separation of crystals took place. This evapo-
ration and treatment with 95% alcohol was repeated three times,
the mass of crystals was drained on a Buchner funnel, and
washed with alcohol. The whole was then recrystallised from
dilute alcohol, and the crystals dried. The weight of this sub-
stance, with the addition of a smaller amount obtained in the sub-
sequent treatment of the solution, was 15gms., equivalent to —
0*65 per cent, of the dried (at 100°) leaves.
0*60 per cent, of the air-dried material.
Purificatio7i of the crystals. — The whole of the substance was
now dissolved in water, in which it was exceedingly soluble, and
alcohol was carefully added to the point of incipient precipitation.
On cooling the solution to 0°C., the substance slowly separated
in fine transparent crystals. During this separation, the super-
fluid was decanted at intervals, till finally there were obtained
twenty separate fractions. The first, tenth, and twentieth frac-
tions, when dried, gave melting-points between 188° and 189'5°C.
(uncorrected), thus proving the presence of a single substance
onl^^ The combined fractions were recrystallised three times,
and dried in a desiccator.
Properties of the Crystals.
The following tests are described in the order in which they
were performed, and show the method by which the constitution
of the compound was gradually elucidated.
Preliminary tests. — (1) On fermentation with a very active
preparation of emulsin, no hydrocyanic acid was evolved. The
compound is, therefore, not the active principle of the plant.
(2) When heated, the substance melted, charred, and burned
entirely away without residue.
It consisted of the elements carbon, hydrogen, and oxygen
only. The crystals possessed a very sweet taste, and were ex-
(;essi\'ely soluble in water; from which facts it may be inferred
that the compound contains a nunlbei' of hydroxyl groups. When
BY JAMES M. PETRIE. 853
examined by the microscope, the crystal form was similar to that
of cane sugar.
Molisch's reaction with sulphuric acid and </.-naphthol or
thymol gave no colour, and Fehling's solution was not reduced.
The compound is, therefore, not a sugar.
(3) The melting-point, taste, and crystalline form are identical
with those of the hexahydric alcohol, dulcitol; the latter, how-
ever, yields mucic acid when oxidised with nitric acid, whereas
no mucic acid could be obtained from this substance.
(4) By treatment of the solution with phenylhydrazin acetate,
no hydrazone or osazone could be obtained.
Silver nitrate in ammoniacal solution gave no precipitate in
the cold; but, on warming, the solution slowly darkened with
precipitation of the silver.
On boiling witli dilute acids and alkalies, and subsequently
recovering the compound, no apparent change in its properties
was noted.
{h) Quantitative determinations. — The crystals, which had been
formed in dilute alcoholic solution, and dried in a desiccator at the
oi'dinary temperature, were heated at 110°C. for two hours, but
showed no decrease in weight, and then at 150°C. for 30 minutes,
with a similar result. The substance, therefore, contains no
water of crystallisation.
The solubility showed that 1 gm. required l"9c.c. of water at
2rC., or 53%.
(6) The melting-point, as carefully determined on a standard
Anschiitz thermometer wholly immersed, was 190°C.
(7) Ultimate analysis of the substance yielded the following-
results : —
0-1262gm. gave... 0-083 H.,0 and 0-1983 CO..
Equivalent to ... 7-3% H and 42-9% C.
C-Hi^O,. requires 7-2% H and 43-3% C.
This formula, which conforms most closely to the figures ob-
tained foi' the substance, is possessed by the meth^d-hexoses,
simple glucosides, and certain derivatives of benzene.
(8) A determination was made of the number of methoxy
66
854 METHYL LiEVO-INOSITOL IN A POISONOUS PLANT,
groups present in tlie molecule, by Perkin's modification of the
method of Zeisel. By heating in pure liydriodic acid, methyl
iodide distilled over into silver nitrate solution.
0*2 gm. gave ... 0-2315 gm. silver iodide.
Representing . . . 00306 gm. CH...O.
C.Ji^ ^0^. requires 0'0310 gm. for one CH.jO group.
Tlie substance can, therefore, be represented by the foi-mula
CH.. -C^^H^ ^O,;, and the residue remaining in the Zeisel apparatus
should possess the formula C^HjoOg. In order to recover this
substance for examination, the solution, from which the methyl
iodide had been driven off, was lieated on the water-bath to
remove hydriodic acid, and then ev^aporated to dryness. By
extracting this residue with alcohol and cooling to 0°C., white
crystals were recovered.
Crystals dried in desiccator and weighed... OvlGOgm.
Crystals dried at llO^'C. and weighed ... 0"160gm.
C-H^^O,. — 0-2 gm. requires ... ... O'lSGgm.
The crystals are, therefore, without water of crystalli.sation.
The low yield may be accounted for by partial decomposition
during the boiling with hydriodic acid, as the odour of benzene
and phenol was distinctl)^ detected.
(9) Pro])erfieti of the demethylatfd substance. — After three crys-
tallisations, the substance gave a melting-point of 238°C., and
charred at 239X\, carefully determined on an Anschiitz standard
thermometer with the column submerged.
This substance also gave a negative result with Molisch's test,
proving the absence of all open-chain carbohydrates; and since
benzene and phenol were identified as decomposition-products of
the ester, the possible cyclic compounds may next be considered.
The formula C,;Hj^Og is contained in the inositol ring, for
the identification of which the following reactions are specific: —
{a) Scherer's test gave positive reactions with this substance,
and likewise with the original methyl derivative. When a little
of the solution is evaporated with nitric acid, neutralised with
ammonia, and calcium or barium chloride added, a brilliant rose-
red colour appears. (Liebig's Annalen der Chemie u. Pharm. 81,
1852, 375).
BY JAMES M. PKTRIE. 855
(b) Gallois' test gave positive results witli both substances.
When the crystals are treated with mercuric nitrate, a yellow
precipitate first forms, which, on evaporating to dryness, gives a
deep red colour; on cooling, this colour slowly disappears, to
return on being reheated. (Fres'. Zeitschrift fur anal. Cheniie,
iv., 1865, 264).
These two charactei'istic reactions are due to the oxidation of
inositol to a (luinonoid substance known as rhodizonic acid, whose
salts with calcium, barium, and mercury, possess the bright
colours described.
The reduced substance is thus proved to be one of the inositols,
and the original compound isolated from the plant-extract is its
methyl ester.
(10) Optical propfrfifis. — A polarimetric determination of the
two substances was made with a Schmidt and Haensch polari-
meter reading to one-hundredth of a degree.
Methyl inositol 0"5gm. was dissolved in 10 c.c. of distilled
water at 16°C., and a Ijevo rotation w^as recorded of - 4*01 in a
1 dcm. tube.
The specific rotatory power [o]^- = - 80-2; [MJ^J- = - 155-0.
The solution was boiled for two minutes, and after cooling to
16°C., was again read in the polarimeter. No change was ob-
served, such as is due to mutarotation among the hexoses.
Inositol, the de-methylated compound, 0"0741gm. was dissolved
in 10 c.c. of water at 16°C., and showed a Ifevo rotation of
- 0'48° in a 1 dcm. tube.
Specific rotatory power [a]^' = - 64-8; [MJ^J' = - 116'7.
(11) Hydration. — The Isevo-inositol was obtained by crystallisa-
tion from cold aqueous alcohol, and contained no water of crys-
tallisation. When crystallised from water, it was also obtained
in anhydrous crystals.
Now Maquenne and Tanret have described some important
differences with regard to the water of crystallisation in the
isomeric inositols."^ They found that —
* Reeherches sur I'inosite, Maquenne — Annales de ehemie et de physique,
xii., 1887, 94; Comptes lendus, ex., 1890, 87.
856 METHYL L.EVO-INOSITOL IN A POISONOUS PLANT,
Inactive-inositol
from dil. alcohol, or M-ater under 50°C., gave ... crysts. with 2H2O.
from dil. aleoliol, or water above 50°C., gave ... aiihj'drous cry.stis.
Dextro-inositol
from dil. alcohol, or cold water, gave ... ... anhydrous crj'sts.
from cold water seeded with hydrate crysts. , gave crystal with 2H.,0.
from hot water, gave ... ... ... ... crysts. with 2H..0.
L;evo-inositol
from cold water, gave ... ... ... ... crysts. with 'iH.^O.
from hot water, gave ... ... ... ... crysts. with 'iH.^O.
from dil. alcohol, gave ... ... ... ... anhydrous crj-sts.
Dl-inositol
from cold water, gave ... ... ... ... anhydrous crysts.
It is to be pointed out that the inactive and dextro forms give
reverse results with the same treatment. Maquenne, in com-
paring the dextro- and l^evo-isomers, could always obtain the
former from cold water in anhydrous crystals, but was quite
unable to obtain the same with the hi^vo form.
The following results were obtained with the k«vo-inositol
from Heterodendron, and are of interest when compared with the
figures in the previous table.
gm.
1 . Heated at 1 1 0°C. for 1 hour 0 'oSOo
2. Dissolved in cold water, dried in desice. at 25^C. for 2 daj^s .. 0'5522
,, ,, ,, 25°C. for 1 day ... O'ooll
25°C. for 1 day ... 0*5o08
3. Dissolved in water at 70°C., crystd. at 70'C., dried in desice.
at 25°C. for 2 days 0'5678
at 2o°C. for 1 day 0-5650
at 15°C. for 3 days 0-5530
at 15°C. for 3 days 0-5512
4. Heated at 100°C. for 3 hours ..." 0*5508
5. Diss, in water, dried in the open at \ai°C. ... ... ... 0'5508
Inositol ■2H2O requires ... ... ... ... ... 0-5660
The crystallisation from cold water, therefore, left anhydrous
inositol when kept over sulphuric acid, or dried in the open.
When cr3'stallised at 70'C., and subsequently kept over
sulphuric acid for two days, the crystals contained an equivalent
of two molecules of water; but since this water was graduall)'
lost at the ordinary temperature standing over sulphuric acid, or
BY JAMES M. PETRIE. 857
ill the open, appai'ently it was uul coinl)iiied as hydrate in the
crystals.
Tlie liy(h-ates <)l)tained by Ma<|ueiiiie and others were only
decomposed at 100°C. If the hydrate exists in the above case,
it is decomposed by drying at the ordinaiy temperature.
The l;*^vo-inosit()] of this research, therefore, was obtained in
anhydrous crystals only. The hydrates prepared Ijy the French
chemists could not be obtained.
The substance isolated from HebrorhDidron olcrvfoliiDit is thus
})roved to be the methyl ester of l(e.vo-rof(Uor}j inositol.
(12) Crystal Form, — -The outward structure and measurements
of the crystal forms are intimately related to the internal struc-
ture of the isomeric molecules, and, therefore, form an essential
part in the elucidation of the individual members of a group.
The methyl inositols apparently have never been examined by
crystal lographers, and indeed, as far as the author can ascertain,
only inactive inositol crystals have been examined by the goni-
ometer.
The goniometric determinations of the crystal forms of 1. -methyl
inositol were kindly made by Dr. C. Anderson, Mineralogist to
the Australian Museum, and are here included.
CrYvStal Measurements of Methyl L.Evo-ixosrroi..
By Charles Anderson, M.A., D.Sc.
The crystals are small, the largest being about 2 mm. in length.
They belong to the orthorhorabic system and are very uniform
in development and habit: of the five crystals measured, fuur
sliow the forms a(lOO), 6(0 10), m(llO), ^(011), wdiile one has, in
addition, one face of the form /'(lOl), and they are all tabular
on a. The faces are by no means perfect, being interrupted and
wavy, the signals are only fair, and, consequently, the measure-
ments are not m close agreement. The crystals were measured
on a two-circle goniometer, the reducing lens being used.
858 METHYL L^VO-INOSITOL IN A POlSONOtTS PLANT,
m"
in
a
Text-tig. 1.
The axial ratios were talculated from the following angles.
<^
P
i No. of
I obs.
Form.
Mean.
Limits.
Mean.
Limits.
wllO
52°44'
5Vm'—o3°2T
39°26'
39°0'— 39°49'
' 17
, 8
The elements deduced from these angles are
a:b:c = 0-7609:l:0-8224.
BY JAMES M. PKTRIE.
859
Forms and angles.
Measured.
Calculated.
Form.
Symbol.
<t>
P
(ji j p
ft
100
S9°o4'
90°4'
WO'
900'
1,
010
0°o'
90°22'
0°0'
go'^o'
nt
110
o2°44'
90°2'
90°0'
q
Oil
OW
39°26'
0°0'
r
101
88°33'
47°48'
90°0'
47^13'
The Inositols.
Position of the group. — The relative positii^ii of the group, and
the mode of occurrence of its members in nature, are of consider-
able interest to the biochemist, especially since the discovery of
"phytin" in plants by Paladin, in 1895.
The basis of inositol is the hexamethylene ring (CH..),..
Hexamethylene, C„H^^ (Text-fig. 1 ), occurs only in the hydro-
carbons of the petroleum of Russia, Galicia, Baku, East Indies,
and California, in the fraction boiling about 80°C. It has not
been detected in plants or animals.
By the substitution of hydroxy! groups (OH) in the hexametliv-
lene ring, the following series of compounds is obtained: —
(OH), (OH).^, (OH).. — synthetic compounds only.
(OH)^ — betite, isolated from beet sugar residues.
(OH)- — quercite, in oak and other plants.
(OH),; — inosite, in many animals and plants.
Inosite, or inositol, has, therefore, the constitution ^)i a liexa-
hydroxy hexamethylene C,.H,. ' (OH),., and although its formula
may be written C\;H^^O,., it is nevertheless, in its relationships,
far removed from the carbohydrates.
Tha couistituJion of Inositol. — The configuration of the inositol
molecule, or, in other words, the arrangement of its at<jms in
space, admits of eight different geometrical groupings resultinij
in eight possible isomeric foi-ms. When these and their mirror-
images are built up in models, it is found that seven of the forms
may be superimposed on, and, therefore, coincide with, their
860
METHYL L^VO INOSITOL IN A POISONOUS PLANT,
mirror-images. Tliis condition, arising from a certain degree of
molecular symmetry, is accompanied by inactivity towards polar-
ised light. Tliese seven forms having their asymmetric carbon
atoms internally compensated, are, therefore, optically inactive
in the polarimeter. One form alone is found to possess an
entirely asymmetric molecule, and, in consequence, this arrange-
ment can exist as active dextro and lt«vo compounds, and, in
addition, their (f/- or racemic inactive combination may also exist.
Constifution of the methyl psters. — As has just been stated, the
active forms of inositol are the result of one particular arrange-
ment of the hydroxy 1 groups. This arrangement, which may Ije
readily discovered in the models, is that where the six hydroxyl
groups occupy the positions 1, 2, 4, on each side of the ring.
This form (Fig. 2) and its mirror-image (Fig. 3) constitute the
dextro- and l?evo-inositols.
Fig. 1. Fig.2, Fig.3. Fig. 4.
H H OH OCH,
and
inositol.
Text-fig. 2.
methyl leevo-
inositol .
The methyl ester, C^H,(OH)- (OCH..^) is obtained by sub-
stitution of a methyl group in one of the hydroxyl groups, and
again from the models it can be proved that substitution in tlie
hydroxyl 1, 2, or 4 results in three possible and different com-
pounds being obtained. The first of these is represented by
Fig. 4. The corresponding three positions in the mirror-image
(Fig. 3) — which are identical with the alternate three positions
below^ in the other form (Fig. 2) — produce their optical anti-
BY JAMES M. PETRIE. 861
podes. The possible existence of three dextro- and three Ifcvo-
inethyl esters is thus deteimined. It now remains to ascertain
(a) whether tlie few inositol esters wiiicli have been is(jlated up
to the present time, represent the same stereo-isonier, (6) whether
the compound obtained h'om Ifeterodendroii is identical with any
of the others, t)r represents the second or third isomer.
Occurrence in iiafure. — It has been previously stated that
theoretically there can exist ten stereo-isomeric forms of inositol
as a maximum possible number: — -
7 inactive by internal compensation (meso),
2 active, dextro and Ipevo,
1 inactive, racemic or dl,
10 isomeric forms.
Only the inactive inositols have yet been found existing in the
free state in nature, but esters of both active and inactive
inositols are found.
{a) Inactive inositol. — This form, widely distributed in animal
tissues, and already well known to physiologists, possesses the
formula of one or other of the seven internally compensated
molecules, and it is worthy of note that this is the only form
found in the animal kingdom. It has always been referred to in
physiological chemistry as one substance, with definite and con-
stant general properties. But no one so far has troubled to ex-
amine minutely, material from widely different sources or organs,
as to the particular properties which would differentiate these
inactive isomers, such as crystallographic measurements, or optical
properties.
It was discovered in animals, in 1850, by Scherer, in extracts
of flesh,* and in plants, six years later, by Vohl.f This author
was examining the sap of unripe pods of Fhaseolus vulgaris, and
after completely fermenting the sugars, and distilling off the
alcohol, he found that the solution still possessed a very sweet
taste. He then separated a manna-like substance,
* Liebig's Annalen der Chemie und Pharni., Ixxiii.
t Ibid., xcix., 1856, 125.
which he called
, 1850, 322.
R A R
862 METHYL LiKVO INOSITOL IN A POISONOUS PLANT,
phaseo-maniiite. In the following year, the same chemitst proved
the itlentity of his maniiite with Scherer's inosite from animals.
The occurrence has been recorded of three other substances,
which are believed to be isomeric with inactive inositol, since
they, although differing widely in crystalline form, melting-point,
and solubility, possess the same general characters. These are
the scyllite of Stfedeler, from certain elasmobranch fishes, the
quercinite of Delachanel, from the oak, and the cocositol of
Mueller, from the cocoanut.
The compounds of inactive inositol which have been found in
nature are : —
Bornesite — the methyl ester, obtained from caoutchouc.
Dambonite — the dimethyl ester, obtained from caoutchouc.
Phytin — the phosphate ester, an essential constituent of all
plants and animals.
(h) Dextro-inositol occurs only as the methyl ester, pinite. It
was discovered by Berthelot,"^ in 1<S56, in the resins from Oregon
pine, and has since been found in senna leaves, and caoutchouc.
(c) Lcevo-iiiositol is likewise found only as the methyl ester,
and the following is a complete record of its occurrence: —
1. In quebracho bark, Aspidosppnna quebrarho (Apocynacete),
discovered by Tanret, of Paris, in 1889,t and named by him
quebrachite.
2. In Hevea brasUiensis (Euphorbiaceae), in the aqueous solu-
tions of the latex after coagulation of the rubber,; and in Para
rubber. §
3. In Grevillea robusta (Proteacea3).|| It is associated in the
lea\es with the glucoside arbutin.
4. In Heterodendron o/ece/olimn (Sapindacete), this paper.
(d) Racemic inositol was discovered in mistletoe by Tanret in
* Annales de chiniie et de physique, xlvi., 1856, 60.
1 Coniptes rendus de I'Aead. des Sciences, cix., 1889, 908.
+ de Jong, 1906, thro. Wehmer's "Die Pflanzenstoffe."'
§ Pickles and Whitfield, Proe. Cheni. 8oc. Lend., 1911, 54.
Bouiquelot et Fichtenholz, Journ. pharni. et de chirnie, Paris, vi.,
1912, 346.
BY JAMES M. PKTRIE.
8H3
1907.* It was found to exist free in the berries and leaves, and
associated with much (uieso) inactive inositol, but no active
isomers were present.
The Active Forms of Inositol.
The following Table shows the physical constants Ijy which
the active forms and their compounds are identified.
melting-point.
spec. rot.
power.
(1) Dtxtro- methyl ester.
From Pinus (Maquenne)
186°C.
+ 65 '5
Pinus (Combes) ...
18(5 -.1
657
caoutchouc (Combes) ...
187
66
caoutchouc (Girard
181
64-7
{'!) L<vro-methyI ef<ter.
Fiom Quebracho (Tanret)
190X\
-80
Rubber (Pickles and Whittield)
191-2
80
Gievillea (Bourquelot) ...
190
80-3
Heterodendron (this paper)
190
80 2
(8) Dtxtro- 1 11 osito/.
From pinite (Maquenne)
247°C.
+ 65
pinite (Berthelot)
•245
caoutchouc (Combes) ...
240
67 6
caoutchouc ( Gi rard )
28.-)
64-7
(4) Litro-inositol .
From Quebraclio (Tanret)
288 C.
- (>5
Quebracho (Maquenne)
247
65
Rubber (Pickles and Whithi
-Id)
237
—
Grevillea (Bourquelot) ...
247
65
Heterodendron (this paper)
238
64-8
Table showing the amount of metliyl Ipevo-inositol obtained
from the difterent sources: —
Aspidosperma quebracho ... 0*1% of dried leaves,
Hevea hrasiliensU rubber ... (2 "5% of the rubber).
GrerifJea rofmsfa ... ... ... 0*4% of dried leaves.
Hettrodendroii ole.<i'folinm ... 0"65% of dried leaves.
From the first Table, it is apparent that the It^vo-methyl
inositols (2) from the four difierent sources, have identical melt-
ing-points and specific rotatory powers, and therefore, in all
probability, represent one only of the three possible stereo-
isomers previously mentioned,
* Comptes rendus de I'Acad. des Sciences, cxlv., 1907, 1196.
864 METHYL L.EVO-INOSITOL IN A POISONOUS PLANT,
AV'heii converted to hevo-iiiositul (1), however, there would
a}>}>e;ir to be two groups of melting-point figures, one 10° higher
lljcin the other, but since there can be only one possible /-inositol,
this dili'erence nuist be otheiwise explained.
The Table also shows that while the dextro- and Itevo-inositols
(3 and 4) are optical antipodes of one another, their esters (1 and
2) are not. The optical properties especially ai'e so very diver-
gent that, in all probability, the methyl group occupies a different
position in the two compounds. Tlie compound isolated from
Heterodendron is, therefore, shown not to be an optical isomer
of Maquenne's pinite.
Biochemical Relationships and Significance.
(a) The chemical aspect. — Since the researches of Maquenne,
cited in the previous paragraphs, no subsequent w^ork has shown
any relationship between the inositols and the carbohydrates,
other than the sweet taste and the molecular formula common to
both. Perhaps one exception to this is found in Neuberg's iden-
tification of furfural among the products of decomposition, when
inositol is boiled with acids."^ Although furfural is also obtained
from the hexoses and heptoses in small amounts (about 0'2 /^), it
is characteristic of the pentose sugars. It must be also remem-
bered that the production of furfural is the basis of Molisch's
group-test for all carbohydrates, and with this reagent the
inositols gave negative results.
However, it seems probable from the results of many workers
that the hexamethylenes form a kind of stepping-stone between
the open chain compounds and the true benzene ring derivatives.
Ojjoi chain camps. Closed ring camps.
hexose sugars hexamethylene derics. Benzene derivs.
dulcitol (juercitols phenols
mannitol inositols
sorbitol
The hexamethylene derivatives are much more easily decom-
posed than the simple benzene compounds. In fact, it has been
* Biocheni. Zeitschrift, ix., 1908, 551.
BY .TAMKS M. PETRIE. 865
found byDrechsel that fungi and bacteria maybe grown in solu-
tions of the former, especially the oxidised forms (juercitol and
quinic acid. It has also been proved that when phenol is ex-
posed for some time to the alternating electric current, it is first
comerted to hexamethylene derivatives, before being finally
oxidised and split up into various fatty acids.*
Though there are many instances of the closing up of open
chain carbon compounds into ring compounds, such as the con-
version of citral into C3^mol, the cyclo-citrals, and terpenes;
geraniol into dipentene, etc.; yet no such conversions have been
accomplished between the carbohydrates and the inositols
Griffin and Nelson, in their researches on inositol and pinite,
tried by various methods to close up the hexose chain, and to
open out the inositol ring, but were entirely unsuccessful. t
{/)) TJiP biochemical aspect. — On the other hand, in favour of
the biochemical possibility, there exist the important observa-
tions of Neuberg: that quercitol and inositol may be reduced in
a few minutes to open chain carbohydrates (reducing Fehling's
solution, etc.) by the action of sunlight and a catalyser such as
uranium salt, also by the action of the alternating current.!
These processes, however, are all reverse reactions, resulting
in cleavage of the hexamethylene ring. Concerning the direct
synthesis — carbohydrate to inositol, we have no evidence at all,
and Maquenne had no experimental basis for his belief that the
alcohol mannitol was the source of inositol.
Rosenberger observed tlie appearance of inositol in post mortem
tissues, where previously no inositol existed; and he assumed the
pre-existence of an "inositogen" from which, by enzyme-action,
the inositol was formed. v^
The inactive inositol combines with inorganic phosphates, and
in this form exists as "phytin" in nearly all living organisms.
This substance is always accompanied by the enzyme phytase,
* Journ. fiir prakt. Chemie, xxxviii., 1888, 60.
t Journ. Amer. Chem. Soc, xxxvii., 1915, bjo'i.
X Biochem, Zeitschrift, xiii., 1908, 308.
§ Zeitschrift fur physiol. Chemie, Ivi., 1908, 373.
866 METHYL L.EVO-INOSITOL IN A POISONOUS PLANT,
which effects its cleavage and yields inositol again, in the free
state. Starkenstein's investigations show that the source of free
inositol in tissues is the pliytin, and that inositol is a decompo-
sition-product of the phosphoric acid metabolism in both plants
and animals."^ Indeed, much work has been done in elucidating
the conditions of this transformation on the side of the phosphoric
acid, but again, as to the inositol side, nothing is known.
The few definite observations concerning the part played in
metabolism, by inositol, are here summarised : —
1. In iin7'ipe seeds, inositol and quercitol accumulate just at
the time when the transport of carbohydrates to the fruit begins.
2. As the fruit 7'ipens, Vohl observed that inositol and quercitol
disappear, and are changed into "phytin" (not carbohydrate).
3. On the yerinination of the seeds, inositol again makes its
appearance, both when grown in the dark and in the light.
4. During the metabolism of the growing ptmit, inositol disap-
pears gradually with the rest of the reserve-substances.
Thus it comes in at the beginning and later passes out again,
without a clue to its precursors or katabolites.
AVhen fed to animals, or injected into the blood-stream, inositol
is about three-fourths decomposed, and the remainder may be
recovered from the urine unchanged. Mayer injected large doses
into rabbits, and obtained, from the urine, racemic lactic acid.
It is likewise decomposed by fungi into butyric and lactic acids.
This inactive inositol, which occurs so widely in fresh green
plants, has been shown by many workers to be present in much
larger quantities in young growing plants (and animals) than in
the adult forms. It almost entirely disappears from plants when
they are slowly dried.
The esters of active inositol, on the other hand, do not vanish
on drying the plants. When we consider the great rarity of
their occurrence, and the fact that the active forms have never
been identified in nature as free inositol, it almost leads one to
assume for them a different origin. Such an origin would be
more in common with that of certain well known plant-con-
* Biochem. Zeitschrift, xxx., 1911, 98.
BY JAMES M. PETRIE. 867
stituents, which also possess side-chains in the 1. 2. 4. positions
on the benzene ring, corresponding to the positions of the
hydroxy! groups of tlie active inositol esters: a few of these ma\^
be mentioned, such as vanillin, eugenol, safrol, coniferyl alcohol,
protocatechuic and caffeic acids.
In conclusion, the author desires to express his indebtedness
to Professor Sir Thomas Anderson 8tuart, in whose laboratory
this work has been done.
►Summary.
The endemic Australian plant, Hetprodendron oleccfoliuin Desf.,
Family Sapindacea?, contains the methyl ester of Ifevo- rotatory
inositol.
The amount isolated was equivalent to 0'65% of the dried (at
100X\) leaves.
This substance is not optically isomeric with the pinite of
Maquenne, which is the methyl dextro-inositol, possessing a
different melting-point and optical rotation.
It is apparently identical with Tanret's quebrachite, and has
been previously recorded from three plants only — Aspidosperma
quebracho (iVpocynacere), Ilfivpa brasiliensis (Euphoi-biacea?), and
GreviUea rohusta (Proteacene).
The occurrence of this compound is, therefore, exceedingly rare,
and is in great contrast to the occurrence of inacfii^p inositol,
which exists as a plastic substance in most plants.
HeterodendroH also contains a cyanogenetic glucoside.
868
AN^ ACCOUNT OF 80ME OBSERVATIONS UPON THE
LIFE-HISTORY OF FHOMA CITRICARPA McAlp.,
The Cause of the ''Black Spot" Disease in Citrus Fruit in
New South Wales.
By G. p. Darnell-Smith, D.Sc, F.I.C.
(Plates Ixxxiv.-xc.)
Historical.
A. H. Benson (1) briefly described the symptoms of a disease,
to which he gave the name of the Black Spot disease of the
Orange, in 1895. He stated that it was probable that it was
identical with a disease of the orange in Europe, that had been
alluded to by Sorauer, under the name of "Schwartz" or black
disease of the orange. There is little doubt, however, from his
figure, that it was the disease caused by Phoma citricarpa
McAlp., with which Benson was dealing. Benson records the
disease as occurring at Seven Hills, Castle Hill, Dural, the
Kurrajong, and Emu Plains.
N. A. Cobb (2) described the Black Spot Disease of the orange
in 1897. He gave some excellent figures of affected fruit. He
also figured the spores. According to Cobb, "these spores arise
after the manner of those of the genus Glceosporium. It is pos-
sible, therefore, that the Australian form is the C olletotriclmm
adustum of Ellis." He further stated that the spores were borne
in large numbers in tandem-fashion from the mycelium at the
base of the interior of the pycnidium in a manner entirely similar
to those of the Bitter Rot of the apple, and gave the size of
spores as 7-8 x 10-15/x.
The fungus causing the Black Spot Disease of Citrus fruit was
described as a new species under the name of Phoma citricarpa
by D. McAlpine, in " The Fungus Diseases of Citrus Trees in
BY G. P. DAHNELL-SMITH. 869
Australia,"(3) November, 1899, and recorded in Vol. xvi., 854,
of Saecardo's Sylloge Fungorum(4). IMc Alpine appears to have
written a letter to Saceardo, in which he stated tliat the Phoma
looked somewhat like a Phyllosticta.
In the Melbourne Herbarium there area number of specimens
marked Phyllosticta citricarjxi — see Phoma. As no Phyllosticta
citricarpa appears to have been described, it is probable that it
is to the letter written by Mc Alpine that Saceardo i-efers in his
footnote.*
McAlpine .states that the Anthracnose di.sease in Florida, due
to C olletotrichum adiistum Ellis, and the black disease of oranses
in Italy known as *'La Xebbia,' due to the fungus Pleospora
hesperidearum Oatt, are quite distinct from the Black Spot dis-
ease of Oranges found in New South Wales, due to Phoma
citricarpa McAlpine.
He states that the disease has not been met with in Victorian
orchards.
Mc Alpine's description of the fungus is as follows : Phoma
citricarpa, n.sp. — " Spots dark brown, at first whitish or greyish
towards centre, but may ultimately become of one uniform
colour, round, sunken, solitary or confluent, varying in size from
1 mm. to 5 mm., and, when confluent, forming large irregular
patches (fin. or more)."
" Hyphae permeating rind, hyaline, septate, branched, 4-5A//.
broad."
" Perithecia, solitary or in groups, somewhat circularly
arranged, minute, black, but dark brown by transmitted light,
punctiform, globular, erumpent ; pore about 20/x diameter,
although it may be somewhat elliptical, 100-120//. diameter.
Sporules hyaline, somewhat variable in shape, elliptical to ovate
or even pear-shaped, with conspicuous granular contents, 8-11 x
4J-6/X, average 9J x 5|/x. (Stained a light green by potassium-
iodide-iodinej; basidia hyaline, slender, about 6/>. long."
" On ripe or still green Oranges, Lemons, and Mandarins,
winter, spring, and summer. New South Wales."
* For this information, I am indebted to Mr. O. C. Brittlebank, Plant
Pathologist to the Department of Aj^rieulture. V'ietoria.
870 LIFE-HISTORY OF PHOMA CITRICARPA,
"The round, sunken, conspicuous spots are generally of a
ruddy-brown tint, and paler in the centre where the pustules are
seated."
" This is a distinctive species in the small size of the perithecia,
as well as in the size and shape of the sporules."
N. A. Col)b(5), in 1904, described some attempts to germinate
the spores of Phoma citricarpa. He states that -" The spores
of this disease do not germinate under the laboratory conditions
ordinarily brought to bear in their examination, and, in this
respect, they differ from most spores found in connection with
prevalent fungus diseases. The following observations, though
they are inconclusive, are inserted here out of regard to the
rarity with which I have observed these spores to germinate. . . .
Spores of this fungus were ringed in a small supply of water and
numerous air-bubbles. After twenty-four hours, the spores had
failed to germinate in those parts where they were completely
surrounded with water. Spores located at the edges of air-
bubbles where, on one side, they had access to air, each sent out
into the air a single very fine unbranched mycelial thread about
one micromillimetre wide, though slightly wider at the free end.
The mycelium was colourless, and so fine as, under the circum-
stances, to preclude observation as to septa; none were seen. At
the end of twenty-four hours, the length of these hyphje averaged
several times the length of the spores."
General Symptoms of the Disease.
The disease, which is now commonly known in New South
Wales as Black Spot or Anthracnose of Citrus Fruit, appears as
minute black spots upon the foliage throughout the year. Upon
the fruit, it is seldom seen till the beginning of August. In
September, particularly after hot westerly winds, it may quite
suddenly make its appearance upon the fruit throughout an
orchard. Dark brown, irregular spots first make their appear-
ance upon the skin. The spots vary in size from one-eighth of
an inch or less to half an inch or more in diameter. Later, these
become depressed, and some may become confluent. The inner
part of the spot next becomes of a light cream-colour, and some-
BY G. P. DARNELL-SMITH. 871
what parchment-like in texture. Shortly after, minute black
pycnidia are formed beneath the surface; and, breaking through,
form small irregular fissures or minute holes, at which points the
spores are discharged through the necks of the pycnidia.
The disease almost invariably makes its appearance on the
sunny side of the tree, and upon that side only of the fruit which
is exposed to the sun. So constantly is this the case, that, in
an affected orange, it is almost always possible to draw an equa-
torial line dividing the sunny from the shaded side of the fruit,
and, on the former side only, will black spots be found. Even
on the sunny side of the tree, if the fruit is well shaded by
foliage, it is seldom affected by the disease, even when exposed
fruits around it are badly marked.
That the rind has some principle that may inhibit the growth
of spores, is suggested by the incidence of the disease in the
various varieties of citrus fruit. It is common on the orange
(navel, valentia, siletta), it is found less frequently on the
Emperor mandarins, and quite exceptionally upon the thorny
mandarin.
The infection of the fruit only upon one side of the tree sug-
gests that it may be related to the prevailing wind, or to the
effect of too much sunlight or heat upon the rind.
The development of the disease only upon the sunny side of
the tree indicates that it is the sun, rather than the wind, that
exerts an influence.
To test the effect of diminishing the amount of sunlight fall-
ing upon the trees, and to protect them from scorching winds,
two trees in an orchard were completely covered-in with hessian
in the form of a tent. They were covered in at the beginning
of June, that is, at the commencement of the ripening period.
The fruit on these trees did not show black spot on the fruit
for two or three weeks after the uncovered ones, and then the
spots developed very slowly, remaining a dull brown colour; and
they did not pit the fruit to such an extent as fruit exposed to
the weather. The spots upon uncovered fruit soon become black.
The following season, two trees were protected from the sun by
putting up a screen of hessian upon the sunny side only. Here
872 LIFK-HISTORY OF PHOMA CITRICARPA,
again, the disease was slower in developing upon the fruit, and
the spots remained of a dull brown colour. Since dew would be
just as heavy upon one side of the tree as another, its action,
except in conjunction with the sun, may be left out of account.
As regards the wind, the prevailing winds would tend to blow
spores upon the east side of a tree in the Gosford district, where
these observations were carried out: there is very seldom a north-
west wind.
The spores from infected leaves, falling upon the fruit, become
uniformly distributed in the dew-drops. This uniformity of dis-
tribution is shown in the equidistant positions that the ripe
spores take up, when introduced into hanging drops.
From the foregoing considerations, it seems probable that the
spores distributed on the surface of the fruit effect an entrance,
with their germ-tubes, into the rind, when its natural inhibiting
power, due to physical or chemical causes, has become impaired
bv exposure to the heat of the sun.
Culture op the Fungus.
By successive transportation of groups of spores to sterile
drops of water, isolated spores were obtained. These were trans-
ferred to various culture-media, and pure cultures in Petri-dishes
or test-tubes, were at length obtained. These were grown in an
incubator at a temperature of 22''C. The media in which the
fungus was most successfully cultivated were ordinary nutrient
agar, glucose-agar, and agar impregnated with watery extract of
orange-peel. The growth of the fungus upon each of these three
media is very different.
Upon ordinary nutrient agar, growth proceeds very slowly.
Little patches of dark brown hyphie arise, creep over the medium,
and penetrate it. But they seldom travel far from the site of
inoculation, and, if several spores have been intr(^uced into a
Petri-dish, it is possible to obtain several isolated plants before
the hyphse have become interlaced.
Upon glucose-agar, the other extreme is reached. The plant
• (M'ows so luxuriantly, that, in a test-tube, the agar quickly
becomes a carbonaceous-looking mass, tilled with quantities of
BY G. P. DARNELL-SMITH. 873
dark brown liyphfe; on tlie sin-face, aerial liyplifp are sent up,
covering the wliole witli a light grev nivceliuni (Plate Ixxxiv.,
fig.l).
Upon agar impregnated with water-extract of orange-peel, an
intermediate type of growth was obtained (Plate Ixxxiv., fig. 2).
The fungus grew well but not luxuriantly, and there was little
production of aerial hyplije.
In these cultures, pycnidial formation began after about seven
days, and ripe spores were obtained after thirteen days: but, in
the glucose-agar, where growth was luxuriant, spore-produotion
was much delayed.
Production of Pycnidia and Spokes.
When grown upon a medium that is thoroughly moist, the spores
that are formed in the pycnidia are expelled, and appear upon
the ostiole in a mass. This mass is whitish in colour, viscid and
semi-translucent. Usually it is globular, but it may take the
form of a truncated cone. When placed in water, the spores do
not readily separate from the mass, 'i'hese spore-masses may be
found in nature if the fruit is continually surrounded by a damp
atmosphere. Much more generally, however, the spores are
produced in pycnidia surrounded by a dry atmosphere. They
do not then emerge till a drop of moisture falls upon the pycni-
dium, whereupon tliey escape in a thin stream. Observed in a
hanging drop, they are seen to separate and spread themselves
at spaces almost equidistant from one another over the surface
of the drop, as if they were mutually repellant; and no two
spores are found to remain in juxtaposition. This separation is
probably brought about by surface-tension. Normally, the pyc-
nidia are produced upon the peel of oranges, and tlie ostioles
open at the surface. A .surface-view of a pycnidium, with an
ostiole in the centre, is shown in PI. Ixxxv., fig.3.
If, however, affected oranges be kept under a bell-jar, and
remain free from attack by other fungi, they shrink somewhat,
and the peel becomes harder. The mycelium of the fungus
penetrates the whole of the pulp, which becomes black in colour,
and pycnidia are formed in abundance throughout the tissue.
874 LIFE HISTORY OF PHOMA CITRICARPA,
The hypliae even penetrate the outer coat of the testa of the
seed, and pycnidia are there produced (PL Ixxxv., fig. 4). A
somewhat similar phenomenon is observed when the fungus is
grown upon an orange rind-extract agar-slope in a test-tube. At
first, the mycelium grows near the surface, and pycnidia are
produced tliere; but, later, the hyph?e produce submerged pyc-
nidia, which may eject their spores as spore-masses into the
surrounding medium; in these cases, the ostiole appears to have
no particular orientation.
It is noteworthy that oranges affected with Phoma citricarpa
and free from any other disease, if kept under a bell-jar, give off,
in the course of several days, very little moisture; but if they
are attacked by Penicillium italicum (to which attack they are
peculiarly liable in the presence of Phoma cit7'icarpa), a large
quantity of moisture condenses upon the surface of the bell-jar
in a few hours.
Structurk of the Mycelium.
The mycelium exhibits much diversity. The extreme tips may
be pointed or round, the hyphse being thin, hyaline, and almost
devoid of septa. Further back, the liyphse become somewhat
suddenly thicker, the septa become more numerous, and the
colour is olive-green. In the older hyphse, the septa are very
numerous, the colour is dark greenish-brown, and the contents of
the cells granular; the cells may be oblong, or round, and often
carry numerous, short, round, protuberances (PI. Ixxxv , fig. 5;
PI. Ixxxvi., fig.6).
Structure of the Spores.
Two kinds of spore are produced. Both kinds may be pro.
duced in a pycnidium or one kind only. They differ considerably
in size. The large spores are those described by McAlpine^
whose measurements, in regard to the size of these spores and
the pycnidia, T can confirm. These spores are usually hyaline,
with granular contents; they have, however, frequently a greenish
hue. The cell-wall is very thin. They may have one or two
nuclei, generally there are two, and these are placed opposite to
one another adjacent to the cell-wall in the region midway
BY G. P. DARNELL-SMITH. 875
between the ends of the cell; frequently they are joined together
by a narrow band which stains deeply with hsematoxylin. When
the spore is about to germinate, the protoplasm becomes much
vacuolated and the nuclei appear to fragment (PL Ixxxvi., fig.7).
The second kind of spore, which has not been previously de-
scribed, is much smaller, measuring only 7-5 x 1 -G/x.
The extremities are usually slightly thicker than the middle,
the spores being somewhat dumb-bell shaped.
The spores have the following dimensions :
L.= length, c. = centre-width. E. broadest-end.
Range: (L.) 6-0 - 9-3 x c. (1-0- 2-0) xE. 1-3 x 20/x.
Average : (L.) 7'5 x c. 1-3 x E. 1-6/ji.
A highly refringent granule is usually present at each end.
Sometimes there are three or four granules present, or there may
be none at all. The spores themselves occasionally have the form
of simple rods, or they are moniliform or hourglass-shaped. If
present, they are discharged from the pycnidia with the larger
spores in the viscid masses previously referred to, or they may
be discharged in a stream from a ripe pycnidium when moistened
(PI. xc, fig. 25). They do not separate from one another in the
manner of the larger spores; indeed, they sometimes lie side by
side like rouleaux of red blood-corpuscles (PI. Ixxxvi., figs. 8, 9).
As I have not been able to induce these spores to germinate,
or to determine their functions, I shall speak of them as "x"
spores.
The presence of "x ' spores in the pycnidia of various Sphce-
ropsidce is known in a few genera.
F. A. Wolf (6), in describing Ascochyla hoi'torum, which was
formerly known as Phoma solani, states that, in the pycnidia, he
finds typical conidia, 6-10 x 2"5-4//,, together with "a second type
of spore which is hyaline, continuous, frequently curved or
hooked at one end, 14-17 x 2-2 S/y.. These spores may occur in
the pycnidium together with the pycnospores, or alone in other
pycnidia." He continues, "Morphologically, at least, they
are identical with the stylospores of Nitschke in Diaporthe, the
" B " spores of Diedicke in Phoniopsis, the scolecospores of Spear
and the paraphyses of Reddick in Fusicocann, the pycnidial form
^76
LIFE-HISTORY OF PHOMA CITRICAHPA.
of Cytosporella vitiada. Only failure lias thus far met the
various attempts to germinate these bodies."
Gehmination of the Spores.
The normal pycnospores do not germinate readily in water.
After several days, a few may send out an attenuated hypha,
which seldom develops branches, and soon ceases to grow.
In a nutrient solution (containing amm<niium nitrate, 10 gram;
dihydrogen potassium phosphate, Go gram; magnesium sulphate,
0*25 gram; iron chloride, trace; cane sugar, 5'0 grams; water,
100 c c.) the spores assumed a dark olive green colour, and only
a few commenced to germinate after several days. The hyphse
produced were a very dark green colour; they appeared stunted
and unable to elongate (PI. Ixxxvi., fig.lO). When a solution of
equal parts of peptone-water and this solution was tried, ger-
mination was somewhat better, but unsatisfactory. In an
aqueous extract of orange-peel, however, the spores germinated
freely. The rapidity with which the spores germinated in this
medium appeared to depend largely on the age of the spore If
placed in the solution immediately they were discharged from
the pycnidium, the spores had produced a germ-tube in twelve
hours. If, however, the spores were three days old, they took
several days to germinate; and, if much older, many of them
failed to germinate at all. The germinating spore usually sends
out a germ-tube from the side, but it may produce a germ-tube
from one end, or from both ends simultaneously. The granules
in the spore assume a greenish hue, and pass, for the most part,
into the germ-tube. A septum may be produced in the germ-
tube early, or the production of septa may be deferred (PI. Ixxxvi.,
tig. 11). The hyphai soon branch, their ends being at first lound,
and, innnediately behind the tips, the cell-contents are usually
highly granular (PI. Ixxxvi., fig. 12).
By taking special pre(;autions to prevent the hanging drop
from drying up, and to have every part of the apparatus and
instruments used sterile, T have been able to follow the develop-
ment of a spoi-e in a hanging drop from the production of a germ-
tube to the formation of a pycnidium by the mycelium, and the
IBT G. P. DARNELL-SMITH. 877
discharge of spores from tins pycnidiuni. Such a pycniflium is
shown in PI. Ixxxvii., Hg.l3. A noticeable feature of theliyphtv
is the ease with which they anastomose. Anastomosis may occur
in two ways. A branch from one hypha may grow out and fuse
with another lying parallel or nearly parallel to the one from
which the branch originated, or branches may arise from two
adjacent hyphjf, approach each other, meet, and fuse. Examples
of this are seen in PI. Ixxxvii., fig. 13. The pycnidium hero
figured took a little over three weeks to form, and ultimately
discharged "x" spores only. In other hanging drop cultures,
where noimal pycnospores have been discharged, they have never
been observed to germinate, without transference to a fresh
medium, the mycelium having apparently exhausted the medium
upon which it was growing before the production of pycnidia.
Development of the Pycnidia.
By means of cultures in hanging drops, and serial sections of
cultures upon agar, it has been possible to follow tlie develop-
ment of the pycnidia in detail.
The beginning of pycnidia-formation consists in one or more
adjacent h37phaj producing lobulated branches of much greater
diameter than the ordinary hyphie (PI. Ixxxvii., figs. 14, 15).
These lobulated branches stain rather more deeply with hsema-
loxylin than the ordinary hyphjfi. 'J'he lobulated branches become
greater in number and interwoven, and many septa develop (PI.
Ixxxviii., figs. 16, 17). This leads to the formation of a firm, com-
pacted pseudoparenchyma; the protoplasm which lines the cell-
walls contains numerous deeply-staining particles, and becomes
especially distinct (PI. Ixxxviii., fig. 18).
Soon, the parenchymatous body ig distinguishable int(j outer
layers of small, thick-walled, brownish cells: and an inner region
of larger, thin-walled, parenchymatous cells. In the inner region,
we, later, find groups of hypha3 with thin lumina, and, in their
place, arise, at a later date, the loculi of the pycnidia.
In the transition-stage from parenchyma to hyphte, the walls
of the parenchyma cells frequently appear to be dissolved, and a
mass of naked protoplasm, with deeply staining granules, is seen
(PL Ixxxix., fig.l9).
878 LIFE-HISTORY OF PHOMA CITRICARPA,
The following stages may be distinguished in the production
of a pycnidium :—
1. Production of lobulated hypliae, which become interwoven.
2. Formation of a pseudo-parenchyma.
3. Dissolution of cell-walls of the pseudo-parenchyma at certain
foci, followed by formation of hyphse.
4. Establishment of loculi at these foci, into which spores are
abstricted from the hyph^e.
5. Gradual replacement of the pseudo-parenchyma by spore-
bearing hyphie.
6. Shrinkage of the spore-bearing hyphae.
7. Pycnidium completely full of spores borne upon attenuated
stalks.
The transition from pseudo-parenchyma to spore-bearing hyphie
is shown in PL Ixxxix., figs. 20, 21. A portion of the pseudo-
parenchyma frequently lines the pycnidium until a late stage.
In the natural state, when growing upon the orange, the
pycnidiaaregenerally unilocular (PL xc, fig. 22). But, in cultures
upon orange-agar, they are frequently bi- or trilocular, in which
case, the loculi are surrounded by a common wall (PL xc, fig.23).
In bilocular pycnidia, portions of the inner parenchymatous
tissue have become firm, and the groups of thin-walled hyphse
have become separated.
Pycnidial formation has been briefly described in the case of
Diplodia zece by Van der Bijl(7), and in the case of Phoma
lavamluhe by W. B. Brierley(8).
The origin of the pycnidium by the interlacing of lobulated
hyphse with the formation of a pseudo-parenchyma is very similar
in Phoma cilricarpa and I'h. lavaaduUn. The later stages in the
development of the pycnidium of Ph. citricarpa resemble the
description given of that of Diplodia zece. The marked visiijility
of the protoplasm lining the cells of the pseudo-parenchyma, and
the disappearance of the cell- walls previous to the formation of
hyphae, described in the case of Fh. citricarpa, is not mentioned
by the authors quoted.
The formation of hyphse from naked protoplasm is not un-
known, as it occurs in the Myxomycetes, when the capillitium,
BY G. P. DARNELL-SMITH. 879
consisting of a system of solid or of tubular threads, appears
before the spores are formed. The capillitium has l)een described
by Harper &, Dodge(9), as originating in vacuolar spaces in the
cytoplasm, which elongate and take on the tubular form of young
capillitial threads. They find that the capillitium is formed by
the deposit of materials in the vacuoles from which the capillitial
threads are formed; and that radiating threads run out from
the larger granules, which are deposited by the process of intra-
protoplasmic secretion. These radiating fibrils suggest, rather
strongly, that they are cytoplasmic streams which are bringing
materials for the formation of the capillitial wall and its thick-
enings, which are laid down as spirals, suggesting that the pro-
cess is comparable with the ordinary process of cell-wall forma-
tion, but along internal plasma-membranes, rather than external.
Development of the Spores.
The stalks bearing the normal spores are unicellular, hyaline,
and rich in protoplasmic contents, sometimes excessively so.
Paraphyses have not been distinguished, though stalks vary
much in length; and sometimes old stalks, or stalks upon which
the spores have not developed, have the appearance of paraphyses.
The spore develops as follows : —
The stalk elongates, and the spore is abstricted from its end;
at this stage, the spore is pear-shaped. 'I'he stalk is slightly cup-
shaped at its extremity occasionally, and uni-nucleolate (PL xc,
fig.24). The spore increases in size, and becomes binucleolate;
at the same time, the stalk shrinks to a mere thread. On their
discharge, the spores may show no traces of their point of attach-
ment; or each spore may have, still joined to it, a minute thread.
The "x" spores develop by abstriction in concatenation from
minute hyphaj bordering the loculus of the pycnidium (PL xc,
fig.24).
I am indebted to my assistant, Mr. VV. A. Birmingham, for
much careful work during the progress of this investigation.
Summary.
1. " Black Spot" is a serious disease of Citrus fruits in New
South Wales, due to the fungus Phoma citricarpa.
880 LIFE-HISTORY OF PHOMA CITRICARPA,
2. It develops only on the sunny side of trees, and upon the
parts of the fruit exposed to suidight.
3. Pkoma citricarpa produces two kinds of spores, normal
pycnospores, and "x" spores.
4. Normal pycnospores germinate readily in suitable media:
it has not been possible to induce the ''x" spores to germinate.
5. The details of the formation of the pycnidia, anfl of the
pycnospores have been worked out.
6. Spraying with Bordeaux (6 - 4 - 50; followed up by spray-
ings with weaker solutions, controls the disease.
7. A study of the life-history of the fungus indicates that spray-
ing need not commence till the fruit is half grown.
LITERATURE CITED.
1. Benson, A. H. — "Black Spot of the Orange." Agric. (iaz. X. S.
Wales, Vol. \i., Pt.4, p.249 (1895).
2. UoBB, N. A.— "Black Spot of the Orange." Op. ell.. Vol. viii., Pt.4.
p. 229 (1897).
3. Mc Alpine, D.— The Fungus Diseases of Citrus Trees in Austialia (1899).
4. Saccardo — Sylloge Fungoruni, Vol. xvi., 854.
5. Cobb, X. A. — Letters on Diseases of Plants. Second Series, p. 72
(1904).
6. Wolf. F. A.— ''Egg-Plant Rots" Mycol. Central!).. Vol.4, p. 27S
(1914).
7. Van der Bljl. Pail A. — "A study of the "Dry Rot" disease of Maize.
caused by Diplodia zt<f." Science Bulletin, Xo.7. Dept. of Agri-
culture, Union of S. Africa,
8. Briekley, \V. B.— Kew Bulletin, 1910, Xo.5.
9. Annals of Botany, xxviii., 1-18, Jan. 1914.
KXIVLAXATIOX OF PLATES LXXXIV.-XC.
Plate Ixxxiv.
Fhoma cifric((rpa.
Fig. 1. — Pure cultuie in gluco.se-agar. The cidture grows ver^^ rapidly,
produces numerous aerial hj'plije, forming a grey-coloured felt, but
does not readilj' produce pj'cnidia.
Fig.2. — Pure culture growing upon orange-rind-extract agar. The culture
grows at a moderate rate, remains dark in colour, produces few
aerial hj'plue, and forms pycnidia readily.
BY G. P. DARNKLL-SMITH. 881
Plate lxxx\ .
Fi.i^.3. — Smfaee-view of a pyrnidiuni. sliowint^ the ostiole in the centred
( > 480).
Fig. 4. — Section of the testa of an orange -seed which has been penetrated
bytlie liypha^ of P/ionta cift-intr/ia, and in which two small pycnidia
have been produced.
Fig..>. — Young hj'phte, showing elongated cell; ( x 4S0).
Plate Ixxxvi.
Fig.6. — Old hyphre, showing dark brown-coloured cells, which are more or
less circular, Mith numerous protul)eranees; ( x 480).
Fig. 7. — Spores in various stages of growth : a, a j'oung spore showing two
nuclei; b, c, d, spores about to germinate, .showing vacuoles and
fragments staining deeplj' with hfcmatoxylin.
Fig.8. — "x" spores arranged in rouleaux.
Fig.9. — "x" spores showing highly refringent dots.
Fig. 10. — Pycnospores germinating in nutrient solution, showing stunted
hyph^e with a tendency to bud; ( x 480).
Fig, 11. — Pj'cnospores germinating normally in orange-rind-extract, shoM -
ing germ-tubes, septa, and granular contents; ( x 480).
Fig. 12. — Establishment of a mycelium from a germinating spore; ( x 480).
Plate Ixxxvii.
Fig. 13. — Formation of a pycnidium from the mycelium growing in a hang-
ing drop. The hyph^ of the mycelium are shown anastomosing b^'
the union of adjacent branches or by the direct fusion of one hypha
with another by a branch; ( x 480),
Figs. 14, 15. — Formation of lobulated branches, first stages in the develop-
ment of a pycnidium; ( x 480).
Plate Ixxxviii.
fie/. IH. — Formation of lobulated branches, first stages in the development
of a p3''cnidium; ( x 480].
Fig. 17. — Section through a young pycnidium, showing the development of
septa in the lobulated branches, and the establishment of a pseudo-
parenchyma; ( X 480),
Fig. 18. — Section through an older pycnidium, showing the protoplasm,
with granular contents, lining the cell-walls of the pseudo-paren-
chyma; ( x480).
Plate Ixxxix.
Fig. 19. — Section through a pj'cnidium at a later stage showing dissolution
of the cell-walls of the pseudo-parenchyma before the establishment
of spore- bearing hyphte; \ x 480).
882 LIFE-HISTORY OF PHOMA CITRICARPA.
Figs. 20, 21. — Portions of sections through a mature pycnidium, showing
the outer layers of dark brown, thick-walled cells, lined bj' the
remains of the pseudo-parenchyma from which the spore-bearing
hyph£fi arise; ( x 480).
Plate xc.
Fig. 22. — Section through a pycnidium upon orange rind. The pycnidium
contains mature spores. The slightly increased development of the
outer layers of thick-walled, brown cells just beneath the ruptured
epidermis, is noticeable; ( x 480).
Fig.23. — Section through a trilocular pycnidium grown upon orange-rind-
extract agar. There is a common outer covering, and the loculi
are separated by walls composed of fine hj'phai; ( x 120).
Fig.24. — Section through a pycnidium in which "x' spores are being
abstricted from fine hyphfe; ( x 120).
Fig. 25. — (Iroup of normal p3'cnospores and "x" spores discharged from a
pycnidium; ( x 480).
883
THE OCCURRENCE OF AN INVERTED HYMENIUM
IN AGARICUS CAMPESTBIS.
By G. p. Darxell-Smith, D.Sc, F.T.C.
(Plates xoi.-xciii.)
In 1917, some inuslirooiii-growers, who were growing mush-
rooms on a large scale in an old disused railway-tunnel, brought
to me, for examination, a number of mushrooms that were un-
saleable on account of their appearance. The mushrooms were
grown upon large beds of manure, the making of which I had
superintended. The manure was fairly moist, it was lightly
compacted, and the average temperature was 22''C. Both the
stipe and the cap of the specimens were somewhat tough. The
cap was quite unusual in appearance. Instead of having the
normal, comparatively smooth skin, its surface was broken by
one or mor^ black protuberances, 1*25 cm. or more in diameter,
that looked like boils (Plate xci., fig.l). It was this appearance
that spoilt the sale of the mushrooms. Samples of mushroom-
spawn from Sydney, from Adelaide, and from France, were grow-
ing in the same tunnel, and providing normal mushrooms; it was
only a particular sample of spawn imported from France that
was giving rise to these abnormal specimens. A close inspection
of the black protuberances showed that they were composed of
sinuous, labyrinthiform gill-lamellse (PI. Ixci., fig.2). They had
the appearance of small inverted caps, but no appearance of a
stipe could be found. Sections through the cap showed that
these structures were quite separate from the normal hymenium,
which was present on the undersurface of the cap (PI. xcii., fig. 3).
Sections for microscopical examination were prepared and
stained. They showed that the structure of the hymenium on
the undersurface of the cap was quite normal.
Sections through the hymenium on the upper surface of the
884 AN INVERTKD HYMENIUM IK AGARICUS CAMPESTRIS,
cap, however, showed it to be quite abnormal in character (PL
xcii., fig. 4). They showed that the "gills" had a most irregular
outline, that they varied much in width, and that many of them
were hollow. Spores in abundance were produced, both on the
external surface of the gills, and on the walls lining the internal
cavities. These spores were produced upon enlarged cells, and
borne upon sterigmata, as in the case of normal gills. While,
however, the normal number of sterigmata is four, cells bearing
only one, or only two sterigmata were found, as well as cells
bearing the normal number of four (PI. xciii., figs. 5, 6, 7). Crops
of mushrooms, showing the abnormal development above de-
scribed, are not unknown. W. A. Smith, according to Wors-
dell(l), found, on more than one occasion, crops of mushrooms,
every individual of which had an inverted cap on its surface.
According to Worsdell, an inverted hymenium may arise (1)
through the congenital formation of an inverted cap, or caps,
from the earliest stage onwards on the upper surface of the
primary cap; (2) through the formation of inverted caps by local
invagination of the margin of the primary one. It is held, how-
ever, that these two valuations really represent the same pheno-
menon, of which (1) represents the final and completed stage of
(2) arising congenitally and isolated.
In my specimens, no trace of the formation of inverted caps
by local invagination could be found. Moreover careful examina-
tion of mushrooms in the " button "-stage showed, that these
irregular gill-lamellae, on the upper surface of the cap, were in
process of formation in the very earliest stages, and long before
the velum had separated from the stipe (PI. xciii., fig. 7). There
can, therefore, be no question of invagination of the cap in these
specimens.
The question arises, what light, if any, does this inversion of
the hymenium throw upon the phylogeny of the Agaricacese? Is
it an expression of a partial reversion to an ancestial character^
That certain spawn has particular characters peculiar to it, is
the experience of professional mushroom-growers. In commer-
cial practice, under the influence of the atmosphere and the heat
of the manure-beds, the mycelium gets weaker and weaker, and
BY G. P. DAUNEr,L-SMITH. 885
eventually perishes. "Virgin spawn" obtained fiom tlie fields is,
therefore, transferred to beds of manure, wliere it spreads and
supplies the grower with a "frank spawn" for growing the edible
mushroom. But "often twelve to fifteen kinds of vii-gin spawn
as found in the fields or on heaps of refuse are discarded as use-
less before a good strain is obtained "(2). That the production
of inverted caps was a feature inherent in the spawn under dis-
cussion, is shown by tlie fact that all mushrooms developed from
it produced abnormal caps, whereas mushrooms produced fiom
other spawn, under identical conditions, did not.
This phenomenon, together with the characters of the spoi"0-
phores, lends support to the hypotheses that have been put
forward as to the evolution of the cap, and as to the probable
origin of the Agaricaceie, which may be stated briefly as follows:
(a). Evolution of the ca-p
1. The production of the horizontally extended, llatteued cap-
form of fruit from an original, cylindric, dome-shaped form.
2. The relegation of the hymenial tissue to the lower surface
of the cap.
3. The formation of "gills " from the original pore or alveolar
structure.
(6). Evolution oj the Agaricacea^.
The most primitive type of fructification is probably that of
Glavaria, in which a cylindric or club-shaped branch is uniformly
covered with hymenium, which extends down the stalk as well
for some distance.
"Cases in which the hymenium covers the whole of the upper
exposed surface, in the form of a semi-alveolar structure, or
labyrinthiform gill-formation, are seen in Tremella and Ncema-
telia, and these plants pertain to the Protobasidio-Mycetes; the
Tremellinese have, according to Maire's classification, branched
off laterally from the Auriculariacese, and it is from these latter
that the Agaricacese and Polyporace?e have descended.
In this connection, it is interesting to note that, in one of my
specimens that was fairly large, but was obtained in the "button-
stage" before rupture of the velum, the whole of the upper surface
of the cap is covered with sporogenous tissue (PI. xciii., fig. 9).
68
886 AN INVERTED HYMENIUM IN AGARICUS CAMPESTRIS,
Moreover, this sporogenous tissue is completely covered by a thin
membrane, so that there can be no question of its having attained
its position through invagination.
The examination of these abnormal mushrooms, therefore,
leads to the conclusion, that they represent a partial reversion to
an ancestral character. One would hardly expect a complete
reversion; and that it is only partial, is demonstrated by the
manner in which the spores are borne.
The discharge and disposal of the spores in the Agaricaceje,
according to Buller(4), are brought about, as follows : —
The special conidiophore, or basidium, usually bears four
spores, which are discharged successively, and each spore becomes
violently detached. The violent discharge of the spores prevents
the adhesive spores from massing together, and from sticking
fast to the gill-surface. At first, the spore is shot out horizon-
tally; then, under the influence of gravity, it describes a sharp
curve, and then falls vertically. The path described by the
falling spore has been appropriately called a sporabola. After
falling, under the influence of gravity, in the still air between
the gill lamellae till they reach the exterior, the spores are borne
away by the breeze. Basidia, being four sterigmata with four
spores attached, were found in my specimens in the inverted
hymenium; and it is obvious that, if the spores were discharged
in the normal way from the sterigmata and then started to fall
under the action of gravity, they would not get free from the
hymenium at all, but would fill up the spaces between the gills
We have, therefore, a reversion to an ancestral condition so
far as the macroscopic characters are concerned, with a retention
of the modern condition so far as the microscopic characters are
involved.
LITERATURE CITED.
1. WoRSDELL, W. C. - Principles of Plant Teratology, Vol. i., p,33 (191o).
2. Aquatias, p. — Intensive cultui'e of Vegetables on the French Sj'steni,
p.153 (1913).
3. WoRSDELL, W. C— Principles of Plant Teratology, Vol. i., p.3U(191o).
4. BuLLER — Researches in Fungi, p. 144 (1909).
BY G. P. DARNELL-SMITH. 88 <
EXPLANATION OF PLATES XCI.-XCIII.
Plate xci.
Fig. 1. — Abnormal mushroom, showing protuberances of the cap. At the
4)ase of the stipe, two small "buttons" are seen.
Fig.2. — Abnormal mushroom, showing hymenium on the surface of the
cap.
Plate xcii.
Fig. .3. — Section of the same.
Fig. 4. — Transverse section through the liymenium on the surface of the
cap, showing the irregular character of the gills, and the large
spaces in their interior; ( x 120).
Plate xciii.
Figs.S, 6, 7. — Basidia bearing varjnng numbers of sterigmata (4, 2, 1) from
the hymenium in the surface of the cap; ( x 480).
Fig. 8. — Section of small "button" much enlarged, showing the formation
of lens-shaped areas in the cap containing sporogenous tissue before
the separation of the velum.
Fig. 9. — Section of a large "button" (nat. size), showing sporogenous tissue
covered with a membrane over the whole upper surface of the cap.
The velum has not yet split.
886
DONATIONS AND EXCHANGES.
Received during the period November '28th, 1917,
to November 27th, 1918.
( Frotn the respectivp. Societies, etc.^ unless other^cise mentioned.)
Accra.
Geological Survey of thk Gold Coast of W. Africa —
Report for the Year 1916(1918).
Adelaide.
Department of Chemistky of South Australia —
Bulletin, No.7(1917).
Department of Forestry, University of Adelaide —
Bulletin, No.6(1918).
Department op Mines: Geological Survey of S.Australia —
Annual Report of the Government Geologist for 191 6(191 7).
Review of Mining Operations in the State of South Aus-
tralia during the Half-years ended December 31st, 1917,
and June 30th, 1918, Nos.27-28(1918).
Public Library, Museum, etc., of South Australia —
Records of the S.A. Museum, i., 1(1918).
Report of the Board of Governors for 1916-17(1917).
Royal Society of South Australia —
Transactions and Pi'oceedings, and Report. xli.(1917).
Albany, N.Y.
New York State Library —
Annual Report of the N. Y. State Museum. Ixviii., 1914
(1916).
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Ann Arbor.
Michigan Academy of Science —
Seventeenth and Eighteenth Annual Repoils, 1915, 1916
(1916).
Baltimore.
Johns Hopkins University —
Hospital Bulletin, xxviii., 321, 322; xxix., 323-332( 1917-18).
Barcelona.
Junta de Ciencies Naturals de Barcelona —
Musei Barcinonensis Scientiaruni Naturalium Oijeva. Series
Biologico-Oceanographica, i. (1917)^Ser. Botanica, i -ii.
(1917) — Sei'.Geologica, i.(1918) — Ser. Zoolugica, xi.(1917).
Basle.
Natukforschende Gesellschaft in Basel —
Verhandlungen. xxviii.(19l7).
Berkeley, Cal.
University of California —
Publications.— ^oton2/, v., 12-14; vi., 15-16(1 918). — ^n«5o-
mology, i., 8; ii. (complete) [1 917-18].— (reo^y/, x. 13, 18,
20-28; xi., l--2{\m-U).~Physwlo<jij, v., 3(1918).— ^o-
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Five Reprints : (a) " Non-influence of Injections of pure
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Immunology, ii., No.4, June, 1917].— (6) "Stability of
Emulsions,"' lirc, by I. C. Hall [Journ. Phys Chem., xxi.,
No.8, Nov., 1917]. — (c-) " Note on the aerobic Culture of
Anaerobes," l^c, by L. J. Ellefson and I. C. Hall [Science,
N.S., xlvi , No. 1197, Dec, 1917]. — (c^ "Occurrence of a
Positive Intracutaneous Reaction," etc., by F. P. Gay and
A. J. Minaker [Journ. Amer. Med. Assocn., Ixx., Jan ,
1918].— (e) "Automatic Water- Level for Arnold Steril-
isers," Ijy I. C. Hall [Journ. Bacteriology, iii., No. 1 , Jan.
1918].
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Botany Bulletin, No. xx.(1918).
donations and exchanges.
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DiRKCTOu OF Fisheries, Bengal, Bihar, and Orissa —
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well and P. Prashad [Journal of Parasitology, Vol. iv.,
March, 1918].
Geological Survey of India —
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I
892 DONATJONS AND EXCHANGES
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894 DONATIONS AND EXCHANGES.
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Sec. D, xii., 4-6; xiii., 1-5(1917-18).
Contents and Index of Vols, i.-x., 1906-15(1917).
Marseilles.
Musee d'Histoire Naturelle de Marseille —
Annales. T.xiii.(1914); xv. (1915-16).
Massachusetts.
Tufts College —
Tufts College Studies (Scientific Series), iv., 5(1917).
Melbourne.
Australasian Journal of Pharmacy — Vols xxxii., 384 ;
xxxiii., 385-395(1917-18). From the Publisher.
Commonwealth of Austrat.ia : Advisory Council of Science
and Industry —
Report of the Executive Committee for the Year ended
June 30th, 1918(1918). ^
Commonwealth of Australia: Dept. of Trade and Customs —
Fisheries : Biological Results of the Fishing Experiments
carried on by the F.I.S. '* Endeavour," 1909-14. Vol. iv.,
Parts 5-6; v., 1(1918). \^Received through the Australian
Museum^ Sydney\
Commonwealth Bureau of Census and Statistics —
Official Year Book of the Commonwealth of Australia. No.
10(1917).
DONATIONS AND EXCHANGES. 895
Pocket Compendium of Australian Statistics, 1918(1918).
Department of Agriculture of Victoria —
Journal, xv., 12; xvi., 1-11(1917-18).
Report of the Council of Agricultural Education, and of
Dookie and Longerenong Agricultural Colleges (1918).
Field Naturalists' Club of Victoria —
"Victorian Naturalist." xxxiv., 8-12; xxxv., 1-7(1917-18).
Public Library, Museums, &c., of Victoria —
Report of the Trustees for 1917(1918).
Royal Australasian Ornithologists' Union—
"The Emu,' xvii., 3-4; xviii., 1-2(1918).
Royal Society of Victoria —
Proceedings. New Series, xxx., 2(1918).
University of Melbourne —
Calendar. 1918(1917).
Mexico
Instituto Geologico de Mexico —
Anales. Nos. ii.-iii. (191 7-18).
Boletin. Num.36( 19lJ>).
Modena.
La Nuova Notarisia — From the Editor, Dr. G.B.De Tom
Serie xxix., Gcnnaio, 1918(1918).
Monaco.
Musee Oceanogkaphique de Monaco —
Bulletin. Nos.326-339(1917 18).
Nantes.
Societe des Sciences Natuiielles dh; l'Ouest dv. la France —
Bulletin, 3"'^Serie, iv., 1-4, T.p..i:c.(19H).
Ne"w Haven, Conn.
Connecticut Academy of Ahts and Sciknces —
Transactions, xxi., T.p.ttc; xxii., pp.249 267, T. p. itc.( 1918).
Ne^v York.
American Geographical Society —
Geographical Review, iv., 5-6; v., 1-6; vi., 1-3(1917-18).
896 DONATIONS AND EXCHANGES.
American Museum of Natural History — .
Bibliography of Fishes, by B. Dean, enlarged and edited V)y
C. R. Eastman. V^ol ii.(iyi7).
New York Academy of Sciences —
Annals, xxvii., pp.215-243( 191 7).
Ottaw^a.
Geological Survey of Canada —
Map of Whiteburn Gold District, N.S. (Piibln. No. 1690)
[1917].
Memoirs. Nos.95, 99-103(1917-18).
Museum Bulletin, No 27(1918).-
Paris.
"Journal de Conchyliologie.'' Ixiii., 2-4(1917-18).
Museum d'Histoire Naturellk de Paris —
Bulletin, xl., 7, T.p.itc: xli., 1-7, T.p.itc; xlii., 1-8, T.p.ii:c.;
xlii., 1 : 1914-17(1914-17).
SOCIETE EnTOMOLOGIQUE DE FrANCE
Bulletin, 1917, 15-21, Tp.JL-c: 1918, 5-14(1917-18).
SOCIETE ZOOLOGIQUE DE FrANCE
Bulletin, xxxix;xl., 1-10,'r.p.ctc.: xli. 1-10, T.p.itcd 914-16).
Pavia
IsriTUTO Botanico dell" R. Universita di Pavia^
Atti. ii. Serie. Vol. xv., l( 1 918).
Perth, W.A.
Geological Survey of West Australia' —
Bulletin, Nos.71 (Text and Atlas), 73, 74, 76(1917).
Government Statistician, West Australia —
Quarterly Statistical Abstract. Nos.207-21 1( 1917-18).
Philadelphia.
Academy of Natural Sciences —
Proceedings. Ixix., 1-2; Ixx., 1(1917-18).
Entomological News, and Proceedings of the Entomological
Section of the Academy of Natural Sciences, xxix., 1
(1918).
donations and exchanges. 897
American Philosophical Society —
Proceedings. Ivi., 1-7, List, T.p.ctc: Ivii., 2-4(1917-18).
Zoological Society of Philadelphia —
Forty-sixth Annual Report of the Board of Directors, April,
19^8(1918).
Plymouth.
Makink Biological Association of thk United Kingdom —
Journal. N.S. xi., 3-4(1917-18).
Pusa, India.
Agricultural Research Institute —
Memoirs of the Department of Agriculture in India. — Bac-
teriological Series, i., 7(1917). — Botanical Series, ix., 2
(1917). - Entomological Series, v ,4(1 917).— Bulletin, Nos.
72, 76, 77(1917-18).— Report of the Agricultural Research
Institute and College, Push, 1916-17(1917). — Report on
the Progress of Agriculture in India for 1916-17(1918). —
Report of Proceedings of Second Entomological Meeting,
February, 1917(1917).
Richmond, NSW,
Hawkesbury Agricultural College —
H. A. C. Journal, xiv., 12: xv., 1-11(1917-18).
Rio de Janeiro.
Escola Supekiok de Agkicultura e Medicina Veterinaria—
Archivos. i., 1(1917).
Instituto Oswaldo Ckuz —
Memorias. viii., 2-3; ix., 1(1917-18)-
Riverside, Cal.
Graduate School of Tropical Agriculture, and Citrus
Experiment Station (College of Agriculture, Uni-
versity OF California) —
Publications. Vols. i.-ii.(15 Nos.) — Bulletin No. 23a, College
of Agriculture, Agricultural Experiment Station, Uni-
versity of California Publications (1912;.
898 DONATIONS AND EXCHANGES.
San Francisco.
California Academy of Natural Sciences —
Proceedings. Fourth Series, ii., Pt. i., No.U; Pt. ii., No. 12;
T.p. of Vols, iv.-v.; vi., 8-9, T.p.&e.; vii., 1-13; viii., 1-4
(1917-18).
Sendai, Japan.
ToHOKu Imperial University —
Science Reports. Second Series (Geology), iii., 2, T.p.tfec;
iv., 3; v., 1(1918).
Stockhoim.
Entomologiska Foreningen I Stockholm —
Entoinologisk Tidskrift. xxxviii., 1-4(1917).
Sydney.
Australasian Antarctic Expedition, 1911-14 —
Scientific Reports. Series C : Zoology and Botany. Vols,
iii., 2; iv., 3; v., 2-6; vi., 1 (1918). [deceived through the
University of Sydney\
Australian Museum —
Annual Report, 1916-17(1917)
Records, xii., 1-6(1917-18).
Botanic Gardens and Domains, Sydney —
Annual Report for 1916(1918).
Flora of the Northern Territory. By A. J. Evvart and O.
B. Davies. With Appendices by J. H. Maiden, A. A.
Hamilton, and E. Cheel. (Melbourne, 1917)
Critical Revision of the Genus Eiicalyptus . iii., T.p.&c;
iv., 3-5(1917-18). By J. H. Maiden, Government
Botanist, (fee.
Bureau of Statistics —
Official Year Book of New South Wales, 1916, Nos.17-20;
1917, Nos.l-16(1917-18).
Chief Secretary's Department : Fisheries —
Report on the Fisheries of New South Wales for the Year
1917(1918).
donations and exchanges. 899
Department of Agriculture, N.S.W. —
Agricultural Gazette of New South Wales, xxviii., 12, T.p.
ifec.;xxix., 1-11(1917-18).— Science Bulletin. No.l5(1918).
Department of Public Health—
Report of the Director General for the year 1916(1918).
Education Department —
Education Gazette of New South Wales, xi , 12; xii., 1-11
and two Supplenieiits(19 17-18j.
Education Department : Teachers' College —
Records of the Education Society. Nos.28, 31-34, 36, 38
(1917-18).
Education Department: Technical Education Branch —
Annual Report, 1916(1917).
Technical Gazette of New South Wales, vii , 2; viii., 1
(1917-18).
Forestry Commission of New South Wales —
Australian Forestry Journal, i., 1-4(1918).
Bulletin. No.l3(Aug., 1918).
Maiden's "Forest Flora of New South Wales." vii. ,2( 1918).
Report, 1916-17(1918).
"Sylviculture." Chapter i.(1918).
Microscopical Society of New South Wales —
Rules, cfec.(1918).
New South Wales Naturalists' Society —
"Australian Naturalist," iii., 15; iv., 2-4(1917-18).
Royal Society of New South Wales —
Journal and Proceedings, li., 1917 (1918). — Presidential
Address by Dr. J. B. Cleland, May 1st, 1918(1918).
Royal Zoological Society op New South Wales —
"Australian Zoologist." i., 4-5(1917-18).
"Scientific Australian," xxiii., 2-4; xxiv., 1(1917-18). From
the Publishers (Bishop Bros.).
University of Sydney —
Calendar, 1918(1918).
900 DONATIONS AND EXCHANGES.
Taihoku.
Bureau OF Productive Industries, Government of Formosa,
Japan —
Icones Plaiitarum Formosanarum. Vol. vii.(1918).
Tokyo.
College of Science, Imperial University of Tokyo —
Journal, xxxvii,, 9-10, T.p.&c; xxxviii , 6, T.p.tfcc; xxxix.,
7 8; xL, 1-6, 8; xli., 1-3(1917-18).
Tokyo Zoological Society —
Annotationes Zoologicje Japoneiises. ix., 4(1918).
Toronto.
Royal Canadian Institute —
Transactions, xi., 2, T.i).ci:c.( 1 917).
ToTvnsville.
Australian Institute of Tropical Medicine —
Collected Papers, No.2(1917).
Tring, Herts.
Zoological Museum —
Novitates Zoologicae. xxiv., 3(1917).
Tunis.
Institut Pasteur de Tunis—
Archives. T. x., 3(1918).
Urbana, 111.
University of Illinois —
Illinois Biological Monographs, iii., 2-4, T. p. c^l^c. (1916-17).
List of Books and Articles published by the Corps of In-
struction, University of Illinois, May, 1915-April, 1916
(n.d.).
Washington, D.C
Bureau of American Ethnology —
Bulletin. Nos.63, 66(191 7-18).
Carnegie Institute of Washington —
Department of Experimental Evolution: Annual Report of
the Director, 1917 [Reprint from the Year Book, No. xvi J
(1917).
DONATIONS AND EXCHANGES 901
Depaktmkxt of rioMMRRCE : U. S, Coast and Geodetic
Survey —
Hypsometrv : " Use of mean Sea-level as the Datum for
Elevations." By E. Lester Jones. [Serial No. 60: Special
Publication No.41](1918).
National Academy of Sciences —
Proceedings, iii., 11-12; iv., 1-9(1917-18).
U. S. Department of Agriculture—
Bulletin. Nos4S0, 491. 535, 536, 564, 571, 598, G09, 616,
640, 643, 645, 647, 671, 689, 707, 723, 731(1917-18).
Entomological llepiints (13) from the Journal of Agiicul-
tural Research, xi., 3, 8: xii., 2, 5, 9; xiii., l(two), 3, 1 1 ;
xiv., 2, 3, 5, 9(1917-18).
Separate, No.747 : " A wasted Sugar-Supply," hy E. F.
Phillips [Year Book, 1917].
Year Book, 1917(1918),
U. S. National Museum-
Bulletin. Nos.71 Pt. vi., 98,99 Pt. i., 100 Vol. i., Pts. i.-iii.,
101, 102 Pts i.-iii., vi., 104 Pt. i.(1917 18).
Contributions from the U. S. National Herbarium. T.p.ttc.
of Vol. xviii.(1914-17); xx., 3(1918).
Proceedings. li.-liii.(1917).
Report for Year ending June 30th, 1916(1917).
Wellington, N Z.
Dkpartment of Minks: Nkw Zkalano GeolocxIcal Survky —
Bulletin. N.S. Nos. 19-20(1 9 I 8).
Palseontological Bulletin. No.6(1917).
New Zealand IJoard of Science and Art —
Bulletin, Xo.l(1918).
N.Z. Journal of Science and Technology, i., 1 4(1918).
New Zealand Institute —
Transactions and Proceedings, xlix., ]., 1916-17(1917-18).
902 DONATIONS AND EXCHANGES.
PRiVATK DONORS (and authors, unless otherwise stated).
Janet, C, Paris. -Four Pamphlets: (a) "L'Alternance Sporo-
phyto-Gametophytique de Generations chez les Algues "
(Limoges; 1914). — (b) " Documents sur la Guerre, Bulletin
d'information public par la Chambrede Commerce de Paris,"
Nos.20, 21 (1915).— (c) ^'Note preliminaire sur I'Oeuf du
Volvox globator" (Limoges, 1914).
Meyrick, E., B.A, F.R.S., Marlborough, England —" Exotic
Microlepidoptera." Vol. ii., 1-5(1916-18).
RoBKRTSON, Professor T. Brailsford, University of California,
Berkeley — Two Reprints : (a) " The Strategics of Scientitic
Investigation" [Scientific Monthly, Dec. 1916]; (b) "The
Utilisation of Patents for the Promotion of Research "
[Univ. of California Chronicle, Vol. xix., No. 4].
Smith, R. Greig, D.Sc, Sydney (donor) — Report of the Dove
Marine Laboratory, Cullercoats, Cumberland, 1916-17
(1917); Pamphlet: "Agriculture in Japan," by Professor
Koide (Sydney, 1918).
Veitch, R., B.Sc, F.E.S., Fiji— Agricultural Report No.3, of
the Colonial Sugar-Refining Company, Ltd., Fiji ["The
Cane Beetle Borer in Australia"] (Sydney, 1917).
Waters, A. W., F.L.S., F.G.S., Bournemouth — One Reprint:
" Some Collections of the Littoral Marine Fauna of the Cape
Verde Islands, &c. — Bryozoa" [Journ. Linn. Soc, Zoology,
XXXV., May, 1918].
[Printed off, March 21st, 1919.
(liJlS.)
(a) GENERAL INDEX.
Address, Presidential. Afareh 27tli,
1918, 1.
Agaric, blowflies bred from, ex-
hibited, 415.
Asariens, inverted hymeninm in,
883.
Alder-flies, descriptions of Aus-
tralian, 819.
Algje. new and rare freshwater.
497.
Angophoras, shoot-bearing tnm<jnrs
of, 191.
Apparatus, wing-coupling of Lepi-
doptera. etc.. 286 — External
brea tiling-, of larvre of jNfuscoid
flies, 658.
Arcella, seasonal distribution of.
237.
Armistice, and the cessation of
war, 712.
Australia, Lower Afesozoic Rocks
of, 37 — Carboniferous Trilobites
of, 437 — Proposed "miniature"
on Pulbah Island, 416, 495.
Australian Alderflies, 819 — Coleop-
tera. 673— Culicidte, 826 — ^Nlecop-
tera, 395— Megaloptera, 819—
Neuroptera, No. 5, 116; No. 6,
750; No. 7, 787— Sawflies, 668.
Balance Sheet, etc.. 1917, 30.
P>anksia. peculiar roots of. exhibit-
ed, 235.
Beans. Soy. seeds of. exhibited.
611.
Beetles, introduced, of economic
importance, exhibited. 361.
Benson, W. N.. Geology and Petro-
logy of the Great Serpentine Belt
of N.S. Wales. Pt. vii.. The
Geology of the Loomberah Dis-
trict and a portion of the Goonoo
Goonoo Estate. With two Pa-
Ifeontological Appendices by F.
Chapman, 320, 363— Pt. viii..
The Extension of the Great Ser-
l^entine Belt from the Nundle
District to the Coast, 593.
"Black" Si)ot of Citrus fruits, 868.
Blowflies, bred from an Agaric, ex
hibited. 415.
Brachyscelid galls, exhibited. 361.
416.
Bramble, cut-leaved, exhibited. 235.
P.reathing-apparatus. external, of
la r Vie of ^Nluscoid flies, 658.
Bretnall. P. W., welcomed on his
return from the War, 415.
Brotherus, V. F., and W. W. Watts,
The Aiosses of N. Queensland,
544.
Caddisflies. Canadian, exhibited,
612.
Callistemon fruits showing de-
layed dehiscence, exhibited, 713.
Cambage. E. H., Notes on the
Native Flora of N.S. Wales. Pt.
X., The Federal Capital Terri-
tory, 673.
Campbell, J. H., Hon. Treasurer's
Financial Statement, and Bal-
ance Sheet, 30, 31— Re-elected
Hon. Treasurer, 135.
Canada, Lepidoptera. Caddisflies.
and Lacewings from, exhibite<l.
611. 612.
Carboniferous insects, affinities of.
123— Trilobites of Australia, 437.
Chactctes, Note on a new species
of, 392.
Chapman, F.. Devonian Foramini-
fera. Tamworth District, N.S.W.,
385 — Note on a new species of
Chaetetes, 392.
, H. G., Presidential
Address, 1 — See Exhibits.
(heel. E., sec Exhibits.
11.
INDEX.
Chorhta aiistraUs, win.2:-venatioii
of. 395.
Citrus fruits. Black Spot of. 868.
Cleland. J. P>.. remarks on a Little
Peng-uin seen in Xeutral Bay,
495— A'sJre Exhibits.
("limbins Plants, stems of. 600.
Coal-seam, fossil insect-wins from
roof of. 260.
"Cobblers' Pegs" (Erif/rrnn). two
forms of, exhibited. 610.
Coleoptera, descriptions of new
Species. 715.
Collins, ]\[iss ^[. I.. On the leaf-
anatomy of f^raerola era ssi folia,
with special reference to the
Ei)idermal Secretion. 247.
Connnentry. affinities of fossil in-
sects from. 123.
Complex, Panorpoid. Introduction.
265: Pt. i.. 286: Pt. ii.. 626.
Coi)epod, ori.irin of yolk in the ova
of a. 136.
Correlation of Lower ^Niesozoic
liocks of Australia, 37.
Cubitus, structure of. in win.2:s of
^Fyrmeleontidpp, 116.
Cnlicidre. contril»utions to a know-
led.2:e of. 826.
Dehiscence, delayed, of Cnlliste-
, mon-fruits, 712.
Devonian Poraminifera, 385.
Distril)ution of Lower IMesozoic
Pocks of Queensland. 37 — Sea-
sonal, of Arcella. 237.
Dodd. F. P.. elected a Member, 416.
Donations and Exchanges. 36. 135.
234, 285, 362, 416, 495, 610. 713.
Dragonfly, sections of caudal gills
of. exhibited. 495 — Large, picked
up on the pavement. 612.
Duck, abnormal gizzard of, ex-
hibited. 361— Xote on. 415.
Dun. W. S.. re-elected a Vice-Pre-
sident. 135,
DunstaniidtH. the family, 568.
Efhidna, Note on the temperature
of, 844.
Elections. 135. 363, 416.
Elcftrical conductivity of milk.
613.
Eiiidermal secretion of t^rarmla.
247.
" Epigraphie Medicale," exhibited.
415,
Erifirrov. two foi'ms of, exhibited.
610.
Eucalypts. shoot-])earing tumours
of, 191.
Eucalyptus, a new Species or Form
of, 747,
Exhibits : —
Chapman, H. C samples of
yeast 612.
Cheel. E., ])eculiar roots of
Fakca. Banksia. and CrcrU-
Ica : seedlings of TcphroRia :
two forms of O.ralia conucii-
lata : cut-leaved or parsley-
leaved bramble ; /?o.*f< hrarfeafa:
flowers of ^ti/phelia tuhi flora.
varying in colour, 235-236 —
T'\^'o forms of "Cobblers'
Pegs" : seeds of Soy Beans.
610-611 — C alii fit r m o n /> a oh //-
phifllN!^. showing delayed de-
hiscence of the fruits. 712.
Cleland, J. B„ four Parts of
Blanchard's " Epigraphie Mtkli-
cale": blowflies bred from an
Agaric, 415,
Fletcher, J. J., Brachysctdid
galls. 361 — Plants, other than
Eucalyjits and Angophoras.
with shoot-bearing tumours.
41.5 — Melaleuca Deanei and
Petrophila .<<e.'^.<iilis, 416 — Water-
storing Mallee-roots, and
water therefrom, 496.
Froggatt. .T. L., House-flies bred
from rotten liver, 361.
, W. W.. Immature Spiny
Stick-Tnsect ; grass infested
with larv.ne of a Cecidomyid
fly, 234 — Three introduced
beetles of economic import-
ance, 816 — Brachyscelid galls ;
a Californian, Cynipid gall ;
and an undetermined living
snake. 416 — An Indian wood-
borer introduced in the wood
of boxes, 610.
I
tNDtX.
Exhibits :—
Hedk-y. C, i>lH.t(.,-r;ii)li <.f ilic
cenotaph in iin'iiiory of Major
and Mrs. limes, in the old
churcli at Port .Ma((iuarie. 2;;i).
Hull. A. F. 15.. a collection of
Land and I'resh water Mol-
lusks collected in France, .'ioj.
8teel. T.. an abnormal j;izzard of
a Muscovy duck, 301. 415.
Tillyird. II. J., win^' of fossil in-
sect from ( oal-seam, 361 —
Sections of the Saccoid Caudal
CJills of tlie larva of a Java-
nese Dragonfly, 49o — ( anadian
Lacewiufjs and Caddisflies ; a
larj;e Dragonfly ] ticked up on
the pavement. 012.
i'urner. A. J., a collection of
Canadian J>epidoptera. 01 J.
, F.. an acclimatised.
Abyssinian grass, 234 — An in-
troduced, European (ieraniuni.
301 — A rare Xew South Wales
plant {TrkninUnn), 010 —
A teratolo.gical specimen of aji
introduced IMantago. 713.
W aite, E. E., the first J'art of the
Kecords of the S. A. Museum.
230.
Fat-content of milk, 013.
J'ederal Capital lerritory. flora of,
673.
Fellows. Finnean-Macleay, sum-
maries of yeai's work, 11-13.
J'ellowships. appointments to, 13 —
Announcements. 610. 713.
Fletcher, J. J., scv Exhibits.
— and C . T. Musson, On cer-
tain Shoot-bearing Tumours of
JCucalypts and Angophoras; and
their mo<lifying influence on the
natural (irowtli-lial)it of the
IMants. I'Jl.
Flies. House, lued from liver, ex-
hibited, 301 — Flies, Muscoid, ex-
ternal breathing-apparatus of
larvjc of, 658.
Flora of Lower Mesozoic Hocks. 37
-Of New South Wales, 073— Of
Federal Capital territory, 073.
Foraminifera, Devonian, 385.
France, mollusca from, exiiibited.
235.
Froggatt. J. L.. A Study of the
External rji-eathiug-appai-atus of
ilie la rv;r of some ]Muscoid F'lies,
05S— ,svr Flxhibits.
. W. W.. re-elected a Vicc-
Tresident. 135 — Remarks on
migrations of Mice, preyed ui'on
ly Marsupials. 361 — Notes on
Australian Sawfiies (Tciithrcdi-
iiidar). 668,— >S'cc I'lvhibits.
! luii. i>ies(rved. springiuir of tins
of, 409.
on Live Oak, ex-
<iall. ( ynipid
hibited. 416.
(ialls. Brachyscelid. exhibited. 36 1
416.
Oeology of the Lowei- Mesozoi*-
Kocks of Queensland. 37--Of the
(ireat Serpeiitine Delt of N. S.
Wales. 320. 363. 593— Of th<'
Loomberah District and a por-
tion of the (Joonoo (Jooiioo F^s-
tate. 320. 363.
<l(raniu)ji niollc. spe'imen of. ex-
hibited. 361.
<iillies, C. D.. elected a Afember.
135 — On the Seasonal Distrilm-
tion of some Queensland Species
of Arcclla. 237.
(.ills, section of ( audal. of Drag-
onfly, exhibited. 495.
(iizzard. abnormal, of a du<k. ex-
hibited. 361 — Note on. 415.
•*(;iow-worms.'" remarks on. 236.
(ioldflnch. Lieut., (i., news of, 285
- Letter from. 712.
(Joonoo (ioonoo Estate, geoloirv of,
320. 363.
(irass. Abyssinian.
(Ji-ass-fly. larv;c of.
(irevilk'fi. peculiar
hibited. 235.
(irowth-habit of plants modified b\
slioot-bearing tumours, 191.
Hakea, peculiar roots of. exhibited.
2 3 5
Hall. ('.. On a new Species or Form
of Eucalyptus, 747.
exhibited. 234.
exhibited. 234.
roots of, ex-
IJ^DEX.
Hallmaiiii. E. F.. Liiniean Maeleay
Fellow ill Zoology : summary of
year's work, 11 — lietiremeut, 11.
Hamiltoii. A. G., elected a Vice-
J 'resident, 135 — llemarks on
"Cjlow-worms," 236.
Hedley, C, elected a Vice-Presi-
dent, 135.— See Exhibits.
Jlvniiptera Heteroptera, fossil, of
Queensland, 568 — Note on the
origin of the Heteroptera, 568.
llolmetabola. phylogeny of, 265,
Hopson, J., Junr,, elected a Mem-
ber, 416,
llouseflies bred from liver, exhibit-
ed, 361.
Hull, A, F. U., ,b'ct' Exhibits.
Hymeuium, inverted, in Agaricus.
883.
India, wood-borer introduced from,
exhibited, 610.
Innes. Major and Mrs., photograph
of cenotaph in memory of, ex-
hibited, 236.
Inositol from an Australian plant,
850.
insects, Carboniferous, from Com-
meiitry, 123 — Holometabolous,
phytogeny of, 265 — Mesozoic, of
Queensland, 117, 568,
Insect-wing, fossil, from roof of
coal-seam, 260.
Johnston. S, J,, congratulations to,
135; thanks from, 234,
Kesteveii, H. L., The Origin of Yolk
in the Ova of an Endoparasitic
C'opepod, 136,
Lacewiiigs, Canadian, exhibited,
612,
Lambert, C. A., and J. Shirley,
The Steins of Climbing I'lants.
600.
Larvie of Muscoid Hies, external
breathing api)aratus of, 658—
Feeding on liver, 361,
Lea, A, M., Descriptions of new
Species of Australian Coleop-
tera, xiv., 715.
Leaf-anatomy of Scaevohi, 247,
Lepidoptera, wing-coupling appa-
ratus of. 286 — Canadian, ex-
hibited, 611.
L'Estrange, W. W,, and Iv. G,
Smith, The "Springing of Tins
of Preserved Fruit, 409.
Life-history of Psychopsia eltfjuns,
787.
Loomberah District, geology of,
320, 363.
iMacquarie. Lake, proposed "minia-
ture Austa-alia" at, 416, 495,
Mallee, water-storing roots of, and
water therefrom, exhibited, 496,
Marsupials reported as preying on
mice, 361.
^Nlecoptera. studies in, 395,
Megaloptera, Australian. 819.
Melaleuca Deanei, exhibited, 416.
Mesozoic Insects of Queensland,
No. 3, 417 ; No. 4, 568.
Mesozoic, Lower, Eocks of Queens-
land, 37.
Methyl Ifjevo-inositol from an Aus-
tralian plant, 850.
Mice, recent migrations of, report-
ed, 361.
Milk, relation between fat-content
and electrical conductivity of.
613.
Mitchell, J,, The Carboniferous
Trilobites of Australia, 437,
Mollusca from France, exhibited,
235,
.Mosses of North Queensland, 544,
Museum, South Australian, Pe-
cords of, exhibited, 236,
Musson. C, T., and J. J. Fletcher,
On certain Shoot-bearing Tum-
ours of Eucalypts and Ango-
phoras; and their modifying
influence on the natural Growth-
habit of the Plants, 191,
Myrmeleontidie, structure of cubi-
tus in, 116.
Neuroptera, studies in Australian,
116. 750, 787.
Neuropteroidea, phylogeny of, 265.
New South Wales, Serpentine Belt
of, 320, 363, 593- Notes on flora
of, 673.
INDEX.
Xote oil a new species of Chactctcs,
392—011 the origin of the Hete-
roptera, 568 — On the tempera-
ture of Echidna, 84-t.
Notes and Exhibits. 235, 361, 415,
416, 495, 610, 713.
Notes on native Flvni of New
South Wales, 673.
Odonata, Mesozoic, 417.
Origin of yolli in ova of a Cope-
pod, 136— Of the lleteroptera,
note on. 568.
Ova of Copepod, origin of yolk in,
136.
Odalis condculata, two forms of.
exhibited, 235.
J'anorpoid Complex. 265, 286, 626.
Panorpoidea. phylogeny of, 265.
JVtrel. Little, seen in Neutral liay,
495.
I'etrie. J. M., Linnean Macleay
Fellow in Biochemistry : sum-
mary of year's work, 11 — lie-
appointed. 1819-19. 13.
. The Occurrence of JNIethyl
Lievo-inositol in an Australian
Poisonous Plant. 850.
i'etrology of the Cireat Serpentine
Belt. 320, 363. 593.
I'ctroijhUu- aesiiUiff, exhibited, 416.
I'lwma vitricarpa, life-history of.
868.
Phylogeny of Holometabolous in-
sects. 265.
Plantago, teratological, exhibited,
713.
]*lants. Climbing, stems of. 600 —
Exhibited, sec Exhibits.
Playfair, Cr. I., New and rare
Freshwater Alga?, 497.
i^oisonous plant yielding inositol.
850.
l*riestley, H., elected a Member.
135.
I'rotodonata. Mesozoic. 417.
Psychopsidic. the family, 750.
Psj/chopsis (Jcquns, life-historv of.
787.
Pulbali Island, proposed "minia-
ture Australia" on, 416, 495.
(Mieensland. Lower :Mesozoic Ilocks.
37— Species of Arcella, 237 —
Mesozoic Insects, 417, 568—
North, Mosses, 544.
Becords of S. A. Museum, exhibit,
ed, 236.
Boots, peculiar, exhibited, 235—
Waterstoring, of a Mallee, ex-
hibited, 496.
Ifom hracteafa. exhibited, 236.
Sawtlies. Australian, notes on, 663.
Scievola. leaf-anatomy of, 247.
Search for toxin <i)roducers, 142.
Secretion, epidermal of Sctevola,
247.
Seedlings with peculiar roots, ex-
hibited ; of Tephrosia. exhibited.
235— Of other plants than Euca-
lypts and Angophoras. with
shoot-bearing tumours, exhibit-
ed. 416.
Serpentine Belt of N. S. Wales,
320, 363, 593.
Sherrie. ]\Iiss H., elected a Mem-
ber, 362.
Shirley. J., tind C. A. Lambert, The
Stems of Climbing Plants, 600.
Smith. (;. P. D., An Account of
s(m»e Observations upon the
Life-history of Phnma citricarpd
McAlp.. the cause of the "Black
Spot" of Citrus-fruit in N. S.
^\'ales, 868— The Occurrence of
an inverted Hymenium in Agari-
cus cam pest ri.s, 883.
.. B. CU-eig. Macleay Ba<-
teriologist to the Society: sum-
mary of year's work. 11 — Con-
tril»utions to our Knowledge of
Soil-fertility. No. xvi. " The
Search for Toxin-producers, 142
- IJemarks on water from Mal-
lee-roots, 496.
• and W. W.
L'Estrange. The "Springing" of
Tins of Preserved Fruit. 409.
Snake, livhig. exhibited. 416.
Soil-fertility, contributions to a
knowledge of. 142.
Soy Bean seeds, exhibited, 611.
i^DEX.
"Sprin^^iiij;" of tins of iireserved
fruit. 401).
Steel. T.. note on an abnormal giz-
zard of a duck. 415 — »Vrc Ex
liibits.
Stems of climbiny: plants. GOO.
Stick-inseet. immature, exhibited.
234.
Studies in Australian Xeuroptera.
116, 750, 787— ^te<X)ptera. 395.
i<t!ii)h('}i<i tiihiflora, flowers of.
variable in colour, exliibite<I.
236.
Sydney Harbour Colliery, fossil
insect-wing from. 260.
'lamwortli district, Devonian Fora-
minifera from the, 385.
Taylor. F. H.. Contributions to a
knowledge of Australian Culi-
cida?, No. iv.. 826.
T. C, congratulations to,
234~thanks from, 285.
Temperature of Echidna. 844.
Tephrosia, seedling of, exhibited,
235.
Tillyard. E. J., Linnean Macleay
Fellow in Zoology: summary of
year's work. 12 — Ke-appointed.
1918-19, 13— Congratulations to,
135— aS'cc Exhibits.
Studies in Australian
Neuroptera. Xo. 5. The Struc-
ture of the Cubitus in the Wings
of the Myrmeleontid;e, 116 ; N(*.
6. The Family Psychopsidie.
with Descriptions of new Cenera
and Species, 750; Xo. 7. Tlie
Life-history of Psi/rhopsis vl' -
(jana, 787— On the Affinities of
two interesting Fossil Insects
from the Upper Carlxmiferous of
('(mimentry. France. 123 — A Fos-
sil Insect-wing from the roof of
the Coal-seam in the Sydney
Harbour Colliery, 260— The
I'anorpoid C omplex : a study of
the Phylogeny of the Holometa
bolous Insects, with special re
ference to the Subclasses Panor-
poidra and yeuropteroidca [In-
troduction], 265; Pt. i.. The
Wing-coupling Apparatus, with
special reference to the Lepidop-
trra. 286; Pt. ii., The Wing-tri-
chiation and its Eelationship to
the Ceneral Scheme of Venation,
626 — Studies in Australian
Mecoptera. No. ii.. The Wing-
Venation of Chorista australia
Klug. 395 — Mesozoic Insects of
(Queensland. No. 3, Odonnta
and Protodouata. 417; No. 4.
ffoniptera Heteroptera : The
Family Dunstaniidae. With a
Note on the Origin of the
Ifctcrnptcra, 568 — Australian
Mcf/aloptera or Alder-flies; with
Descriptions of new Genera and
Species. SI 9.
Toxin-producers, search for, 142.
Trichiation of insect-wings. 626.
Trichinium. a rare species of, ex
hibited. 610.
Trilobites. Carboniferous. 437.
Tumours, shoot-bearing. 191, 415.
Turner. A. J., ace Exhibits.
, F., sec Exhibits.
Waite. E. U.. sec Exhibits.
Walkom. A. B., The Geology of the
Fower ^lesozoic Eocks of
Queensland, with special Kefer-
ence to their Distribution and
Fossil Flora, and their ( orrela-
tion with the Lower Mesozoic
Kocks of other parts of Aus-
tralia. 37 — Congratulations to,
135; thanks from. 234.
Ward law, H. S. H.. Linnean ISIac-
leay IVllow in Physiology: sum-
mary of year's work, 12 — Re
appointed. 13.
— . 'i'he IJelatiou be-
tween the Fat-content and the
Electrical C onductivity of Milk.
613 — Note on the Temperature'of
Evhidna uculcata, 844.
Water from ^Iallee-rt>ots. exhibit-
ed. 496; remarks ou, 496.
Water-storing roots of Mallee. ex
hibited. 496.
Watts. W. W.. and ^'. F. J3rotherus.
The Mosses of N. Queensland,
544.
INDEX. ni.
Wiiit: of fossil iiis«^<t from C'oji] Myrmeleontidce. 116.
seam. :.'60 ; t'.xliibittMl. :{(51. \\ <»o<lI><M-er. iiitro(lu<e«l. Indian, ex-
\Vin:z-coui)lin;r apparatus of in- Inlnted, 610.
se<ts. :,'S6 — Winir-trichiation and
venation. G:.M» Win ^'-venation of ^'east. samples of. exhibited, 612.
Choriata. :59.'). "^'olk. ori;rin of. in ova of a Cope-
W inu's. structure of cubitus of. in pod, 136.
(/>) BIOLOGICAL INDEX.
Names in italics are synonyms.
I'AGE. Page.
Abrus i»recatorius 601 Adenocalymma nitidum .. .. 600
A(
acia
710
armata
704
buxifolia
704
Dawsoni
704
dealbata
..
704
decurren.s r. mollis .
704
diffusa . . ...
704
704
implexa
..
laniirera r. venulosa .
. . .
704
nielaiH»xylon
684.
704
obtusata
696.
704
)>enninervLs
690,
704
jtravissima
, 696,
704
rubi<la
704
sicnliformis
704
rvuulo>i<i
704
verui<iflna
704
,\(
•;ena ovina
704
sanjzuisorbie
704
A(
antbaceie
600
A(
■antliaclisis
119.
122
A(
•antliocladium extenuatum,
561
sp. (?)
561
A(
•ilihylla simplicifolia . .
691.
700.
707,
711
\<
•ri<lii<he
'^63
\(
ridioidea
■'61
A(
•rotriche a;jj;rewita
690,
707
serrulata
707
A<
•t inocystis cornu-bovis
595
A(
tinomyces .. 117. 149.
1.55,
166
cbromojrena
147.
148
odorifera
147,
148
A(
tinostroma australe. .
336,
380.
381.
382,
384
clatliratum
382
.\dia utiles lindsa.voides .... 89
Adiantum a*thiopicum . . . . 701
Aeroluyopsis sijrmatopbylla. . 559
Aeroplana . . . . . . . . 426, 431, 434
mirabilis . . 426, 427, 429, 430,
433, 435
Aeroplauida' 435
Aeroplanoptera 425, 434
.ll^schna 436
flinder^^ii HHis 436
.Eschnidiid.'B 423
^^i^schnidiopsis 436
flindersiensis 435,436
Agaricace;e 884. 885, 886
Apiricus campestris 883
Agrionidie 421, 434
A.irr<»myzidtp 688
Aira caryophyllea 702
Ajuixa australis 708
Albertia australis 90
Alethopteris 56
australis 85. 86, 87
sp 85. 91
Aljrie 497
Alisma i>lantaj:o 702
Alismatace-cV 702
Alveolites depressa 393
sp 331, 595
Amaryllida<ea' 703
Amceba 172, 173
Umax. 176. 17 7. 180. 181, 189
\m|»elidea> 600
Amphientomum. . .. 132. 133. 131
paradoxum 130,131
Amplexiformia 297
Vlll.
INDEX.
Page.
Aimbieiia 5<>3
australica 503. 541
r. coiistricta . . . . 504, 541
laxa 505
V. hortensis 505. 541
oscillarioides 507
r. crassa 504. 5 41
r. steiiospora 507
variabilis 504
r. cyliiidracea . . .. 504. 541
Volzii 507
Aiiaj;allis arveiisis 708
Aiiastellorhina aiij;iir . . 415. 658.
6G2. 664. 666, 667
Amlropogon affinis 702
refractus 702
Aiigophora 191, 227, 688
c'ordifolia 227
iiitormedia 227. 803
laiiceolata .. .. 212, 227, 232
siibvelutina 227
Aiijruillaria dioii-a 702
Aiiobium domesticum . . . . 361
Aiioniobryiim <-ymbifoliimi . . 554
AnomodoH Torcoac 563
Anopheles 826, 827
aiiniilipes 827, 828
atratipes 827
barbirostris 827
r. bancrofti . . . . 827, 843
corethroides 826. 827
puiHtiilatiis 826
stij^matieus 826, 827
Anthistifiii cilkitti 702
Aiithophysa vejj;etaiis .... 508
Aiitliropliyojtsis sp 91
Anti^'oiioii leptopus 601
Aphaniptera 265. 269. 28S
Apochrysidje 638. 783
Apocyiia<-ea« 600, 862, 867
AraUa<eiH 707
AraiK-aria 75
AraiK-arioxyloii spp 05
Araiiearites 75, 90
polyearpa 72. 79. SI
sp 85
sp. A S9
sp. B 90
Arcella 237, 238
artot-rea.. 237, 238. 239. 240.
241, 242, 245, 246
deutata 237
Page.
Areella discoides . . 237, 238, 239.
240, 241, 243, 244. 245, 246
mitrata, 237, 238. 239, 240, 241,
244, 245, 246
vulgaris . 237, 238, 239, 240,
241, 242, 243, 245, 246
Arcliicliaiiliodes, 291, 635, 636, 637
dubitatiis 820
.irnttiferus, 292, 626, 636, 820
Arcliii)anorpa, 633, 635, 636, 650,
651, 653
maguiiiea , . , . 269, 631, 657
Arisoceplialus 732
basalis 733, 735
flavipes 733,735
rufipeiiiiis 733, 734
Arisiis 732
carina ticeps 735
Armigeres 829
breinli 828, 843
inalayi 829
Aroideae 601
Artcriopterij.v, 759, 760. 764, 765,
766, 768, 773
chyaus 767, 777, 784
Artliroiormiis Scliimperi, 545, 549
Arthropoda 278,280
Artliro]K)diuiii i)aiiiciilatum .. 702
Artnulo /'hnif/niites 702
Ascalaphidie \ . 291. 631, 638, 815
A.scaris 140
Aselepediaeeic 600
Ascochyta hortoruiii 875
Asparagus ra<eun)siis. . .. 601,608
Aspfi-gillus sp 147, 14S
Asperula oligantha , . , , 691, 708
r. S(x>i)aria 708
Asi)i(liKiH <icul((iti(ni 701
Aspidosperma (luebraclio, 862, 863,
867
Asplenium liabellifolium . . .. 701 j
Aster crlmi.sia 709 |
AntcrfKlictioii orafiim .. 537, 538
Astrorliizidu' 387
Astrotriclia ledifolia 707
Atrypa sp 331,336,595
Auliiia rotiformis 383 ^
Auriculariacoje 885 J
Austinia luzonensis 561 M
Australina pusilla 703 ■
Austrosialis 821, 823, 824 ■
ignicollis 822, 823 j
INDEX.
IX.
Page.
JJacilliis iiiuiR-tiis 144
mycoides 143, 169
l»rodi;j:iosiis. 14o. 147, 155. 157,
101. 108 171. 172, 18:J. 189.
190.
pyotyaueus 173
si». .. 148. 149. 15:3. 153. 154.
155. 157, 158, 161. 169
Ijackliuiisiu myrtifolia 415
JJacterium tiimefacieus . . . . 197
r.cwkea Ciuimiana,.. 690, 691, 707
iiaiera aiistralis 89
bideiis . . . . 65, 68, 70, 79, 81
j^'inkgoides 65, 79. 81
gracilis 89, 90
ipsvicieii.sis . . 65. 69, 70. 79, 81
miiltifida . . . . 69, 70, 85. 87
Simmoudsi, 65, 69, 72. 74, 76,
79, 80. 85, 87
SI) 86
stormbergeiisis 68, 70
Balmc>i 760. 761. 784, 785
liahnviiini .. 760, 761
Baiiisteria chrysopbylla . . 601, 602
LJaiiksJa margiiiata.. 695, 703. 711
serrata 235
JJarbella eiRTvis 560
I'.arbiila calyciiia 552
IJartrauiiaccte 556
Jiaidiinia corymbom 601
scaiideiLS 601
Beaiimoiitia graudifiora . , . . 600
Bedfordia saliciiia . . . . 701. 709
Beiiiiettitales 65, 72
Beiiiiettites 65. 69, 79
Carriitbersi 69
.SI) 81
Ijernardia Chodati 533
tetraedrica 533. 543
Ijerotbida- . . . . 638. 639. 783, 784
iJertya ok'a'folia 705
Jiescberolhi brvrifoliu 557
( yrtopiis 557
Bigiioiiia fioribuiida 600
picta . . . . 600. 602. 604, 609
Tweediaiia . . 600, 602, 604, 609
veiiusta 600
LUgiiouiacete 600
Billardiera scaudeus 704
Biroiiella 826. 827
gracilis 826, 827
Page.
lUttacidcC 290, 640, 656
lUecbiiiim cai)eiise 701
cartilagiueiim 701
discolor 701
peiina-mariiia 701
Borragiiiaceie 708
Bossiur'a buxifolia 705
Walkeri 705
Bostrychiis icqualis 610
Bougainvillea lateritia.. 601, 607,
609
Sanderiana 607
r. variegata . . . . 601, 609
spectabilis. 601. 602, 606. 609
Bracbycbitoii popiilneus, 697, 706
I '.racbycome discolor. . .. 691,709
scapiformis 709
scapigera . . . . 690, 691, 709
Sieberi 709
sp 690
Bradiyhcna 255
Brachyloma dai)liii< tides . . . . 707
Bracliymeiiinm acuminatum . , 554
Wattsii 554
P.rachymetopiiia 483, 485
Bracbymetopus. 438, 439. 482. 485
duustaiii.. 471, 486. 488, 492,
493. 494
lodiciisis 488
^Nfaccoyi 486
SJ). iud 488, 492
strzclcckii. 438. 439. 443, 482.
483. 486. 487. 488, 491
l!racbyi>byllum. . .. 56, 75, 85. 90
cra.ssnm . . . . 72. 75, 76, 79, 81
gippslaudicum 90
SP 90
Brachytbeciaceic 567
Bromus maximus r02
Brunelbi vulgaris 708
Brunonia 24S
Bryaceu- 554
3>i\voiiia 608
laciiiiosa . . 601, 602, 605, 609
Bryum argeuteum v. iiiveum.. 554
Baileyi 555
erythrocarpoides 554
kurambc 554
leptotbrix 555
l)impamic 555
subatropurpureum .. .. 555
subolivaceum 555
a.
IKDEX.
Page.
Bryiim siibpathyiKuna 555
Biuldleia inadajjcastarensis . . 601
BiilMiie bulhosa.. .. 689. 692, 702
BinhUUa lornnta 525
Bursa ria spiiiosa 704
•
C'aladeiiia alba 692,703
(limorplia 689.692,703
testacea 703
C alama.irrostis anuula 702
Callicostella k;erni)a(liii . . . . 562
CaUiphoni occanhic . . . . 415, 658
nififacUs 658
rariix s 658
viUom 657, 658
Callistemon lopliaiitluis. . 690, 707
pachyphylhis 713
paliulosus 707
piiiifolius 415
rigidus 713, 714
saligims 415
Sieberi 691, 707
vimiiialis 714
Callitri.s cakarata .. .. 676, 701
sp 834
Calomf/ui 838
prk'.stlrii 838
C'alopteryiridu* 495
Calotis scabiosifolia 709
/•. inU'^rifolia 709
Calycothrix tetraj,'(>iia 707
Calymperai ea' 55<>
C'alymperes Keiiiiedyanum . . 552
molucceiiso 545, 552
Motleyi 552
iiigresceiis 553
serratnm 552
teiiernm 552
(alifmiKridiiini Mudlcri .. .. 551
C alyptoiliyiK'lius viridis . . . . 694
(■alyi)t<>tbe(iniii a<iitnm .. .. 560
himiile 560
('alffthrir f<tni(/oii(( 707
CampaimbKoa' 708
( amphoiiliylliim tiexuosum .. 380
Camptocha-te biisbanka . . . . 561
vaga 561
Campylodontiimi Haves<eiis . . 561
Campylopus Wattsii 547
Woollsi 547
Caiid<>llea senulata. 690, 700, 709
Caiidulleacea.- 690, 709
Page
( aprifoliaceiE 708
Carabida" 436
( ardamine liirsiita r. teiiiii-
folia 704
tntififolia 704
( ardiophoriis 733
Cardiiiis pycnocephalus .. .. 709
Carex apiu'essa 702
(iaiidichaiidiaiia 702
pseudo-cyperiis 702
tereticaiilis 702
( ari»olithes spp 90
Carroiiia multisepalea. . 601.^602.
605' 609
Carteria 498, 514
bulhilina 516
granulosa 516, 541
multifilis 515, 516. 541
nignlosa 514, 517, 541
r. angulata 515, 541
r. iuiP(iiialis .. .. 515, 541
( aryophyllacea^ 689.703
( assiiiia aciileata 709
longifolia 709
quiiiqiiefaria 709
( assytha melaiitha 701
pha'olasia 701
Castiiiida-.. .. 297, 308. 311. 314
I'asiiariiia Ciuiiiinghamiaiia, 692.
703
glaiK-a 693
Luohmaiiiii. 682, 693. 703. 711
(juadrivalvis 703
sp 415
strict.a 692, 703
( asuariiiOce 703
Caiistis fiexuosa 691
{ ocidomyia s\> 234
Celmisia longifolia.. 691. 700, 709
Centanrea calcitrapa 709
solstitialis 709
Centipeda (unninghamii . . . . 709
Cephenode.s jauus 319
( erastium vulgatiim 703
(■(MaUKUis 9t)
C luetetes 335, 392, 393
depre.ssiis 393
Coldfussi 380
radians 392. 393, 39J
stelliformis. . 336. 380. 393. o'.)i
( lurtomitrinm entodonloides. . 562
Cieheebii 562
INDEX.
XI.
Page.
( luetomitrium nematosiim .. .")6"2
(halkodoma 278.280
( li:irn<'ium cernssifonne .. .. 529
r. miniminii . . . . 528. 529
uiittiila 528
lonsj;ii>es 529
oniithooephalnni. . .. 528. 529
l»yi-if(>rine 529
Charajria :}02. 313
eximia 302, 319
Chauliodina' 820
( heihmthes tennifolia 701
('lieii-(>lepis( ■?) setosus .... 90
( henoi»<)diaceip 703
( henoiKtdiuin tl•ian^'lllal•e .. 703
(himariha 394
(hlamvdoimnias. 498. 509. 517, 518,
532
alboviridis 520
alpina 518, 541
anjiulosa /•, obesa .. 521, 541
raiidata 518. 541
ijlobulosa 519
Lismorensis /•. i?ra<ilis. 517,
541
longistigma 523
maculata 518, 541
r. oblonga 519. 541
r. pbuiktonica. . ..519, 541
media 521
metasliicma 519, 541
paupercula 520,541
pisiformis r. ocellala. 521. 541
pusilla 520. 541
reiiiforinis 517. 541
rotiila .. . . 517. 518. 532. 541
stellata 518
subcaudata 519
( hloeoii 278
Chloiis truiuata 702
( hloroj;onium minimum.. 521, 542
Chloronionas alpma 518
Chlorophyceie 514
( horetrum spicatum 703
Chorista.. 290. 397, 399, 401, 404.
406
aiistralis.. 395. 398, 403, 405.
407, 408, 657
( boristida?.. 288, 290, 312, 316. 640
Cbrooeoecaeete 499
( hrysopa 396
Chiysopida% 633, 638. 756, 783. 791.
806, 816, 817
Page.
Cicada (?) lowei 87
Cicadid.'i- 589
Ciiicbona 603
Cla<lomiiioii palleiis 557
Cladopblebis australis, 64. 68. 70.
71. 74, 75. 78, 85. 87. 90
denticulata 74
y. australis 89. 90
Koylei.. 64. 68, 70. 71. 75, 78.
85, 87
sp 68, 85, 87
C'lavaria 885
Clematis aristata 703
microphylla 703
Closterium lunula 513
Codiolum i^r 'jj:aiium 530
('(jf'lastraceu' 540
('(TelastiMim reti<ulatum . . , . 540
('(plospermum paniculatum. . 601.
602, 603, 609
CflEHiuosa trianjiularis. 634. 635, 652
Coleoi»tera. 265. 266. 267. 268, 295,
715
Coleopteroidea 265
Collembola 799
Colletotriehum adustnm.. 868. 869
Coml)i'etaee;p 601
Comes])erma retusum.. .. 691, 705
Compositje, 601. 690. 691. 709, 710
Coui-bidium kniubtii 336
Coniferales .. .. 72. 73, 75, 78, 79
Coniojjtei'is delicatula .. 64. 78, 80
bymenophylloides r. aus-
tralica 89
Coniopterygida^ 638
Conites sp 90
ConjugatiF 510
ConvolvulaceiP 601, 708
Convolvulus erubescens . . . . 708
Copeognatha 132
Coprosma hirtella 708
Coreida? 576, 582
( orydalidsp. 291. 316. 636. 656, 819
Coi-ydalime 820
Corydalns 125
Corynopbyllus andersoni.. .. 725
fortnumi 725
interoeularis 724
modestus 725
Cotula lilicula 709
Craspedia Eichea 709
Crassulaeeae 704
Creagrini ,. .. 116. 121, 122, 130
xu.
INDEX.
Page.
Creagris 436
Crcagris 12?-, 436
plumbeus 123, 436
Creoleon ^36
Creoleontini -136
(ristellaria vetiista 386
Crncifersp T04
Crustacea 278
rryphsea tenella 556
(ryphfpacesp 556
Crypliidium ^[nelleri 557
Cryptandra spineseens 7()5
Cryptodus 723, 724
al>errans 723
cavioeps 723,724
gigas 724
grossipes 723, 724
tasmaniaiius 724
Crifptof/k'iia an(/iilo,m 522
('nipiomo)ias Lima 510
( ryptostegia grandiflora, 600, 602
Ciu-umis myriocarpus 708
f'liciirbitai-epe 601. 708
Cn]ex ahdominalia 835
bitjeiiiorhyiichus 835
foncolor 835
consimilis 835
(■)/Ut)(i)ieii.<^ 838
fatigans 836
flnrifroiis 832
fnsous 836
notoftcriptux 834
parr us 835, 836
sitiens 834. 836
tigripes 835
r. fusca 835
tigripes 835
townsvillensis 836
vishnui 835. 836
C'ulioada 826
flavifroDS 832
tasmaniensis 834
wilsoni 833. 834, 843
Culiccisa ahdominalis 835
fusca 836
("nlieidit^ 654. 656, 826
Curculionidae 746
(' lianophuccae 499
Cyatheaeete 64. 701
C'yathophyllnm 336
sp 335, 376. 377, 595
Cycadophyta. 65, 66. 69, 72, 73. 74,
78, 79
Page.
Cycadopteris soolopendrina . . 85
Cyclodictyon lepidum 561
('ylindrospermum 505.507
Goetzei 506. 507. 541
rec'taiigiilare. 505. 506. 507, 541
r. parvicelUila . . 506. 541
stagnale 505
r. australe 505. 541
Cymatop.sociis 132
Cymlxmotus Lawsonianus. . .. 709
("ynodou daet.ylon 702
Cynoglossum australe 708
Cyperaeefp 702
Cyperus Gunnii 70™*
lucidus 702
sanguineo-fuscus 702
Tyrotyphus fascieulatus . . . . 746
vai'iegatus 746
Cyrtopodacete 557
Cystiphyllum australasioum. . 595
vesuulosum 337
Cytosporella viticola 876
Daetylothece arcuatum..
Dampiera Brownii
DauaM»psis Uugliesi, 64, 68
528.
Dania sp
Danthonia carphoides
penieillata r. semiannu-
laris
Datura stramonium
Daucus braehiatus
Daviesia corymbosa
ulieina 690,
Dendroleoiitiiine 117,
Derbinae
Derris scandens
Desmodium varians
Dei/eaiia Forsteri
Dianella revoluta
tasmaniea 689,
Diaporthe
Difksonia antarctica. . .. 683.
Dicranaceie
Bkranella Dietriohitp
pycuoglossa
r. longifolia
Dieranoloma austro ,seopari-
um
dicarpum
serratum
Wattsii
542
248
78.
80
336
702
702
708
707
704
705
122
431
601
705
702
702
702
875
701
545
546
546
546
547
546
546
546
INDEX.
Xlll.
Page.
Diotyeiicnemis 728
minis 729
DiotyophylUim 68
Davidi 71. 78. SO
niirosnm, 64. 6S, 70. 71. 74. 7").
78
Diflisciis humilis 707
Dilarid.'v 638. 783
DilleniaeejH 249.706
Dilhvynia erioi folia r. phyli-
coides 705
Dittoijj/.ris hicrifi 510
I)il»hyi»hyllum porteri . . 336, 595
I'oldistiim 336
8p 595
Diplodia zese 878. 880
Diptera . . 265. 269. 272. 273. 274.
282. 294. 314, 317. 399, 400. 404,
572, 574. 635, 641. 648. 652, 653.
654. 656.
Dii)teridinjp 64, 71
Di>4faria australis 705
Disforbiiia 388
Distichophylliim Baileyaiium . . 561
Ditriclinm affine 545
Dinris maculata 703
sulpliurea 703
Dodonrpa attenimta 705
viscosa 705
DorcHH luteus 715
Dracophyllum sefiindura . . .. 700
Drppanacra 116, 120
luimilis 291
Drepanepteryx .. 116. 120, 756, 757
Drepaiiidse 307. 311
Drepanomina 116, 130
Drimys aromatica 704
Drosera aurieiilata 704
peltata 696. 704
Droseraepfp 704
Dryopteris punctata 701
Dimstania. 568. 569, 570, 571. 572.
573, 574. 575, 577. 578, 579, 580.
581, 582, 583, 584, 585, 588, 589.
590
pulehra . . 568, 576, 579, 582.
584. 586, 592.
Dunstaniidfie . . 568. 575. 576. 581.
585, 583
Dimstauiopsis, 576, 577, 578, 579.
580, 581, 582. 583, 584, 588. 591
triasslca . . 576. 577. 582, 584,
585, 586, 592
Page.
Dynastides 724
Dyscolocerus 738
porosus 727
rubriventris 728
Dysmorphoptila 587
Dystrigonistliis 738
feri'ujjjineus 739
laticollis 737
paradoxus 738
r^cliidna aculeata 844
Eoliinopogon ovatus 702
Eflimepteryx 132
Ectropotliecium serrifolium . . 563
umbifulatum 563
verrutosum 564
zollinjjeri 563
Eht^ocarpus cyaueus 415
J^Hakatothrix jjelatinosa . . 527, 542
Elateridfp 733
Eloana 262
Elcanida- 261, 262
Elcanopsis 262, 264
sydneiensis 262, 263
Eleoeliaris aouta 702
cylindrostachys 702
Embidaria 265
Embioptera 127. 128
Endopliyllum 377, 599
schlueteii 595
Endotric'liella Dietriohiiv . . .. 557
lepida 557
Endromida^ 307
Entodoutaceti* 561
Entomophthalmus,. . 731. 732, 735
Epacridacea- 690, 691, 707
Epacris brevifolia 707
micTophylla 691, 707
paludosa . . 691, 700, 707, 711
serpyllifolia 707
Epilobium glabellum 707
Epioplilebia 420, 421, 422
superstes 420
Epiophlebiina^ 422
Equisetales . . 64, 66, 67, 71, 73, 78
Equisetites columnaris . . . . 70, 73
('/. rajmahaleiisis, 71, 73. 75.
78
rotiferum.. 64. 68, 70, 71, 73,
75, 78
sp 64, 89
wonthaggiensis 89
Equisetum spp. . . . . , . 67, 85, 91
XIV.
INDEX.
Page.
l^roihtites mixta 709
quadrulentata 691.700
ICremosphi^ra 53]. o:>4
viridis 531
r. tetraedrica .. .. 5?>1. .)ir.'
Erijieron bonarionsis 611
canadensis 611
linifolius 610
Eriorrania 29S
semipurpurella 649
Eriocraniidje •^9')
I'^riostemon niyoporiodos. 690. 705
lOristalis tenax 661
i:rodium ciciitariuni 705
cyjjjnorum 705
Krythrjpa aiistralis 70S
Esrallonia 255
l':stheria 38
Cofrhlani 88
manj^aliensis 88
Eucalyptus 195. 203. 710
agrgresata 706
albens 689
amv^dalina 229. 70<i
Beliriana .. 221. 223. 229. 233
Blaklelyi 706
botryoides 229
Bridgemana 706
Cambagci 699. 706
camphora 700, 7o6
capitellata 229, S03
eiuerea 229
coriaeea.. 229, 230. 684. 69o.
691, 698, 706. 711
eoi-vmbosa. 204. 205. 208. 229.
232 803
ciebi-a 229.749
dealbata 687. 688. 706
Deanei 229
DrlegatfHsiH. 228. 698. 699, 706
dives 229. 699. 706
dumosa 220
elaeophora 699. 706
eudesmoides 226
eugenioides. 206, 207, 219, 229.
232, 233, 748, 749
eximia 208, 229, 232
fastigata . . 228, 698, 699, 706
ficifolia 230
fnitieetorum .. 221, 224. 229.
233
gigantea 228, 698, 706
Page.
I':ii(aly])tns .uiobiilns .... 230. 689
gracilis 199
(lunnii 230
b:eniast<»ma. 206. 229. 232. 68S.
699, 706. 748
beniipbloia. 204. 211. 229. 232.
233. 740
leucoxylon, 201. 219. 229. 778
longifolia 209.229.232
Lnehnianniana 229
:Nracartbiiri 229
inacrocarpa 230. 232
macroriliyncha . . . . 699. 706
macula ta 220
maculosa 229. 609. 706
^farsdeni 747, 7i9
melliodora 706
microcorys 803
Moorei.. .. 220, 229. 233, 748
n<>va-angli<a 669. 740
Obliqua 698. 74S
oleosa . . 199, 200. 220. r>26, 233
oreades 228
oralifolia 699
pallidifolia 229
lianiculata 202. 229
pilularis 228
piperita .... 206. 229. 232, 233
polyantliemos 699, 706
pruinosa 230
punctata 228, 749
regnans 699
resinfera . . 209, 229, 232. 361.
748
robusta 209, 229, 232
rostrata 201, 229, 680
rubida 229. 706
saligna 229. 803
sideropbloia 229, 748
sideroxylon. 205, 212, 214, 216.
229, 232, 233
Sieberiana 229
sp 203. 227. 228. 233, 668
scpiamosa 229
stellulata 706
Stricta.. .. 199, 220, 229, 233
Stuartiana 230, 706
tereticornis. 194. 195, 205, 211.
218, 229, 232, 233, 749
terminalis 226
vimiualis .. 201, 229, 706, 748
viridis 221. 229. 233
INDEX.
XV,
Page.
Eiioamptodon Ahielleri o47
Encnemid.'v 72.")
Eiulyptnla minor 495
Eiisrereon 587
Enomi>halii.s sp 505
Enphorlda Drnmmonrtii . . . . 705
Lathynis 705
Euphorbia cejip . . . . 705. 863. 867
Euphrasia Rrowuii 708
EupterotifUip 307
Eusc'hemon.. .. 308. 309. 310. 311
Exoc-arpus cupressiformis. 699, 703
stricta 703
Exodictyon snbscabruni . . . . 549
Exsul 573. 574
singularis 573
Extatosoma tiaratum 334
Exuviella Lima 510
Fabrionaeeae 561
Faradaya splendida .... 601. 603
Favosites basaltiea 7-. moon-
biensis 595
r-apillaris 393
orummeri 331. 333
depressus 393
Gothland ica. 331. 336. 379. 595
multitabulata . . 339, 331. 336.
379. 595
plttmani 331
salebrosa 595
sp 336. 379. 384
Festuca bromoides 703
Hookeriana 691. 703
Ficus pumila 601
FiUcales. 64. 66. 67, 68, 71, 73. 74,
78, 80
Finlaya poicilia 838
Fissidens arboreus 549
asplenioides 549
cairnensis 549
DietrichijF 550
Hollianus 550
kerianus 550
kurandae 550
microleeythis 550
pallidum 550
tenellus 550
zippelianus 550
Fissidentacese 549
Flagellaria indiea 601. 608
Flagellariere 601
Page.
Flagellata- 508
Florilmudaria floribuuda .... 559
pseudofloribunda 559
robustubi 560
Fornax castaneus 736
ma.1orinus 737
niger 735
parvulus 737
suturalis 736
Franeeia oblonira 533, 543
ovalis 533
Fraus 303
Frenata 296. 297
Frenatdc 296
Frenelopsis(?) 91
FulgoridtP 431, 587
Funariac^iie 553
Fusarium sp 159, 169
Fusicoeoum 875
Fusulina 386, 388, 389
Calba 731, 741
australitp 741
waHacei 743
(lalbodema 741
Galeola oassythoides. 601. 608. 609
Galium umbrosum 708
Garclvea pliascoides 545
Garovaglia ]\ruelleri 557
Gastrodia sesamoides 703
(ieitonoplesium cymosum. 601. 608
Geminella 498
Gentianaceae 708
Geraniaceap 705
Geranium dissectum 705
moUe 361
sp 198
Ginkgo antarctiea.. . 65. 68, 79, 81
digitata . . 65, 68, 70. 79. 81
dilatata r. lata 85
Huttoni r. magnifolia . . 74
magnifolia, 65, 68, 70, 71, 72,
74, 76, 79, 80
sp 80, 89
Ginkgoales, 65, 66, 67, 68, 73, 73.
74. 78, 79
Gleichenia dubia 85
Glenoleon 119, 122
Globigerina 386,388
Gloeocystis 518
Gloeosporium 868
Glossopteris 45, 81, 264
70
XVI.
INDEX.
Page.
CJIycine olandestiiia 705
hisi)i(lji 611
Latro)>e{iiia 705
(ilyphomitrinm coinmutatum. . 55.3
(JiiJiplialium .iapoiiicuin .. .. 709
piirpu renin 709
(iom])h<»l()l)iiim .urandiHoriim. . 704
Hiie.i^elii 704
iincinatiiiii 704
(iuiiomitiiuiii ;i<-iimiiiat um . . 553
(ioodeiiia lio<l(M-acea . . .. 690. 708
ovata 248, 259
piiiiiatifida 708
(ioodeniaf-eie. . 247. 248, 250. 253,
690, 708
rirahhamia theo])aldi .... 832, 843
Graminea^ 689. 691. 702
(iratiola nana 708
Pernviana 70S
Grevillea anstralis . . . . 691, 703
jnniperina .. .. 695, 703, 711
lani;L!:era 703
montana 235
punicea 235
rolm.sta . . 195. 862. 863, 867
(irjffithides .. 441. 447, 465, 475
ronvexicandatns. 473, 475, 489,
490, 491
dnhius, 438. 440, 441, 443, 462,
481. 494
fluhius 440
eiehiraldi .. 438. 477
globieeps 478
seminiferns, 438. 442. 447, 470
Sf'miniferuH 442, 447
V. aiistrala,n<'(i .. 439. 442,
466, 470. 480, 493, 494
sp. ind 439, 443
svveeti. . 443. 479, 480, 482, 494
Crimmiaeere 553
(iuttiferae 706
(iymnocnemia 121
(rynniosoma rotundarnm .... 662
JTakea acicnlaris 703
dac-tyloides 235
microcarpa 703
sericea 703
Halorrhasacetp 707
TTalorrhagis micrantha . . . . 707
totragyna 707
Hampeella Kurzii 557
pallens 557, 562
Page.
Haplonyrha rolossa 721
gigantea 722
nobilis 722
Haplophragmium 389
Hardenl)ergia monopliylla . . 705
lluriotuia rcticuhtta 540
Harpobirtacns tillyardi . . 290, 395
Ifansmannia P>nchii. 71, 74, 75, 78,
80
ITedycaria angnstifolia . . . . 704
('nnninghaniii 704
Tleliclirysnm 710
apienlatnm 709
baofharoides 709
hnif'teatnni 709
ledifolinm .. .. 690, 691, 709
liicidiim 709
r. viscosnm 709
I'Oi^marinifolinm r, thyr-
soidenm 709
scoi'pioides 691. ;o9
seniipapposnm 709
Stirlingii 709
Helicophynaceje 562
HcliroijJufninn (iii>iU'(ih' .. .. 562
ilelioliti's . . . . 329. 334. 336, 595
interstincta 331
porosa,. .. 331. 336, 380. 595
Heliopora 335
Ifolii>ternni antliemoides .. .. 709
diinoipholepis 709
in<annm 690, 709
Hemorobiidte. . 116. 117. 120, 290.
312, 633, 638, 759, 783, 784. 816.
817.
Hemerobins enelivagns 778
mimicns 775
Hemiopsida 729
longicoruis 726
ventralis 727
ITemiptera. 568. 570, 582, 586, 587.
588, 589, 591, 592
Hepatica^ 701
liepialidie, 295, 297, 301, 302, 303.
304, 312, 313, 315, 318, 569, 570.
572, 647, 649.
Hepialus 301
IIeri)etineni'on Tocco.np 563
Hes])eriidii' 308, 314
Heterocera 309
nctei'odendi'on olesefolinm . . 850.
856, 857, 861, 862, 863, 864, 867
INDEX.
XVll.
Page.
TTptoronenra, .. 297. .304. .308. :ill.
314, 31.-), 317. 647. 0.52
Heteroptera . . 568. 575, 576. 578.
581. 582. 586. 587. 588. 589. 590.
593.
Hovea hrasiliensis. . 862. 863. 867
[lihliertia lim^ai'is r. ohtusi-
f(»lia 7()<')
soi'ityllifolia 7o6
stricta 706
Hiiiiaiitdcladium loriforme . . 561
llil>l>(>lh)ii scrofa .. 304. 306, 319
IIil»ta^v 608
Mada biota 601. 602
Hoduesla 826
caii-iisensis 842. 843
triaii.i,nilala 842. 843
TTolciis laiiatus 702
Holomotahola. 265. 266, 267. 268.
397. 399. 626, 629. 630. 632. 633.
635. 648, 813.
Hoh.uiitrium Mnellcri 546
l>ericlia'tial«^ 54()
Ifitttnilia <.ii(/u(i 561
Honiaii(>(UMi(li(>ii «'.\liinHin .. 561
H.ibeilatiiui 561
lloinolKMira. 297. 298. 315. 317. (U7.
652
lT<.)ii(>i»l(M-a. 131. 431. 569, 570, 572,
574. 586. 587. 588. 58<). 591
Haohcrio fhibclldtd . . . . . . 561
kdrstcnitDKi 561
Ilookeriacejv 561
Uordeiini mniiiuiin 702
Ifovea linearis 69o. 705
HydviK-otyle (.'(indollci 707
laxiHora 707
irydrodictyaceje 536
lfydi'(»t;astrace;e 529
HydroidiilidiP 295
Hydropteridea' 64. 72
liydroittilidii^ 646
Hymeiioinoiuidacefe 508
Ilymenoptera. 132. 265, 266. 267.
268, 278, 280, 294, 399, 400, 815
Ifymenopteroidea 265
ny()i)hila Midiolitzii 552
Hyperifiim japoiiifiim 706
r. jiramiiieiim 706
Hyperme^ethes 631
shnckerti 630
Tlyitnacwe 563
Page.
iryi)ii<)d(Midi-acea^ 567
IlyjuuMkMidi'oii s]»iiiiiierviniii. . 567
II jllttuim niossnidHiautuii . . . . 564
iimhiciilatimt 563
iryiHxlKrris radiata 709
Hypocct'lns 731
llyixMjeiia lateriflora 691
Hyi>oi>lei-y,2ia<'e;v 562
Tlypoprei-yiciiim Mnelleri .. .. 562
Hypoxis liynroiuetrica 703
InaMpiipalida 294
Indii^ofera aiisti'Mlis 705
Ipoiiiiea Horsfaliia' 601
Isodoii iteeuarius 723
Isoptery^iuni .-lust ro-piisilliiin 564
<an<lidiiiii 564
uordc-idh'HUK- 564
sii I ilati folium 564
rsosceliideroii 63!). 656
Tsotoma flmiatilis 708
Ithonidie r)30, 638
.Tiigata 296, 297
J iif/dtdc 296
Junoaceu' 689, 702, 710
Jiincajj:inaee:e 701
.Iniiciis 710
bufoniiis 702
falcatns 700. 702. 711
Foeljei 702
liomaloeanlis 702
laiuin'ocai'itiis 702
pallidiis 702
Itleheins 702
l)olyaiithemos 702
]>rismatocarinis 702
radiila 702
va.iiinatus 702
Kalli.m-ammitidje . . 773. 781. 782
]\ircliiieriella ohesa . . . . 533, 543
sultsoiitaria 533
Koderia plileoides 702
Ivniizea Miielleri .. 700. 707, 711
parvifolia 707
pediUKiilaris . . 690. 700, 707
Kylliiiiiia hnrifolid 702
intermedia 702
Lahiata- 691, 708
T,aceo])teris ]iol.viK»di<^ides , . 74
INDEX.
Page.
Ladenbergia 603
Lagena 386
Lageiiophora Billardieri . . . . 709
Lageniilina 386
Lagerheimia filiata v. f/rari-
iis 53?.
elliptica o33, 543
genevensis r. gracilis . . 533
Lasiocampa 307. 628
Lasiocampidsp 307
La sicca mpina 307, 311
Latliyrns odorat\is 601
Lauracefe 704
Legnepliora Moorii 601
Legiiminosa- .. 601. 690, 704. 710
Lembopliyllacea? 561
Lentibulariacene 708
Lrpidopilum auatralc .. 557, 562
Lepidoptera. 265. 269, 270, 271. 272.
273. 274. 275. 276. 279, 281. 286.
287, 288. 292, 295. 296. 297. 301.
311, 312, 313. 314, 315, 317, 401,
404, 406, 568. 569, 571. 572. 574,
635. 647. 648. 650, 651, 652. 654.
655. 656, 799
Leptoceridie 294
Leptorrhynchos sqiiamatus . . 709
Leptospermiun attennatum. . 700,
707
flavescens 415
r. obovatum 706
lanigerum 707
myrtifoliura 707
scopariiim 707
stellatiim • ' '^^l
triloculare 707
LeskeacejTp 563
Lestidae 418, 421, 422
Leto 303
staceyi 772
Leucobryacese 548
Leucobryum ballinense . . . . 548
brachyphyllnm 548
candidum 548
sanctum 545, 548
var 548
Scbimperi 549
strictifolium 548
vesiculosum 548
viride 548
Wattsii 548
Leucoloma clavinerve . . . . 547
Sieberianum 547
Page.
Leucoloma subiiitegrum , . , . 547
Leucoplianes australe 548
octoblepharoides .... 545. 548
Leucopogon biflorus . . . . 690. 707
Fraseri 690
Hoolveri 707
lanceolatus 707
Richei 247
virgatus 707
Liliacejp 601. 689. 692, 702
LimacodidiP 571
Limnanthemum crenatum . . 708
Liparetrus acutangulus . . . . 717
bituberculatus 719
distans 719
fulvoliirtus 717
incertus 718
intermedius 719, 721
iridipennis 719
majorinus 716
mixtus 718
obtusidens 720
parvidens 720
perkinsi 719. 721
plioenicopterus 718
quinquelobatus 720
tridentatus 720, 721
vicarius 718
Lissanthe strigosa 707
Lissapterus 716
howittanus 716
Lissotes 715
grammicus 715
howittanus 716
luteus 715
Litophyllum konincki . . 336, 595
Lituola nautiloidea 389
Lobelia dentata 708
peduuculata 708
Locustoidea 261
Loganiaceae 601
Lomaria alpina 701
capensis 701
discolor 701
Lomatia longifolia 703
Lo]>li<x-eratomyia 826,838
amiulata 837
cairusensis 837, 843
cylindrica 837,838
Loranthacese 703
Loranthus 696
pendulus 703
INDEX.
XIX.
Page.
J.otiis australis 247, 705
corniculatus 70o
LiK'aiiida^ 71,3
Liicilia sericata, 058. GG4. GGO. GG7
J.iiznla campesti-Ls 689,702
Lycaoii 729
ater 738
J^.v<'oiK)dite,s virturuo 89
Lyj;ieidie 581, 582
Lythracew 70G
J^ythnim liy.ssopifolia 70G
salkaria 70G
JMacrohymeiiium rufiiui . . . . 5GG
Macriunitriiiin aiirescens .. .. 553
Daeiuelii 553
dimori»lmm 553
liemitrichodes 553
microstomum 553
imgioiiifoliimi 553
imsilhim 553
Sayeri 553
Scottiie 553
Wliitelejj:j;ei 553
Macroiiciuatiiue 647, 656
Maerotteiiioyteris 85
.sp 85
wiauamatUe . , . . 85. 87, 91
.Mai;allaiies,. . 760. 761, 764, 765,
767, 768, 773. 785
insolcn.s 779
mcifricki 779
Maj-'iioliaceie 704
Malpijihiacea' 601
Malva rotiiiidifolia 706
Malvarefv 705
Maiisoiioides imiformis .. .. 838
Maiitispida' 633,638
MarattiacecO 64
Marcliaiitia polymurplia . . . . 701
Marc-liantites sp 89
.Mariaiithus prucumheiis. 691. 704
Mastoteruies 125, 128
MatoJiiiia^ 71
Mecoptera, 123. 124. 129. 130. 131.
133, 134, 267, 268, 269. 271, 272.
273, 274. 286, 288. 290, 301. 312.
316. 395. 397, 399, 400. 401. 403.
431, 572, 635. 639, 641, 645, 651.
654. 655. 656, 815.
.Me.uaj;iiatlia .. 125, 126. 127, 128
odouatif ormis . . 124, 126, 128
Page.
^Mejjairiiatliida' 128
.Megalodoii 386
:\lepil()ptera .. 124. 125, 126, 265,
268. 269. 274. 284, 291, 293, 297,
312, 316. 626. 635. 652, 655. 656.
819
Megaiieura 433. 434
monyi 433
^leganeuridu- 434
Megapsycliops, 769, 770, 771, 772
illidgei. 771. 781, 782, 785, 786
Megasecoptera .. 266
^Jeiotliccium IJrotlieri 565
teiieruiu 565
Wattsii 565
.Melaleuca 689
genistifolia 415
nodosa 415
]\Jeliclinis urceolatus 707
]\relodiiiiis aeiitifolins 600
Meiiispermacea' 600, 601
Mentha australis 708
laxiflora 708
satureioides 708
.Menura superba 701
Merismopedium 499
cyaneum 500. 501
duplex 500. 501
elegans r. const rictum .. 500,
501
Kuiiingil 499
puuctatuni 499, 500
/". oblongnni 500
r. vacuolatuni 500
-Merope tuber 290, 640
^Nleropidie 289, 316, 640
Mesocluete undulata 556
Alesojassus 370
Mesopanorpa 4:j5
liartungi 436
M<'.sof)an(tri)a 435
iridiHUHdttcK.'ii.'i 435
M cxKUxuiorp'uJac 436
Mesopanoi-podes 4;j."i
wiaiianiattensis 435
Mesopaiioi-podida> 436
]\Iesopli]er>ia 4;5.5
antino<lalis 417
Meteoriopsis reclinata 560
Meteoriuni lUiileyi 559
floribundum 559
xJc.
INDEX.
Page.
jMeteoriuiii Mi(iiieliaiiimi . . . . 559
Micromns tasmaiiia». OoS. 634. 638
.Mi( Toplasma parallelnm .. .. 595
Microi>tery.iii(bp. 275, 284, 286. 295.
297. 298. 299. 300, 301. 302. 304,
311. 313. 315. 318. 647, 649.
^Nlicroptei-y.iriiia 296.569
:Micr()i)teryx 298, 798. 572
arnncella 319
t]iniil)er.i;ella 319
^licroi'lia^iis 731, 732
niticollis 725
:^ri(■r(>seris lM>rstt'ri . . 090, 691, 709
:Mill€tia aiistralis . . 601, 603. 609
lue.i^aspeniia 603. 603
]Mliiieteoinyia .. .. 826. 830, 831
atra 830
atrijH^s 829, 830
doddi 831
piilcherrinia 830. 8:52
]Mimnlns luoschatiis 708
]Miiemo]iu-a 298
subpurimrolla 649
:NJiiesarclia'a 298
Miuaccie 555
^Iiiiodciulroii coiualuluin .. .. 567
Mniiiin si> 555
^lodiola multiHda 70(i
MoljxMnyia 838. 843
Iirii^stloii 838
]\i«>iiiiuia<'e;r 701
Moiiotoca scoparia 707
M(»iisfera aciimiiiala 601
Moiiireofia 513
lirlpviroiis 513, 514
MiK-idns alcniaiis 644. 828
^lulloriohryinn Wliitelejigei .. 557
Mnrcliisonia sp 33()
Miisca doiiu'stica :'.61
Muscida' 6U
Myi«»daria 644. 653. (556
Myriophylliiin ]>n»i)iii(|uiiin .. 707
s]> 509
MynneltM.n 117, 119, 122
uiiiseriatiis 116
MyiinelfM)iitidJt% 116, 117, 119. 120,
121. 291, 403. 631. 638, 783, 815
Myrtaco;e. 690. 691, 706. 710. 817
]Mynriinii nifesceiis 557
MyxophycojH 498, 499
Aly/<»inyia 827
Myzorliyiichus 827, 843
Page.
X:eii)al('lia 885
Xaiiiiocliorista dipteroides . . 289
Xamioclioristidai .. 131, 289. 312,
316, 640. 656
Xcckcra 561
trichophoroldes 560
Xeckeraceit' 557
Xeckeropsis 560
Nematodes 731. 733
Nematodimis 731
armipennis 731
NeniopteridiF 638
Xeocalamites Carrerei, 64, 67. 70.
78. 80
lut^reiisis ,. 6 4, 67. 70, 78. 80
Xeosalpiiiiius 743
brevis 745
politns 745
XcosfjiKinioniijia 829
Jirrinli 828
Xonroptera. 116. 265, 268. 269. 750.
784, 785, 787
Xeiiropteridinm an st rale . . 85
Xeuropteroidea. 265. 266, 267. 268.
269
Xilssoiiia sp 89
X'ostocact'ji' 503
X<.th(M]jrysa 806
Xdtodoutina 307
AOtopsycliops . . . 759
Xotosalpiii,mis 744
]>ruiiiiOTi.s 742
Xovapus ])arviis 722
simplex 723
\y<tai,'ine;e 601
Xyini»iiida« .... 120. 121. 638, 783
Nyssorliyiicluis 827, 828
Ochlerotatus notuscriptiis .. 834
Octoblephariim all)idum .. .. 548
Odoiiata, 133, 265. 282, 417, 418.
422, 423, 425, 432. 630
Oeretis sp 294 313
(Edicladiacea- 557
(Eiiotheia biennis 247
(Enotheraceie 707
()leandri<num leiitricnliforme, 85.
87
sp 85
Olearia arjr()]»hylla .. .. 701, 709
axillaris 247
INDEX,
Page.
Olearia clirysophylJa 709
mepilopliylla 709
stelliilata G91, 709
()lit;(»t(>ma T. .. 128
sp 1:37
Ompliacomeiia acerba 703
Omphyma.. 336
Oiiiscif,'aster 278
Oocystaeete 531
Oocystis 498. o31, 532, 534
laciistris 532
r. iiataiis 532, 543
r. paliidensis 534
532
534
542
666,
uodvilosa r. crassa . . . .
ovalis r. cyliiidratea . . . .
rotiila 518, 531^
Ophyra iiij^ra.. 658, 660, 665,
667
Orlnilina 386
Orchidarea^ .. 601. 689, 691, 703
Oreouiyrrhis aiidicola 707
Orioua strata 383
(Jrites laiicifolia 601, 703
Ortliis resiipiiiala 45 1
Orthophlolda J 24, 129
Orllioplera . . .. 261.
. 262, 282. 431
Orthorrhyiicliiiim
cyinbifoli-
oides
560
OrthotricluK-ea- . . . .
553
Oscillatoria
502
■ liexa
. . . 501. 540
. . . . 502, 540
simplicis.sima r.
cons trie
ta
, . . . 502, 540
r. irrauiilala
. . . . 502, 540
Oscillate )ria< vie . . . .
501
Osmuiidarea:'
64, 71
()smylida\ 638. 755,
756. 783, 784
Otozamit^s
50, 80, 92, 93
Feistmaiiteli, 72,
75, 76, 79, 81.-
92, 93
Maiidelslchi. 72.
75, 76. 79, 81
obtnsus . . . . 72
, 75. 76, 79, 81
(liieeiislaiidi. . ..
.. 72, 79, 8J
Ottelia prteterita ..
85
Oxaiidacca- 705
Oxalis coruii-ulahi 235, 705
Oxylobium alpestre 696
ellipticniu r. alpiiniiu . . 704
prociiiiiboiiS(( V) .. .. 690, 704
Tiiltenece 704
Page.
Pala-odictyoptera. . 265, 266, 630,
631
PalceoluMinplei-a. 586. 587, 589, 592
Paheontiiia 569, 570
oolitica 569, 571
Pakeoiitiiiidic 570,571,592
Palissya austialis 90
Panax sambucifolius 707
Paiiicimi efl"\isiiiii 702
Paiiorpa 395, 396, 632
coiifusa 627
Paiiorpata^ . . . . 265, 266, 267, 268
Paiiorpida- 124, 640
Paiiorpoidea . . 265, 266, 267, 268,
269
Papilioiiiiia 307, 308, 311
Papillaria amblyacis 559
BaiUi/i 559
flexieaiilis 559
keriuadeceusis 559
iiitidiuscula 559
Pappoplutnim (•(aumiiiio .. .. 702
J'ai-adimstaiiia .. .. 576, 583, 585
atiiiiis 582, 585, 586
Parallelopora dartiii.utouciisis 381
I'areleaiia 262
I'assiflora suberosa 601
Passiflorea* 601
Pecopteris ( ? ) tcnnifolia .. ., 85
Pediastnim 537, 538, 539
aiijjciilosiim /•. areiiosiim .. 539
biradiatiim 539
/•. Piaunii 539, 543
JSoryaiiimi . . . . 538. 539, 543
r. austrak' 539, 543
r. Hayiiaidii . , . . 539, 543
lirainrii 539
clathratiuH 537
duodena rium 538
ilu]>\ex V. (kit hra turn. . .. 538
r. ivticulatinn 538
viKqthjn 537
Udi/nahlii , . . , 53!>
or at urn 538
rotula 539
Schrbterii 538
simplex . . 536, 537, 538, 543
/. clathrata 537
V. clathratHin 536
V. nutlanii 536, 537
^iunnii 538
XXll.
INDfeX.
Page.
IVdiastriim tetras r. Crux
Midicf li 540, 543
tricorinitiim 539
Pelarfroiiium 197, 198
australe 247, 705
Peukillium eladosporioides, 145,
146, 149, 155, 167, 168, 369, 170
172
italkiim 874
sp 148 158
reiiiiisetiim loiigistyliim . . . . 234
Peiitameriis kiiiirhtii 336
sieberi 336
sp 383
VeiitatomuUe . . 577, 582, 591, 592
Terga 668. 671
dorsalis 668
macleaii 671
rericlystus 121
Peridiuiese 509
Perissectis australasite . . . . 302
Peri8Soi)hlel)ia 422, 423, 425
miiltiseriata 422, 424
Peiiaria. 124, 125, 265, 282. 431.
755
Perlida- 124
J'ermocliorisfa .. .. 123. 129, 267
mitclielli 130
Per.sooiiia ehama^peiK-e . . . . 703
Petaliira 423, 434, 435
jlLijaiitea 424. 612
Petripa volnbilis .. .... .. 601
Pftrothemis 423
siiijiiilaris 425
Pliaeotns . . . . 498. 509. 523. 524
australis 524, 542
crassiis 523, 542
.U'laber 523. 524. 542
IMia'iiocerus 739, 740. 741
clavicoriiis 740
subclavatus 739, 740
Phii^toiiides 485
Pbasco^'ale fiavipes 361
I Mia scolomys mitclielli 696
IMiaseoliis vulgaris .. .. 611. 861
riicbalium scpiaimilosum r. al-
piinim 705
Phillipsastriea. . .. 322, 377, 379
giiras 379
graiidis 336, 379, 384
heimalii 383
verneuilii . . 323. 333. 337, 379
Page.
Philippsia .... 438, 439. 443. 465
breviceps 449,489,493
colliiisi .. 442, 443, 444. 446,
447, 489, 490, 494
coiiiiollii 472, 493
roiilteri 447, 473, 489
derbiensis 450, 455
duhku 438, 439, 440, 456, 458,
459. 460, 461, 466, 470, 481,
482, 490, 491
duiigogeiisis 474, 490
eichwaldi . . 438, 440, 451, 455,
472, 478
ckhiraUU 440, 477
elongata.. 454, 456, 458. 462.
489, 491, 492
gemmiilifera 446
graiidis. 439. 441. 443, 453, 454.
463, 464, 465, 489, 492
(jrandi!^, 441, 451, 452, 454, 472,
473
iusigiiis 446
leei 456
minor 456
morgaiieiisis, 471, 479, 491, 492
proxima 451, 489
robiista. 451, 453, 465, 489. 490
rockhamptonensis . . 459, 466.
479, 491, 492
roiulieleiisis 473, 490
seminifera 438
.'■icminifera 447
spp. iml. . . 439, 441, 446, 473
staiivelleiisis . . 458. 460. 461,
463, 470, 473, 474, 479, 482,
490, 493.
stroiuleiisis .... 453, 489, 493
superba 454, 491, 494
trinicatula 446, 458
waterliousei. . .. 455. 491, 494
woodwardi. 437. 439, 441, 442,
443, 465, 468, 470, 472, 479,
480. 488, 490, 491, 492, 493,
494.
iroodiranJi 459
Pbilomastix i^/(/Z;cr 671
macleaii 671
Philoiiotis laxissima 556
pseiidomoUis 556
I'lilebopteris aletliopteroides. . 71.
74, 75, 78, 80
Phoeiiicopsis 94
INDfiX.
Page.
J^li(jeiii<-oi>sis elonjxatus. 72. 73. 79,
81
J'li<»ma 8GU
citrk-arpa, 808. 809. 870. 874,
878. 879, 880. 881
lavaiuliihe 878
solaiii 875
J'liomopsis 875
I'hormidium jrraiide .... 502, 540
iiumarium 503, 541
I 'lira jrmites communis 702
l'hryy:aiieidce 393
I'liry^'aiioidea 205, 208
J'hinochronKjphifvvae 499
rhyllaiitliiis tliymoides .. .. 705
J'liyllopteris Feistmauteli, 40, 72.
70, 79, 80, 91
Phyllosticta 809
citricarpa 809
I'hyllotheca 50, 85
aiistrali.s, 04, 07. 70, 78, 80, 85,
87
coiiciniia 85
Hookeri 85
Pkris hieracioides 709
Pieliis 303, 305
byaliiiatus 303
I'imelea coloraus 700
turviflora 700
jLTlauca 700
ligustriiia 091, 700
/•. hypericiiia 700
linifolia 7O0
paiK'iflora 700
sp 090
I
iiiaceit' 701
iiiiiatella iiitralimbata .. ..501
inns sp 803
iper iiijj:riim . . . . 001. 007. 009
iperaceie 00 1
isicola 140
ittosporacfje 091. 704
Iaj;iantliiis pulchellns . . . . 705
r. tomeiitosus 700
lau'iutliociete 504
laniponnia. 133. 209. 271. 274, 284.
287, 290, 301. 312, 313, 310, 401,
400, 431, 432. 572, 030, 034, 035.
037. 048. 051. 652. 055. 656, 657.
753. 754. 757. 759. 781. 784. 787.
813. 815, 819.
Plaiitairinace.'ie 708
riantajxo laiKOolata 713
varia 708
Page.
l^lasmupura 330, 379
IMatylobiiim formosiim .. .. 705
I 'kHtoptera . . .. 205. 278, 399, 574
Plectrotarsiis 294, 047
^'raveiihorsti 040
PJeospora hesiKiridearum . . .. 809
Pleiiridiiim .sp 545
Pleurocarpi 550
J'leiirococraeeie 527
I'leiirosorus riitifolius 701
Pleiirotus sp 4l5
J'lnsia 305, 309
vertk-illata 305
Poa ciespitosa . . . . 089, 091, 702
Podolei)is caiiesceiis 708
loiijripedata 091
r. robiista 709
I'odozamites ]>arklyi 89
. elliptk-us 89
Kklstoiii 89
lam-eolatus 85
loiiju'ifoliiis 85.89
spathiilatiis 85
sp 85
Polleiiia stygia . . 058. 002, 060, 007
i'olyedrium 534
aciimiiiatiim 535
octaedricuin 535
r. spiiiusum 535
ix Htat/oninn 535
tetraediicuin 535
tcfnifjonNm 530
trigomim 530
IV»I.v.ij:alare;e 091,705
I Vdy;j:oiia(eie 001, 703
l*oly;Lroiiiim minus 703
prostratum 703
J'olypodiacwr 091, 701
J*olypodium diversifolium . . 701
l)inirtattnH 701
scanilcH^ 701
Polyporacea' 885
Polystiflium atideatum .. 091, 701
I'<»lyst(e<lu'ti«l;i>. 038
IN>lyst<jechotos 784
IVdytricliacwe 550
J*olytri(lium jiiniperinnm. . .. 550
I'omaderris 097
apetala 705
elliptica 705
j)liyli(if<)1ia 705
juMinifolia 705
ratemosa 705
XXIV
INDEX.
c*AGE.
I'omax viinbfllatJi 700, 708
PorantlKM-a micropliylla . . . . 705
IN.rina 302
umi>ra<ulata 304
l\»si<l<>iiia 247
rotamoju'etoii iiataiis 701
natan.s 701
perfoliatus 701
tricariiiatiis 701
lV)tam<>^etoiuK-ea' 701
J'othos aureus 601
I'ottiaceje ^•^~
JV)\vellia australis ^62
I'rasophyllum fuscum 703
I'rimiilacetf 708
Pioetidjv -139, 443. 482
Proetus ^^'^
mi.ssourieiisis 450
l*rohemerobiidci\ 773. 781, 782, T83
rroroceiitraeea^ 509
rrosbole 588, 589, 591
hirsuta 587, 588
rrostaiitliera cuiieata . . 691, 708
lasiaiithos 708
Proteaceie .... 091. 703, 862, 867
ProtiK()<<acea* • . 528
ProtocoicoideiJe 498
Prutodonata .. 4 17, 425. 431. 432.
433, 434
i'rotoheiuiptei'a . . .. 587, 588. 592
I'roti»mec«.i»t('ra. 209. 284. 288, 316,
588. 631, 634. 035, 641. 645, 651.
054. 650. 657.
Protoiieuriii;e 421
I»rotui>le(tiiiii 116
I'rotoplectixdi 122
.stria telhim 121. 122
veiuistum 122
Protoi>sy<lu)l»si.s. . .. 781. 782. 785
veiiosji 785
I'rotusialis.. .. 820. 821. 822. 82:;
Protosinlion 530
Prototheora. 301. 302, 648, 650, 652
petroseuia, 301, 648, 650. 651.
057
Prototheoridie. 295. 297. 301. 302.
312. 313, 315. 319. 647. 650. 050
PntnrUn rnhjarifs 708
Psaumiospluera 388, 390
fusca 388
iiemiiigheiisis . . 331. 387, 388.
390, 391
Page.
JNc'udiKteiiis fathieiisis, 72. 75. 76.
79, 81
Pseud(>i»lia'a sp 495
Pseudosknsea . . . 820
l»asalis 829
cairnsensis 829
mnltiitlex 829
Psocoptera 132, 133, 131
Psoralea adsceiideiis 705
Psydiida' 305, 048
Psyclioda sp 043
J»sydiodida\ 573, 042. 646, 651. 650
Psychopsella .. .. 769, 771, 780
.i^allardi 771. 780, 786
J'sychopsidie ., 281, 282, 628, 02'.).
030. 637. 638. 750, 753, 754, 75f.,
757. 762, 764. 773, 781, 782, 783.
787, 813, 815, 816, 817, 819.
J'syfh<»])siiia' 784
I'sychopsiiii, 759. 760, 761. 762, 771,
785
I'.sychopsis. 755, 756, 757. 759, 760.
764. 765, 7G6, 767, 768, 769, 770.
771. 773. 774, 778. 782, 783, 784.
785, 797, 799, 802, 804. 806, 815
l>irinaiia 758, 760, 781
fo< lirnfjii 779
(■(i-Ii\"a;:;us. . 767. 770. 774, 775.
778, 803, 804, 805, 809
ele.uaii.s .. 629, 750, 751, 752.
753. 754. 755, 756, 758, 760,
761, 764, 766, 767, 768, 770,
774, 775, 777. 778, 787, 788.
789, 790, 793, 794, 797, 798,
800. 802. 803. 808. 810, 812,
814, 816, 818.
irrarilis. 766, 770, 775, 776, 786
illi<lfjri.. .. 765, 769, 770, 771
iiis«»leiis , . 755, 756, 760, 761.
767. 770. 774. 775, 779, 781.
784. 816. 817.
iiiarslialli 758. 781
iiieyrkki.. 751. 700. 701, 768,
770. 775, 779, 784
mimica. 755. 756. 759. 761, 765,
708, 770, 774, 775, 776, 777,
785. 787.
tniinUa 775, 777
)i(inmuii . . 760. 707, 777, 787
zel.ra 759
I'UMidiiim a(|uiliiiuin 701
Ptirin aquillmi 701
INDEX. XXV
Page. p^^jp
J'lcrohryidinm 544 JJaphidia 313,630
192
IMer..l.ryoi.sis .^3!) Kapliidiuhp 392,310,650
fihgora . 558 Kajiliidioidea . . :.^05. 208. 20'.), 029.
rtcrof/oniclJa M attsii 50o g^g g.J „.
I'trrof/oniMm flarcsccm .. ..501 jjestio australis '..''"' 702
rteromoiias 522 ,, ^.
(//./fa 522 :f-^f'''":i<-^^^ 691. 702
V. an-sfralis 522 J^li'i^<'l»ila<ea- 502
aii,:,'iil<»sa 522 JJ'if«<<'pil"ni coiivolutareiiiu .. 502
r. aiistralis .. .. 522. 542 Kliaimiactw 705
r. scutiformis .. .. 522.542 Ifliaphidosteuriuni acirulum. . 505
r. vexilliformis . . 522. 542 ovale ,505
lM(>r<»i»liylUiiii almorme. 72. 79. 80 J'seiKlo-lKnnoiDalliim .. .. 505
coutijiinim . . 72. 75. 70. 79, SO saitn.xylopliihim 500
miiltiliiieatimi. 05. 09, 70. 79, Ivhe^'iiiatcMlontaceie 560
81 Jlliijn'docerida' 7;mj
Nathorsti . . 72. 75. 70. 79. 80 lvMii]>idoi)sis -inkiroidos r. lata 85
^1» 85. 87 KMiii)<»j;(>inim allnim .. .. 001, 0O8
Pfen.stylis curtaC') 703 KliizohiuDi sp.. 148, 149. 150. 153,
<>''tnsa (V) 703 154. 155. 158. 159. 161. 107, 109
i'"f'^ ''•'•'^ iniizo^roiiiarea' 556
J'teryp.i.honis 670 |;iiiz,,i;(,niuin lnvvif(,lium. . '.'. 556
ijialis 671
>ifas(iatns 070
(ioheebii 556
pairainaltoiiso 556
Jnil(M»liylluin JWillianisuina) si»ii,iforim. 55(
KMiizonioptoris l':tlu>rid-ei .. 89
Ivliizopprtlia dojiiinica 361
KMiodohryiiiii olix a<riim ., .. 55. »
snltciispalniu 555
i;ii<»i>alo<(M-a 296, 307. 309
Uhyac<>j>lii!a .. 269. 271. 293. 64 7
dorsalis -nro. 645
J{hya<<n>liili<la' 645.646
|ie<-1('n .. 72. 74. 75. 70. 79. So
Pty<li<Mk>ia austialii'usis. . .. 136
rtythoiniiiacoa' 557
i'ultciura fasciculata 705
MiK'lleri 705
l)ro< uiiiboiis 705
I'ulviiinliiia 390
ln'iisoni . . . . 331. 389. 390. 391
bnjM-kijana 389 ,,, , , ■ ,, ...
^I^,p,„^ l-v.;,, I'Jiynchosti— iclla ••(•nv«.hil]f()lia 567
Pvaiosoma nitifaciVs '.". 658." 660. 'Jl».vn<hosl(-iniii nano-pcima-
663. 664, 666. 66^
varipes. 658. 660. 664. 666, 66,
t'liii 567
tonuifoliimi 567
rvnh.)(<.rid;r.. .. .. .. 581. 582 l''l».vplii«ln' .... 041. 042, 045. 650
I'Vthida- 742 IJI'M/Jnis 027. 642, 052
bnnis 627, 642, 657
. K'iclica (inniiii .. .. 691, 700. 707
Quisqualis mdi<a ooi ,.,,^., h,.;„.(t.ata .. 236
tnrltiiiata 236
KaiiniicniaccH' 703 Kosacca' 704
Kainniciilus a<iuatilis( V) .... ;o3 K'oltiaica' 601.691, 708
liirtiis 704 Kulms lacinialus 235
lappaceus 703 j)arvif()lins 704
parviflorus 701 iJiimcx atotosella 703
ph'fxiii.s 701 Prownii 703
rivularis 704 llutaceie 690
XXVI.
IND6X.
Page.
Sabatiiica 208.300,318
iiK'oiiirruellti .. 299, 300. 318.
319, G49
Sarcammiim 388
Saccammiiiiiice 387
Saccharoinyoes cerevesiie. . .. 412
Zopfii 411
JSageiiopteris 94
J'hillipsi 68, 74, 75
rhoifolia, 04. 68, 70, 72, 74. 75.
76, 78
Salvia verbenacea 708
SambiKus (laiidicliaiuliaiia . . 708
Saiiidopliylliim davidis . . . . 595
Santa lace;e 703
Saiitalum obtiisifolium .. .. 415
Sapindaeeie. . .. 703. 850, 862. 867
Saturniidce 572
Saturniiua 307, 311
Scievola erassifolia, 247, 248, 249.
250. 253, 254, 255, 257, 258
microcarpa 248
i'lumieri 248
spiuesceiis 248
Scarabieidie 716
S<h((lonorii>< llookvriunu . . . . 702
Schizoneiira 264
af ricaiia 68
(•/. afrkaiia . . 64. 70, 78, 80
australis 85
j^oudwaneiisis 68
sp.a .. .. 71, 73." 75, 76, 78
sp 71, 73
Schizot)h!/r«ar 499
Schlotlieimia Haileyi 553
Seirpiis mrtUuf/UK'Ha 702
teriiuus 702
iiiuiidatus 702
laciistris 702
p<»lystacbyu8 702
riparina 702
stelbitus 702
Srlerantbus bifionis .... 689. 703
Scruphidariaeeie . . .... 690, 708
Scutellaria hnmilis 708
>'<(utonuila atriprs 829
notoffcriijtfi 834
Scytiiioptera kokeiii 587
S<ytoiiemaceie 507
Scl( naiitrinn obrsum 533
Selliera radieaiis 248
Seinatopliyllacea? 565
Page.
Sematophyllum erythropodiuiii 566
Seiieoio ai(&truU!i 709
dryadeiis 799
lautiis 247
pectiiiatns 691. 709
tamoides 601, 603
Serieo«tomatida.^ 294
Sialidie .. 124, 291, 316, 636, 656.
819. 820
Sialis . . . . 291, 636. 820. 821, 822
Sialoidea 269, 636, 819
Hk'bvra BUlurdicri 707
Sileiie jrallica 703
Siricidie 400
Sisyrida? 638
Sl^usea i>seiidomediofa.sciata . . 838
Smilax anstralis 601, 608
Solaiiacete 601, 708
Solaiiiim iiijj;rimi 708
opaeum 708
simile 708
Wendlaiidii 601. 603. 608, 609
Siiei'iiriilaria rubra 703
Spermopliorella. . .. 638, 639, 656
disseminata 639
Splueropsidio 875
Spluerozya laxa 505
Sphenolepis rhtetiea 69
Sphenopluu'us striatus . . . . 361
Spheuupteris 69, 85, 86
ampla 89, 90
laeiuiosa 65, 78, 80
sp 85,89
superba 65, 72, 79
S]>inifex hirsutus 247
Si>irifer striatus 454
Si»iroi:yra 510
Fullebornii 513
kuusanioensi.s 510, 511
J Jsmorensis . . 510
mirabilis 510, 511, 512
ne^'lecta 512,513.543
r. amylaeea . , . . 513. 543
nitida 513
spp 510. 511
Spreeiana 510. 511
Si>irottenia acuta 527
Sphnlinacea' 553
Spondylomorum quarternari-
um 525, 542
r. rostratum 526, 542
Spongophyllum bipartitum . . 377
INDEX.
xxvn.
Page.
Spon.ijophy 11 11111 Lciuanteum . . ."iO.'j
sp :]?>6. p.TT. nrs. :!st
Star-hyoi>itys 66
.'iiinularioidos. ().'). ()9. TO. 79. SI
SiiuiiioiKlsi ().■), 79. SI
sp 69. 70
Rtiukhousia liiiariifolia . . . . 70")
nionof/iittd 70"i
viiiiiiioa 70."')
StacklKnisiacoa^ 7o")
Stoijouiyia 829
iitni 830
f)ini(t(i]atrr<ili.9 .. .. 829. 830
Stcllaria piiiii^eiis 689. 703
Steiiohiolla 43J. 638
Stonoi»teris eloiiicata. 6."j. 69. 70. 72.
74. 76. 79. 89
si» 89
Stoiiosialis 821. 823
aiistralieiisis 824
Stei)liaiiia lieniaiidiiefolia. 601. ()07
Steniilia acerifolia 19.')
(livei'sifolia 19.5. 706
Sterol! liacoa- 706
Stereodon mo>;.>^inaiiiiiaiiiis .. 061
.siilu-lirysoiraster .564
StertHKloiitea' .563
Stij;iiiapliylkui .iati(»i>ha»f(tliiim 601
Stipa scahra 702
Stomat()|K>(la 278
Stratioiiiyia fliamivleo 662
Stroma topoia 380
coiiceiitrka 382
Stromatoi»orella )>eiisoiii. 336. 380.
381. 381
loombereii.sis. 336. 380. 381, 384
Stroi»homona of. ruirata . . . . 336
Stiiartina Mueller! 709
Stylidiaoesp 709
Stylidium sramiiiifolluiii. . .. 709
Stypaiulra j^laiica 702
Styphelia triflora 707
tiibiflora 236
Swaiiisona teidirotriclia . . . . 70,5
Sy<-oi»teri(l;e 134
Syeopteroii, 124. 129. 130. 132. 133.
134
i<iimm(tnc(i 124, 129
symmetrkiim 129, 130
Syiioarpia lanrifolia.. .. 415. 803
Syiiechoooeous jjrandis . . . . 499
major 499
Page
Synemoii 308. 309. 310, 311
hesperoides 308
Synle.'^te.^ 431
weyersi 420
Syiiiira sraiinlosa 508. 542
Syriiiiropora 332
aiiloi»oi<)ides 331. 595
porteri 331
Syromastes sj».. 576. 577. 578. 580
Syri>hid;e 644. 656
Syrrhoitodoii aiiKPinis 550
<airneiisi.'< 551
croceus 551
fasHculatiis . . ._, 551
fimhriatiilns 551
fiml»riatiis 551
hiimilis ;552
^riielleri 551
Xova' Vales;ia> 551
uiidulatiiliis 551
iiiidiilatns 552
Tabaiiida^ 643, 661
'I'ahaiiiis sp 644
'r:eiiior-hori.«<ta 267
palUda 289
'IVeiiiopteris. 56. 66. 67. 69. 73. 74.
78. 264
Carruthorsi. 65. 69. 70. 72. 76.
79
orassiiiervis. 65. 69. 70. 71, 72.
76, 79
Daintreei. 58. 84, 85. 86, 89, 90,
264
r. major 89, 90
Diiiistaiii 65. 79, 80
fliKtiians 91
leiitrk-iiliforme. 65, 72, 79, 87
MeClellandi 85
sp 56, 85, 260
spatulata. 58. 72, 76. 79, 80,
84. 85, 86, 90, 94, 264
r. ereiiata 89
r. major . . . . 72, 76, 79, 80
Teni.soii-Woodsi. 65, 69, 70, 72,
76, 79. 86
wiaiiamattie . . 65. 79, 80, 87
Tasmo.salpiiigus 743
promisciuis 74 5
qiiadrispilotiis 744
Taxites 56. 75, 90
planus. 72, 75. 70. 79, 81, 35,
86
XXVlll.
INDEX.
Page
Taxites sp 90
Taxithelium kerianiim . . . . 564
Xovjp Valesi.ne 564
papillatnm 564
\Vatt<;ii 565
'IVcoma Hillii 60()
Jasmiiioides 600, 6();!
'l\'iirhi'*Mlini<lje 668
'JVidirosia liraiidifiora 2?)'i
'IVssHlla 508
TrxH(Uu 508
'IVssellaria 508, 509
volvociiia 508, 54:2
Tetiaedron 498. 534
a<umiiiatiini spino.siim . . 5.35
acutiim r. re<tilineare . . 535
caiulatum 535
r. ausrrale 534. 54:'.
coiikiim 535, 543
Lira nil l«»siim 534. 543
hastatiim r. elegaiis . . . . 535
uiiiiimiim /. rertilint^are. 536.
543
obrusnm 536, 543
Muadratiiia f 534.543
re^ulare 535
r. octaedricniii • • . • 535, 543
trij?omim 536, 543
'IVtrastniiii eleiians r. deiita-
tiiiu 540
'LVtratheca ericifolia .... 697. 705
r. thymifolia 697
Texmlariida^ 388
Thelymirra veiio.sa .. .. 693. 7o:>
Themeda Forskalii 70:3
'J'heobaldia 830
atripes 830
Thlnnfeldia 68, 74
acuta 64, 78, 80
Feistmaiiteli. 56. 64. 68, 70, 12.
75, 78, 85, 87, 91
lancifolia. . 64, 68, 70, 72, 75,
78. 85. 87, 91
^rcC'oyi 89
media 91
narraltoeiiensis 85
(Mlontopteroides . . 56, 58. 64.
68, 70, 12, 75. 78, 85, 86, 87,
89, 91.
sp 85. 89. 585
Thiiinfeldieje 64, 7:2
Page.
'riuiidiiim lillipiitannm 563
plumulosiforme 563
rameiitosiim 563
sparsiim , . . . 563
s\d>ere<tiim 563
Til nil he rui a .i;randitl<>r;i .. .. 600
lanrifolia 600
'riiymeheacea' .. .. 690, 691. 706
Thyrididjc 308
Th i/rifliiim hiiniilc 552
riiysanotns tuberosns 702
'rie^heiiKtpanax sainlmcifolins 707
Tilla'a verticillaris 704
'I'olypothiix lanata 507
roxorliyncliires inoinatns.. ..82S
spe<-i<>sns 828. 843
'l'ia<liel()si»eiimini jasminoides 600
'I'rafhylonia planit'olinm . . . . 557
rcciirriilinn 560
'ri-a<liyinene P.illardieii . . . . 707
Jtu mills 707
Irarliypns Xovie Caledonia^.. 560
'l'remaiidi'a<e;e 705
rieinatodon P>aileyi 545
lonii-escens 545
Tremelia 885
'rienielliiue 885
Tiiassolestes 418
epiophlebioides MS. 419. 420
Triassolestiiue 4 is
'I'libonema 529
Tric-hinuim corymbosniii r.
parviflora . . 610
'I riclioma 638
Trk-homatidie. . 431. 6:J8. 783, 784
'Iriehoptera, 267. 268. 269. 270, 271.
272, 273. 274. 284, 286. 292, 293.
295, 297, 302, 312. 313, 317. 399.
400 401. 404, 406. 572. 645, 646,
648, 651, 654, 655, 656. 657, 755
Trichosteleum elegaiitnlnm . . 566
. liamatiim 566
r. semimammillosiim . . 566
kerUumin 564
Tricoryne elatior 702
'I'rifoUnm arveiise 704
'rriillocbiii procei'a 701
'I'rigonoplenrns 731. 733
Trilobita 439
Tristaiiia neriifolia 415
rropidorhynchus corniculatu.s 695
'rry)»lasma, 321, 332, 334, 335, 336
INDEX.
XXIX.
Ti'yi)lasma conjjreixationis
(lelicatuhi
lonsdalei
r. scalarifonnis
sp 335
Page.
. 331
. 3.'! 5
ijol
. 321
vt^rmifoi'uus , .
Tnni<a prolitVra .
Typha aiijj^ustifolia
Typhaceje
331,
Ihiu.s liilli
riiihellitVra' 091,
I iiio 38
Duiistani
eyreiisi.s 88.
iljsvicieiisis
Stirliii.sj:!
wiaiiamatreiisis
( uioiu'lbi
howralensis
( arnei
I. i'aii(>t;eiiia 826.
albescens
antenna lis
• airnsensis
liilli 841,
Id'opria
tibialis 839.
LTrnella
terrestris .*)29,
/•. al»norniis . . . . .■>3o.
I 'rti<a incisa
Li'ticaceje
rstilaj;(»
I'lricnlarla dicliotonia /•. \ini-
iiora
Ura CaMiiioi^'ufiis
703
701
701
130
707
, 88
88
91
88
9(1
88
38
88
88
842
839
840
839
843
840
84;i
529
530
531
70;i
703
203
Vaji:innlina .
Valvnlina ..
biill<)i(le>-
388.
331. 388. 389,
oldons'ji . . 331, 389,
plica ta. . .. 331, 388
Wlleia paiadoxa .. ..
\'erl>ascum blattaria . .
Veibenace.'B
N'ei'onica calycina . . . .
Dei'wentia
390,
390
708
525
388
389
390.
391
391
391
708
708
601
708
708
Page.
Veronica ?:racilis 708
perfoliata 690, 708
N'esicnlaria rivalis 505
Viola betonic.-efolia . . 090, 697, 706
lied era <ea 697, 7()6
\'iolacea' 690, 700
Vitis liyitojjjlauca Goo
opjica 600
sterculifolia, 000. 002, 600, 009
Vlttadinia anstralis 709
Volvocaceie 498, 509, 514
N'ovlox 498, 526
aurens r. hemispluericus, 526,
542
Carteri 527, 542
tertins 527
Volvullna 526
Steinii 526, 527, 512
Walilenbergia .i?ra<ilis. . 700, 708
Weisia viridula 552
\V«Mnzia, 764. 765, 706, 707, 708.
773, 785
cocJiraga 778
Westrin^ia eremicola 708
WillUnn.sonia pcdrn, 72, 74. 79, 80
.S7; 65, 69. 79
Wistaria chinensis. 601, 602. 605.
609
Wormia P.urbidgei 249
Wiinnhra dioica 702
Xantliodiscns Lanterl)aclii, 509, 542
Xantlioleon 119, 121
lielnisi .... 118. 119, 120, 121
Xantliorrluea sp 702
Xerotes tiliformis 702
trlauca 702
lonjiifolia 702
ninlliflora 702
Zaphrentis sp., 331, 335, 375, 376.
383, 384
Zornia dipliylla 705
Z}loi)hhhihu .. 759. 760, 761, 762
Zi/f/ophlchia 784
Zm/oijhlchiiiH .. 759 700, 773, 785
rcrndK.iinuR .. 759. 777, 785
zehra 760
Zygoptera 418
Zyj;ospira sj* 331
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Megapsychops iUidgei (Froggatt).
P.L.S.N.S W. 1918.
PL. LXXVIU
4-6. MajapsycJtops illidgei (Froggatt) . 8. Psijchopsis <jracilis, ii. sp.
9. Psychopsella gallardi, n.g. et sp.
P.L.S.N.S.W. 1918.
PL. LXXVMI.
10. Pstjchopsis (/racilis, ii. sp. 11. PsychopticJla rjallardi, n.g. et sp
•L.S.N.S.W. 1818.
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■ L.S N.S.W. 1918.
1. Anophdesbarbirostris v.'hancro/ti (Giles). 2. Toxorhyachites speciosus (Skuse).
3. Arinigeres breinJi Taylor.
P.L^.N.S.W. 19I9-
PL. LXXXI.
4. Grabhamia theohaldnajloY. 5-7. Cidicada icilsoni, s'p. n.
t'.L.S.N.SW. 191
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PL. LXXXIII.
12
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Phoma ciiricarpa McAlp
P.L.S.N.S.W. 19!!
PL. LXXXVII
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A<jaricus campt'sfi
Issued Srd July, 1918.
»m
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PROCEEDINGS, 1918, PART 1.
CONTENTS.
PAGES
Presidential Address delivered at the Forty-third Annual Meet-
ing, M^rch 27th, 1918, by Professor H. G. Chapman, M,D.,
B.S., President 1-30
The Geology of the Lower Mesozoic Rocks of Queensland, with
special Reference to their Distribution and Fossil Flora, and
their Correlation with the Lower Mesozoic Rocks of other
parts of Australia. By A. B. Walkom, D.Sc, Assistant
Lecturer in Geology, University of Queensland. (Plates
i.-ii., and six Text-figures) ... ... ... 37-115
Studies in Au8ira.\ia,n Neuroplera. No. 5. The Structure of the
Cubitus in the| Wings of the Myrmeleontidce. By R. J.
TiLLYARD,, M. A., D.Sc, F.L.S., F.E.S., Linnean Macleay
Fellow of the Society in Zoology. (With three Text-figures) 116-122
On the Affinities of two interesting Fossil Insects from the Upper
Carboniferous of Comm entry, France. By R. J. Till yard,
M.A., D.Sc, F.L.S., F.E.S., Linnean Macleay Fellow of the
Society in Zoology. (With three Text-figures) 123-134
Tha Origin of Yolk in the Ova of an Findoparasitic Copepod.
By H. Leighton Kesteven, D.Sc, M.B., Ch.M. (Plate iii.) 136-141
Contributions to our Knowledge of Soil-Fertility. No. xvi. The
Search for Toxin-Producers. By R. Greig-Smith, D.Sc,
Macleay Bacteriologist to the Society 142-190
On certain Shoot-bearing Tumours of Eucalypts and Angophoras;
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the Plants. By J. J. Fletcher and C. T. Musson. (Plates
iv.-xxvi.) 191-233
Hon. Treasurer's Financial Statement and Balance Sheet 30-34
Elections and Announcements ... 135,234
Notes and Exhibits ... ' 235-236
Issued ISth September, 1918.
Vol. XLIII. VW
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CONTENTS.
PAGES
On the Seasonal Distribution of some Queensland Species of
Arcella Ehrenberg [Rhizopoda]. Bj^ C. D. Gillies, M.Sc,
Biology Department, University of Queensland. (Four Text-
figures) 237-246
On the Leaf- Anatomy of SccEvola crassifolia [Goodeniace^], with
special reference to the Epidermal Secretion. By Marjorie
Isabel Colli>'S, B.Sc, Demonstrator in Botany, University'
of Adelaide. (Plates xxvii.-xxviii.; and six Text-figures) ... 247-259
A Fossil Insect-wing from the_roof of the Coal-seam in the Syd-
ney Harbour Colliery. By R, J. Tilly ard, M.A., D.Sc,
F.L.S., F.E.S., Linnean Macleay Fellow of the Society in
Zoology. (One Text-figure) ''... 260-264
The Panorpoid Complex. A Study of the Phylogeny of the
Holometabolous Insects, with special Reference to the Sub-
classes Panorpoidea and Neuropteroidea. [Introduction].
By R. J. Tilly ARD, M.A., D.Sc, F.L.S., F.E.S., Linnean
Macleay Fellow of the Society in Zoology ... ... ... 265-284
The Panorpoid Complex. Part i. The Wing-Coupling Apparatus,
with special Reference to the Lepidoptera. By R. J. Till-
YARD, M.A., D.Sc, F.L.S.. F.E.S., Linnean Macleay Fellow
of the Society in Zoology. (Plates xxix.-xxx. ; and sixteen
Text-figures) 286-31g
The Geology and Petrology of the Great Serpentine Belt of New
South Wales. Part vii. The Geology of the Loomberah Dis-
trict and a portion of the Goonoo Goonoo Estate. By W,
N. Bexsox, B.A., D.Sc, F.G.S., Professor of Geology and
Mineralogy in the University of Otago, N.Z., late Linnean
Macleay Fellow of the Society in Geology. (Plates xxxi.-
xxxviii. ; and four Text-figures) ... ... ... 320-360,363-384
With two Palseontological Appendices by F. Chapman, F.R.M.S.,
A.L.S,, Paleontologist to the National Museum, Melbourne.
(Plates xxxix.-xlii.) " 385-394
Studies in Australian Mec.opfera. No. ii. The Wing- Venation of
Ghorista austral is Klug. By R. J. Tilly ard, M.A., D.Sc,
F.L.S., F.E.S., Linnean Macleay Fellow of the Society in
Zoology. (Plate xliii., and four Text-figures) , 395-408
The "Springing" of Tins of Preserved Fruit. By W, W,
L'EsTRANGE and R. Greig-Smith ... 409-414
Elections and Announcements 285,362,416
Notes and Exhibits 361,415,416
Issued 18th December, 1918.
^ Vol. XLIII.
^ Part 3, No. 171
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Mesozoic Irisectsof Queensland. No. 3.. Oc?ontti!a and Protodonata.
By R. J. Till YARD, M.A., D,Sc., F.L.S., F.E.S., Linnean
Macleay Fellow of the Society in Zoologj''. (Plates xliA'.-xlv.,
and Text-figures 11-16) 417-436
The Carboniferous Trilobites of Australia. Bj^ John ^Mitchell,
late Principal, Technical College at Newcastle, N.S.W.
(Plates xlvi,-liii.) 437-494
New and rare Freshwater Algie. By G. I. Playfaik, late Science
Research Scholar of the University of Sydney. (Plates liv.-
Iviii., and eleven Text-figures) ... ... ... ... ... 497-543
The Mosses of North Queensland. By Dr. V. F. Brotherus and
the Rev. W. Walter Watts 544-567
Mesozoic Insects of Queensland. No. 4. HtmijMra Heitroptera :
the Family DiinstajiUda'. With a Note on the Origin of the
Heteroptera. By R. J. Tillyard, M.A., D.Sc, F.L.S.,
F.E.S., Linnean Macleay Fellow of the Society in Zoology.
(Plate lix., and Text-figures 17-22) ' 568-592
The Geology and Petrology of the Great Serpentine Belt of New
South Wales. Part viii. The Extent ion- of the Great Serpen-
tine Belt from the Nundle District to the Coast. By W. N.
Benson, B.A., D.Sc, F.G.S., Professor of Geology and Min-
eralogy in the University of Otago, N.Z., late Linnean Mac-
leay Fellow of the Societj' in Geology. (One Text-figure) ,.. -593-599
The Stems of Climbing Plants. By John Shirley, D.Sc, and
C.A.Lambert. (Plates Ix.-lxvi.) 600-609
The Relation between the Fat-Content and tlie Electrical Con-
dulfetivity of Milk. By H. S. Halcro Wardlaw, D.Sc,
Linnean Maeleaj' Fellow of the Society in Physiology ... 613-625
The Panorpoid Complex. Part 2. The'Wing-Trichiation and its
Relationship to the General Scheme of Venation. Bj' R.J.
Tillyard, M.A., D.Sc, F.L.S. , F.E.S., Linnean Macleay
Fellow of the Society in Zoology. (Plates Ixvii.-lxix., and
Text-figures 17-34) 626-657
A Study of the External Breathing-Apparatus of the Larvtv of
some Muscoid Flies. Bj- John L. Froggatt, B.Sc (Plate
Ixx.) ... 658-667
Notes on Australian vSa-wflies {Tcnthredinidrv). By. Walter W.
Fro(m;att, F.L.S., Government Entomologist 668-672
Elections and Announcements ... ... ... ... ...495,610
Notes and Exhibits 495,610
Issued 26 fh March, 1919.
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Tiie Macleay Memorial Volume [issued October ISth, 1893].
»yal 4to., LI. and 308 pages, witli Portrait, and forty-two plates.
-£3 3.
Descriptive Catalogue of Australian Fishes. By William
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The Transactions of the Entomological Society op New
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live Parts, 1869-73; all published], price £2, net, are also obtain-
able, but neither the Parts nor the Volumes are sold separately.
PROCEEDINGS, 1918, PART 4
CONTENTS.
PAGES
Notes Oil the Native Flora of New South Wales. Part x. The
Federal Capital Territory. By R. H. Cambage, F.L.S.
(Plates Ixxi.-lxxiv.) ... ... ... (573-711
Descriptions of new Species of Australian Gohopttra. Part xiv.
By Arthur M. Lea, F.E^S 71o-746
On a new Species or Form of Eucalyptus. By Cuthbert Hall,
M.U., Ch.M. (Plate Ixxv.y ... 747-749
Studies in Australian Xenroptera. No. 6. The Family Psycliop-
■^idte, with Descriptions of new Genera and Species, By R.
.J. Till YARD, M.A., D.Sc, F.L.S., F.E.S., Linnean Macleay
Fellow of the Society in Zoologj'. (Plates Ixxvi.-lxxviii. ;
and four Text-figures) ... 750-786
Studies in Australian Newroptera. No.7. The Life-history of
Psychopsls dtyaus ((^luerin). By R. J. Tilly ard, M.A.,
D.Sc, F.L.S., F.E.S., Linnean Macleay Fellow of the Society
in Zoolog}'. (Plate Ixxix., and twelve Text-figures)... ... 7S7-<S18
Australian Jhyalopttra or Alder-Flies, with Descriptions of new
Genera and Species,' By R. J. Till yard, M.A., D.Sc, *
F.L.S. , F.E.S., Linne^xn Macleay Fellow of the Soeiety in
Zoology. (Three Text-figures) 81!J-82o
Contributions to a knowledge of Australian Culicidfv. No. iv.
By Frank H. Taylor, F.P],S. (Plates Ixxx.-lxxxiii.) ... 826-843
Note on tiie Temperature of Echidna aadeata. By H. S. Halgro
Wardlaw, D.Sc, Linnean Macleay Fellow -of the Society in
Physiology. (Two Text-figur§s) "... * 844-841J
The Occurrence of Methyl L;evo-Inositol in an Australian Poison-
ous Plant. By James M. Petrie, D.Sc, F.I.C, Linnean
Macleay Fellow of the Society in Biocheniislry. (Two Text-
figures) 850-867
An Account of some Observations upon the Life-history of Phoma
cif ricarpa McAlji. : the Cause of the "Black Spot"' Disease
in Citrus Fruit in New SoutK Wales. By (i. P. Darnell-
S.MiTH, D.Sc, F.I.C. (Plates Ixxxiv.-xc.) ... 868-882
■TheOccurrenceof an inverted Hj'menium in Aijuricn-^ ('(nupeMrl^,
liy(^. P. Darnell-S.hitii, D.Sc, F.I.C. (Plates xci.-xciii.) 883 887
Notes and Exhibits 713-714
Donations and Exchanges ... ... . . . . 888-902
Title-page ... ... ... .. '■
Contents ... ... ... ■■ . ■• ■ ■ • iii-
Corrigenda ... ... ... ... •• •■■ •■ vii.
Li.st of New Generic and other Names - vii
List of Plates viii-
Index i.-xxix.
MBL WHOI LIBRARY
IjdH lADP C
i