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TRANSACTIONS
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BOYAL,. SOCIETY OF | VilGiORaLA.
VOD: Is.< = PART:
1890.
CONTENTS.
THE TRANSVERSE SECTIONS OF PETIOLES OF EUCALYPTS AS
AIDS IN THE DETERMINATION OF SPECIES, sy D. McAtrrnz,
F.C.S., anp J. R. Remrrey. (With Plates 1, 2, 3, 4, 5, 6, and 6a) =
THE VICTORIAN LAND PLANARIANS, sy Artnur Denpy, M.S8c., F.L.S.,
Demonstrator anD Assistant LecrurER IN BioLogy IN THE UNIVERSITY
or Metzourne. (With Plate 7) . - - - - - =
THE EUCALYPTS OF GIPPSLAND, sy A. W. Howitt, F.G.S. (With
Plates 8, 9, 10, 11, 12, 13, 14, 15, 16) - = = = : :
A NEW FAMILY OF HYDROIDEA, TOGETHER WITH A DESCRIPTION
OF THE STRUCTURE OF A NEW SPECIES OF PLUMULARIA,
py W. Banpwin Spencer, M.A., Prormssor orf BroLogy IN THE UNIVERSITY
oF Mretgourne. (With Plates 17, 18, 19, 20, 21, 22,23) - - -
EDITED BY PROFESSOR W. BALDWIN SPENCER, M.A., HON. SEC.
}
_
~' MELBOURNE.
PUBLISHED FOR THE ROYAL SOCIETY
PAGE
81
BY THE SPECTATOR PUBLISHING COMPANY LIMITED, 270 POST OFFICE PLACE.
Aprit, 1891.
ened, SOCOM PY OR VICTORLA.
Patron :
HIS EXCELLENCY THE EARL OF HOPETOUN, G.C.M.G.
President :
PROFESSOR W. C. KERNOT, M.A., C.E.
Vice-PPresidents :
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tbon. Treasurer :
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‘bon. Secretaries :
H. K. RUSDEN, F.R.G.S.
PROFESSOR W. BALDWIN SPENCER, M.A.
bon. Librarian :
JAMES EK. NEILD, M.D.
C. R. BLACKETT, F.C.S.
J. BOSISTO, C.M.G.
R. L. J. HLUERY, F.RB.S.
G. 8. GRIFFITHS, F.R.G.S.
xh W. HOWLED? BGs:
A. H. 8. LUCAS, M.A., B.Sc.
Council :
PROF. ORME MASSON, M.A., D.Sc.
| H. MOORS.
PROF. R. T. LYLE, M.A.
ALEXANDER SUTHERLAND, M.A.
, C. A: TOPP, M.A., ULB.
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TRANSACTIONS
OF THE
Oe we OC mY Ob VICTORIA:
VOR. PARI.
1890.
CONTENTS.
THE TRANSVERSE SECTIONS OF PETIOLES OF EUCALYPTS AS
AIDS IN THE DETERMINATION OF SPECIES, sy D. McAtrring,
F.C.8., anp J. R. Remrrey. (With Plates 1, 2, 3, 4, 5, 6, and 6a) :
THE VICTORIAN LAND PLANARIANS, sy Artuur Denpy, M.Sc., F.L.S.,
DEMonsTRATOR AND AssiIsTANT LecTURER IN BioLoey In THE UNIVERSITY
or Metsourne. (With Plate 7) - - : - - - -
THE HEUCALYPTS OF GIPPSLAND, sy A. W. Howirr, F.G.S. (With
Plates 8, 9, 10, 11, 12, 18, 14, 15, 16) - - = - - .
A NEW FAMILY OF HYDROIDEA, TOGETHER WITH A DESCRIPTION
OF THE STRUCTURE OF A NEW SPECIES OF PLUMULARIA,
BY W. Batpwin Spencer, M.A., Proressor oF BroLtoay in THE UNIVERSITY
or MetBournE. (With Plates 17, 18, 19, 20, 21, 22,23) - - =
EDITED BY PROFESSOR W. BALDWIN SPENCER, M.A., HON. SEC.
MELBOURNE.
PUBLISHED FOR THE ROYAL SOCIETY
PAGE
81
121
BY THE SPECTATOR PUBLISHING COMPANY LIMITED, 270 POST OFFICE PLACE.
ApRiL, 1891.
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ARTICLE I.—Tue Transverse Sections or Pretriones or EUCALYPTS AS AIDS IN
THE DETERMINATION oF Species, By D. McArping, F.C.S., anp J. R. Remrry.
(With Plates 1, 2, 3, 4, 5, and 6.)
(Read Thursday, November 14th, 1889.)
I.—IntrRoDUCTORY.
The Eucalypts form such a prominent feature in the Australian vegetation, and
have such varied uses, that any addition to our knowledge of their characters is sure
to be welcomed.
The primary object of the present paper is to show how transverse sections of
the petioles of Kucalypts may be used as valuable aids in the determination of
species. Sections of thirty different kinds are here described and photographed, and
they show unmistakably, along with a general resemblance, differences which are
more or less constant, and readily recognisable for each species.
Anatomical characters of the leaf have already been successfully used in the
discrimination of species belonging to other divisions of the vegetable kingdom.*
Of course such characters, depending on a single organ, and apart from the aggregate
of characters, are apt to be more or less artificial ; but, when we consider that the
parts shown in transverse section of the petiole are in organic connection with, and
form an essential portion of, the vital machinery of the plant, it need not excite
surprise that they should vary in the different species, and be, to a certain extent,
characteristic for each.
We know that the leaf is simply a lateral expansion or extension of the stem or
branch, usually bearing a bud in its axil, and its tissues are, as a rule, continuous
with those of the stem, so that in it we have an epitome of the parts concerned in
vegetative life—a point deserving of special notice in this connection. Besides, the
petiole, with which we are more particularly concerned, is capable of reproducing
the entire plant from a small portion of it, as in the well-known instance of the
ipecacuanha plant. The vegetative organs which, in some form or another, are
absolutely necessary for the life of the individual, have been too little used in the
discrimination of genera and species, while the reproductive organs have been too
often almost exclusively relied upon. The anatomist (dealing with internal structure)
and the systematist (often entirely occupied with external characters) must combine
their results in order to arrive at a proper conception of the true system of nature.
* See De Bary, Comp. Anat. of Phanerogams and Ferns, p. 298.
2 THE TRANSVERSE SECTIONS OF PETIOLES OF EUCALYPTS
Perhaps one result of this investigation will be to direct attention to the
distinctive nature of the tissues of the leaf in other groups of plants as well as the
Eucalypts. This has already been done, for instance, in the Gnetaceae and Coniferae
by Dr. C. EK. Bertrand* and Prof. M‘Nab.+ The latter confined his attention to the
Coniferae, and, in addition to the general anatomic structure of the transverse section
of the leaf, noted the distribution and number of the resin-canals, and the arrange-
ment of stomata on the surface, as aids in his diagnosis.
While we have likewise described and drawn the transverse section as a whole, and
found the total characteristics of the section of any species to be of great value for
identification, there are certain features very useful for that purpose, even when
taken alone. Thus, the cortical cavities, whether many or few, large or small; the
size and shape of the section; the relative thickness of the epidermis, &c.; but it is
the woody-tissue forming characteristic figures which is the most striking part.
This wood-pattern is fairly constant for each, so that the most enduring part of the
entire section fortunately gives a clue to the affinity sought for.
It would appear that even from an economic standpoint some reliable means
are much needed for determining a given species of Kucalypt, from a readily accessible
portion of the plant, such as the leaf. It is highly desirable, in the interests of
trade, not to speak of science, that one species should not be confounded with
another, for the timber has such specific variations that serious loss and mischief
might in many cases ensue.
Mr. Maiden, of the Technological Museum, Sydney, in his recently published
work on the ‘“‘ Useful Plants of Australia,” writes on this subject, at p. 427, in very
decided terms :—‘‘ Scarcely a branch of Australian economic botany is in a more
confused state than that which pertains to the timber of the Eucalypts. The genus
is, perhaps, the most difficult one in the world, intrinsically, and also because of
accidental circumstances, t.e., difficulty of obtaining flowers and fruit, and irregular
flowering seasons; moreover, the trees vary, according to climate and soil, to such
an extent as to render a definition of the species rather expansive, and, as this
difficulty often extends to the wood, timbers of totally different character are
sometimes reckoned under the same species.”
Such a state of matters calls for some remedy, and it is evident that the present
investigation tends in that direction. The leaves being evergreen are a constant,
and not an accidental circumstance. The deep-seated characters revealed by the
section are not so variable as others more dependent on soil and climate; and a
definition which is included in, and shown by, the transverse section of a petiole,
can hardly be called expansive.
* Anatomie Comparée des Tiges et des Feuilles chez les Gnetacees et les Coniferes. Paris, 1874.
+ Proc. Roy. Irish Acad., Series II., Vol. Il., 1875-77. Remarks on the Structure of the Leaves of certain Coniferae,
and a Revision of the Species of Abies.
AS AIDS IN THE DETERMINATION OF SPECIES. 3
In the masterly monograph on the Eucalypts of Australia and adjoining islands,
the ‘ Eucalyptographia,” by Baron von Mueller, there are numerous references to
the anatomic structure, both of the wood and the leaf-blade, although none of them
are concerned with the petiole. In one instance (Decade 7) he has given transverse
sections of Eucalyptus wood, accompanied by the remark :—‘‘ These microscopic
sections are given to aid in the discrimination of mercantile Kucalyptus timber of
doubtful origin, such as sometimes occurs in the trade.” Here we have a distinct
recognition of the idea, and a practical expression of it, that the minute structure
of the wood may be useful in determining species, and be serviceable from the
timber merchant’s point of view. Similarly in the wood-pattern of the leaf-stalk,
there are serviceable characters for the same purpose.
It may be noted here that the existence of such a work as the ‘ Eucalypto-
graphia’’ invests the present investigation with a special importance. There the
different species of Kucalypts are fully described and carefully drawn, and we have
the advantage of working out a genus from which we can select numerous well-
defined species.
The species selected on the present occasion are such as were conveniently
obtainable from the Melbourne Botanic Gardens. Our best thanks are due to Mr.
Guilfoyle, F.L.8., Director of the Gardens, for kindly supplying us with fresh
specimens when required. Thirteen of these are Victorian, and as there are 38
altogether, according to the ‘“‘ Key to the System of Victorian Plants,’’ we hope to
deal with the remainder in a future paper.
We are also much indebted to J. Bosisto, Esq., C.M.G., for supplying us with
the leaves of some of the oil-yielding species from their native habitats, and for
information contained in various papers read at different times before learned
societies.
Above all, we have to thank Baron von Mueller for verifying each of the
species used in this investigation, and thereby enhancing the value of the work—
the determination coming from such an undoubted authority on this special group
of plants.
The type sections were prepared and the photographs taken by Mr. Remfry.
Since the work has been in hand for over four years (the first sections having
been cut about September, 1885), it has been possible to show the structure of
the leaf-stalk at different seasons, and even in different years; and, as over a
thousand sections have been made, a fair idea of the incidental variation is likewise
given.
4 THE TRANSVERSE SECTIONS OF PETIOLES OF EUCALYPTS
LIST OF FORMS INVESTIGATED, ALPHABETICALLY ARRANGED.
Boranicat Name. Common Name.
1. E. alpina, Lindley a8 a0 00 VY. Alpine Gum
2. ,, amygdalina, Labillardiére .. 30 Vy. hee Tree
3. ,, calophylla, R. Brown oo on Calophyllum-like Eucalypt
4. ,, citriodora, Hooker .. o6 fe Lemon-scented Eucalypt
5. ,, cornuta, Labillardiére 36 BO “Yate”
6. ,, corynocalyx, F. v. M. 06 50 V. Sugary Eucalypt
7. ,, diversicolor, F. v. M. a0 am “ Karri”
8. ,, ficifolia, F.v.M. .. 3 ae Scarlet-flowered Eucalypt
9. ,, globulus, Labillardiére i = V. Blue Gum Tree
10. ,, gomphocephala, Candolle .. ute ‘‘ Touart ”
11. ,, pachypoda, F.y.M... 00 ae Thick-stalked Eucalypt
12. ,, gunnii,J.Hooker .. an a0 V. Cider Eucalypt
13. ,, lehmanni, Preiss .. dc 30 Lehmann’s Eucalypt
14. ,, leucoxylon, F. v. M... ne ae VY. Victorian Ironbark Tree
15. ,, macrorrhyncha, F.v.M. .. 56 V. Victorian Stringybark Tree
16. ,, maculata, Hooker .. 50 oo Spotted Eucalypt
17. ,, marginata, Smith .. te KA “ Jarrah ”’
18. ,, megacarpa, F. v. M... 60 oe West Australian Blue Gum Tree
19. ,, melliodora, Cunningham .. 56 V. Yellow Box Eucalypt
20. ,, obcordata, Turezaninow .. bc “* Maalok ”
21. ,, obliqua,l’Heritier .. . a V. Messmate Stringybark Tree
22. ,, occidentalis, Endlicher ie ai Flat-topped Yate
23. ,, punctata, Candolle .. Se oO Leather Jacket
24. ,, rostrata, Schlechtendal aa a VY. Red Gum Tree
25. ,, rudis, Endlicher Sa Fs 5 Swamp Gum Tree of Western Australia
26. ,, saligna, Smith ae ie 30 Grey Gum
27. ,, sStuartiana, F.v. M. .. ete ote V. Apple-scented Eucalypt
28. ,, tereticornis, Smith .. ah =f VY. Flooded Gum Tree
29. ,, tetraptera, Turczaninow .. no Four-wing-fruited Eucalypt
30. ,, viminalis, Labillardiére He A V. Manna Eucalypt
Thirteen Victorian species (V), out of a total of thirty-eight, recorded in ‘‘ Key to the System of Victorian Plants,”
by Baron Ferd. von Mueller.
IJ.—Mope or Preparation AnD Mountina oF SEcTIONS.
From the fresh branches not less than five of the most vigorous-looking leaves
were selected. They were taken from different parts of the branch, but never from
the growing points. The petioles, with a portion of the lamina attached to each to
show the direction of growth, were placed in methylated spirit, and after a few days
were usually found in good condition for cutting. If still too hard they were boiled
in water for a short time.
The sections were taken from the basal half of the leaf-stalk—from a point
nearer to its base than its junction with the blade. Those intended to be sketched
by the camera lucida were cut by hand, the thinnest being selected from a large
number taken from each petiole, and examined in a mixture of equal parts of
glycerine and spirit.
,
AS AIDS IN THE DETERMINATION OF SPECIES. 5
The sections for the permanent type-slides were cut with a microtome, after
embedding in paraffin in the usual way. To clear the sections for these mounts, a
mixture of glycerine and caustic soda solution was generally used. Warming in
nitric acid was in some instances attended with brilliant results, while in other cases
the sections were ruptured and distorted by its action.
Most of the sections were photographed without any preparatory staining.
Indeed, in some instances, especially in the case of petioles kept for many months in
ordinary tank rain water (and some kept for over three years were not in the least
decomposed or affected by fungus growths), the colour acquired by the specimens—a
rich yellowish-brown tint—was itself a stain not to be surpassed for the purposes of
actinic contrast. When this tint was not acquired, and in the case of fresh material,
the sections were, if necessary, bleached by the action of free chlorine, and then
faintly stained with the stain first proposed by Dr. Frances Hoggan.* It is prepared
by adding pyrogallic acid to tincture of steel. This was found more easy of
application than the iron stain, composed of pyrogallic acid and protosulphate of iron,
recently recommended in the Journal of the Royal Microscopical Society,+ which,
however, gave equally good results to the eye and on the photographic plate.
The Hoggan iron stain was originally recommended as especially suitable for the
treatment of cartilage, in which the desired depth of colour could be produced in a
few minutes.
We found that the time required for the proper treatment of the sections of
petioles varied from five minutes to half an hour, the stain here taking a longer time
to penetrate.
The source of illumination was a clock-work petroleum Jamp with a one-inch
wick, and the light was all that could be desired, but a blue glass modifier was used
in every case, whether the object was stained or not.
Of the mounting media tried, glycerine jelly was found the most suitable.
Balsam was used in a few cases, but it was invariably found to render the epidermal
layers too transparent, and consequently somewhat indistinct. It may be mentioned
that it was found to be essential for purposes of comparison and identification that
the sections should be quite thin, showing little more than one layer of cells in the
cortical ground tissue.
* Marsh, ‘‘ Microscopical Section-cutting,” 2nd Ed., p. 131,
+ Journ. Roy. Micro. Soc., 1888, p. 157.
6 THE TRANSVERSE SECTIONS OF PETIOLES OF EUCALYPTS
III].—CuasstFicatTion oF Tissunrs.
Before describing the sections, it will be well to settle the system of classification
to be adopted for the tissues.
Various methods have been proposed, and more or less generally used. Of these
we will briefly glance at four of the most important, and indicate the one most
convenient for our present purpose.
One system which has been very widely used is the topographical of Sachs. In
this the tissues are mapped out according to their relative positions—the dermal
tissue, the ground or fundamental tissue inside, and the fibro-vascular bundles or
fascicular tissue studding the ground tissue. This threefold division is certainly very
simple and easily applied, but it is not scientific, for geographical position does not
always reveal the essential character of a tissue.
A second system is the developmental of Hanstein, and, at first sight, it seems to
be thoroughly scientific, being based upon development. In this classification certain
meristem layers form certain tissues, and these get special names. There is a three-
fold arrangement of the primary layers, as in animals, viz., dermatogen, forming the
epidermis ; periblem, forming the cortex ; and a central cylinder of plerome, forming
vascular bundles and pith. But the objection to this system is that it does not
always hold good, for, as instances, in perforations of the leaf in many aroids, and in
splitting of the leaf into segments in palms, the original epidermis is replaced by
another, derived from the underlying tissue.
The morphological system of De Bary is purely descriptive, and, therefore, the
safest to adopt in the present state of our knowledge. We know so little of function
as yet that we can do little more than place together parts agreeing in structure,
although they may have little else in common, or no physiological relationship.
Parenchyma is thus one kind of tissue, sclerenchyma another, and so on, and this is
the classification most convenient for present use.
But convenience is only a make-shift, and sooner or later function and structure
together will determine the true position of any system of tissues. ‘This brings us to
the last system to be mentioned, viz., the functional of Schwendener and Haberlandt,
in which a certain structure is associated with a certain function or functions. The
function is here brought to the front, and such terms as bast, cambium, Wc., are used
to denote tissues performing special functions, and not as is usually the case, tissues
haying special positions. Certain tissues may haye a certain position by virtue of
their function and consequent structure, but that is an accidental, and not an essential
circumstance. Relative position and structure are not considered apart, but as
dependent on function. As already stated, function is as yet so little understood that
AS AIDS IN THE DETERMINATION OF SPECIES. 7
the determination of it is often but a guess at the truth, but this guess can either be
verified or proved erroneous, and so advance in this direction will be made. This
mode of classification will be the final outcome of all our studies, and to show its
superiority we cannot do better than take the fibro-vascular bundle as an example.
The F.V. B.,* according to the ordinary view, and as the name bundle indicates,
is a unity, and treated as such. It is composed of a variety of cells, usually having
a definite relation to each other, and either hard and fibrous, or soft and
parenchymatous or vascular.
On the functional view, the F. V. B. is not a unity, but composed of at least two
main elements—a part for mechanical support, or skeleton, and another part for
circulation. The skeleton and the circulating tissues are often associated, just as the
blood vessels have a certain relation to the skeleton in the animal body—the hard
and firm to protect and support the soft and yielding parts.
In describing the sections generally we will adopt a combination of the two latter
modes of classification, giving the separate parts, along with their respective
functions. Perhaps this might be called, for distinction’s sake, the bzomorphic
system, as both structure and function are taken into account, and it is undoubtedly
the most scientific of all, for the structure did not precede and the use follow, but
most probably the necessity for the use arose and the structure was forthcoming.
Since life is the cause and not the consequence of organisation, and as life may be
regarded as the sum total of the functions, and organisation as the sum total of the
structure of the plant or animal body, therefore structure is dependent on—is a
manifestation of—function.
The biomorphic classification of the tissues is based upon the structure, taken in
conjunction with the function, the mechanism in connection with its use, and it is
added mainly to show the importance of certain structures for purposes of
classification. When itis remembered that in a transverse section of even a petiole,
we have structures which serve for protection and support, for the manufacture and
the transport of food materials, and in fact, for nearly all the vegetative functions of
the plant, as distinguished from the reproductive, then such structures become
invested with a higher meaning, and with a corresponding classificatory value.
Adopting a convenient arrangement of parts, which can be seen at a glance in
any of the sections, and which will afterwards be further subdivided, they will be as
follows from the outside inwards :—
1. Epidermis or protective tissue.
2. Cortex, or principally starch-manufacturing tissue.
3. Hard bast, or supporting tissue.
Fibro-vascular bundle /4. Soft bast, or albuminoid-forming tissue.
5. Wood tissue, or circulatory and supporting tissue.
* BF. V. B. is used instead of repeating in full the expression, Fibro Vascular Bundle.
8 THE TRANSVERSE SECTIONS OF PETIOLES OF EUCALYPTS
With the much-debated question, whether the hard bast really belongs to the
vascular bundle or not, we have at present nothing to do. These bundles of bast
fibres always accompany the vascular tissue in these transverse sections, and while
they are often absent in other cases, and may be unessential, yet as a matter of
convenience, and without entering upon the general question, we will use the current
expression of fibro-vascular bundle to include the hard bast and the soft bast, as well
as the woody tissue.
lV.—Sections Descripep GENERALLY.
E. globulus taken as a type.
As each section is photographed and accompanied by its appropriate description,
it will only be necessary here to notice the general character of the sections. For
this purpose we will take Eucalyptus globulus as a type, and compare the others
with it.
When an ordinary hand-cut transverse section is mounted direct in Schulze’s
solution, the differentiation of the tissues is clearly shown. ‘Taking a general view
of the section before entering into details, it is seen to be of an oval shape, one
and a half lines broad, by one line thick, or one and a half times broader than thick.
The epidermis appears as an orange-yellow, or it may be pale crimson border, edged
by a continuous pale yellow line, this latter indicating the presence of a cuticle.
Beneath the epidermis is the purple-coloured cortex, composed of cells with walls
varying in thickness, and as the colour indicates, of cellulose. Next comes the
hard bast, with its orange-yellow lignified cell walls, succeeded by the purple-coloured
soft bast; then the clearly-marked bright yellow cells and vessels of the wood,
traversed by narrow radiating lines, composed of elongated cells of a darker colour.
These are the parenchymatous cells of the wood resembling medullary rays. Next
there is a continuation of the soft bast on the other side of the wood, succeeded by
the hard bast, and then the other tissues as above. In short, if we start from the
curve of the wood, which is more or less central, forming a woody core, there is a
succession of envelopes more or less thick. The soft bast surrounds the wood, and is
wedged in between it and the hard bast, which is surrounded in turn by the cortex,
and the epidermis envelopes all.
Each of the above parts will now be specially dealt with in succession.
1. Epidermis or Protective Tissue.—The cells of the epidermis are well-defined,
and distinct from the underlying tissue. It consists of a single layer of cells,
measuring in thickness 1-570in., and the breadth of each cell on an average is
about 1-800in. (Fig. 1, a.)
AS AIDS IN THE DETERMINATION OF SPECIES. 9
The outer wall is moderately thickened to prevent excess of evaporation, and
generally to provide against injury. In the ordinary unstained cell there may be
distinguished outside of the round or oval cell cavity a distinct clear band externally,
and a somewhat turbid-looking portion abutting on the cell cavity. As the outer
band is coloured a pale yellow by Schulze’s solution, it represents a cuticle, which is
continued right round the section, forming a slight convexity on the outside of each
cell. Itis on an average from 1-3200in. to 1-2400in. in thickness, and shows no
sign of striation.
The turbid-looking portion of the wall is coloured a deeper yellow by Schulze’s
solution, and may even pass into orange-yellow or pale crimson, according to the
amount of colouring matter present. Hence it may be concluded that beneath the
fine sheet of cuticle, which is of the nature of cork, there is a cuticularised layer, a
sort of transition between cellulose on the one hand and cuticle on the other. The
use of the cuticle and cuticularised layer is evident, principally for the purpose of
preventing evaporation. It is very thick on desert plants. Plants in rainless regions,
and Australian plants generally, are well provided with it. The sections of the
Mallee scrub species, to be afterwards referred to, will show this clearly.
The lateral walls unite the cells together, and are relatively thin. They
terminate outwardly at the cuticle, and leave no gaps between. No contrivances for
admitting air were found, although carefully looked for. The inner wall is likewise
thin, and may be composed of cellulose, being coloured purple by Schulze’s solution,
thus allowing nutritive substances to pass in and nourish the living cell. Or it may
be, like the outer wall, coloured yellow, and thus cut off from nutritive supplies, so
that no further growth or expansion can take place. In the one case the epidermal
cell is living, and capable of further growth, in the other it has reached maturity,
and become surrounded by a corky wall.
From the relative thickness and breadth of each epidermal cell, it may be
inferred that it has the form of a prism perpendicular to the surface, and the cuticle
covering produced by each, forms a continuous layer all round the outside.
2. Cortical Tissue, or starch-manufacturing tissue principally.—Immediately
beneath the epidermis there is a single layer of cells, which may or may not have
their outer and lateral walls cuticularised, and which form a more or less distinct
outer border to the cortex proper. This may be called the Hypodermal layer
(Fig. 2).
The remaining cells are of irregular outline, with walls of various degrees of
thickness. The cortical parenchyma contains chlorophyll, and it is here the
manufacture of starch is carried on, as evidenced by the contents of the cells
10 THE TRANSVERSE SECTIONS OF PETIOLES OF EUCALYPTS
containing numerous starch granules. When the section is treated with iodine this
portion becomes visibly violet, showing the presence of starch in abundance.
As the hard bast is approached the walls usually thicken considerably, and the
cell cavity is correspondingly diminished (Fig. 2a). The purple colouration by
Schulze’s solution shows, however, that the cell walls are of cellulose. Here we are
specially concerned with that form of thick-walled parenchyma, known as Collen-
chyma, or collenchymatous parenchyma.
Sometimes the walls are thickened at the angles, or often completely round.
While the epidermis is a protective tissue, this is a supporting one, serving to
strengthen the comparatively slender petiole. It is common in parts of plants, such
as the leaf-stalks, where provision has to be made for increase of length as well as for
strength, and so the cells must be living, and have thin places to admit the necessary
nutriment—they are living mechanical cells. In the old and fully-formed leaf, the
thickening may extend all round (as seen in Fig. 4). If the living protoplasm were
then to disappear, and the walls become thickened and lignified, this tissue would
pass into that of the hard bast.
The cortical tissue is excavated by large and numerous cavities, and crystals are
very frequent, both of which will now be specially noticed (Figs. 2 and 3).
These cavities may be named according to their position, rather than that of
their contents (which has yet to be accurately determined), and so we may speak of
them as cortical cavities. In contrast to these cavities, which are not continuous
lengthwise, and so vary in their number and arrangement at different parts of the
petiole, there are in some species canals which are constant and continuous, and
from their position inside the wood curve, they may be termed central canals.
The contents of these cavities or canals cannot be called resins, for they are
soluble, sometimes very readily in water ; nor can they be called oleo-resins, for the
petioles yield no oil on distillation. They may be spoken of as kinoid, until their
chemical nature is more accurately determined.
The kinos of the Eucalypts have been, and are being, investigated by Maiden,*
and he finds that they are divisible into groups according to their behaviour with
water and spirit. These groups will be referred to in connection with classification.
Cortical Cavities.—These cavities, usually roundish or oval in shape, occur at
irregular intervals, and at some little distance from the epidermis (Fig. 2). They vary
in size and in number. The largest is about 1-80in. in diameter. They are
distinctly visible to the naked eye when held up to the light. The number is usually
from 7 to 8.
* Proc. Linn. Soc., N.S.W. Vol. IY., Series 2.
AS AIDS IN THE DETERMINATION OF SPECIES. 11
The contents are usually of a yellowish, but in some instances of a dark-red
colour, readily dissolved by alcohol, and deeply stained by tincture of alkanet. But
if allowed to harden by keeping in water for a lengthened period, neither alcohol nor
ether affects them.
Each cavity is bounded by much-compressed cells, with their flat faces towards
the cavity. From the appearance presented by these intercellular spaces or excava-
tions, they appear to be formed by the rupture and disorganisation of the cells at
that particular spot, and are hence of lysigenous origin.
In some species there are special canals, not cortical, but enclosed by the curve
of the wood, which will be noticed in their proper place.
Crystals.—Crystals of the well-known octahedral form are seen in great profusion
scattered through the cortical cells (Fig. 8). They vary considerably in size,
sometimes reaching 1-$00in. in length, and, judging from the usual tests, as well as
from their crystalline form, consist of oxalate of lime. Numerous crystals, similar
to the above, likewise occur in the soft bast, but they are much smaller on the
whole. No species examined was entirely free from crystals, although in some
instances they were very scarce.
Longitudinal Section Although it is not the object of this paper to give a
complete account of the histology of the petiole, it was thought necessary to
examine longitudinal, as well as transverse sections, at least in one or two cases.
A longitudinal median section of the petiole of E. globulus, taken through the
shorter axis of the crescentic bundle, revealed the ordinary succession of the various
parts.
The epidermis, after prolonged staining with Schulze’s solution, stood out as a
somewhat pale, lemon-yellow border. The breadth of each cell varied from 1-760in.
to 1-830in., and the length was just a little less, or about 1-920in.
The purple-coloured cortex exhibited much smaller cells towards its outer than
its inner portions, the hypodermal layer usually quite distinct beneath the epidermis.
Cortical cells adjoining the hard bast were often relatively large (Figs. 2a and 0).
The cortical cavities had the same form as in transverse section, being irregularly
round, or oval, or pear-shaped, and the size was about the same. Two adjoining
cavities were measured, one of which was 1-190in., and the other 1-115in. in length.
The F. V. B. showed the hard bast usually in two distinct layers (an outer and an
inner), with intervening soft bast, and the wood with its numerous spiral and annular
12 THE TRANSVERSE SECTIONS OF PETIOLES OF EUCALYPTS
vessels (Fig. 5). The soft bast was carefully examined, and found to consist of
cambiform cells and sieve-tubes (Figs. 6 and 7). The cambiform cells varied
considerably in shape and size. Often they were brick-shaped, and even somewhat
roundish ; at other times they were much more elongated and thinner-walled.
Directly adjoining the wood the cells were clearly marked off from the latter, and
sufficiently thin-walled in some cases to resemble cambium.
The sieve-tubes are shown in longitudinal section. (Fig. 7.) The iron stain,
as well as Schulze’s solution, serves to differentiate them.
F1pRo-VASCULAR BUNDLE.
This is a convenient collective term for the group of tissues already separately
mentioned as hard bast, soft bast, and wood tissue, or if the bast and wood be
simply contrasted, then phloem and xylem may be used.
Since the bundles form the most striking feature of these sections, and are the
parts from the form of which the species of Kucalypts are chiefly determined, they
will be expected to receive a large share of attention and illustration.
The special form of the bundles, or rather of the woody portion of them, will be
considered in the next section devoted to the determination of species, so little will
be said here upon that point.
It is to be premised, first of all, that the xylem portion of a leaf bundle is, as a
rule, towards the upper surface, and the phloem portion towards the lower surface,
just as would naturally happen in a lateral expansion of these tissues of the stem.
Also, that the upper or inner surface of the petiole has often a different contour to
the lower or outer surface. In this instance the upper surface is flattish, while the
under surface is rounded. But in cases such as the fig and walnut, where the petiole
is cylindrical, the bundles are likewise arranged in a circle, and form a cylinder.
As is well known, in most Eucalypts the blade of the leaf is not spread out
horizontally, as is usually the case, but vertically in relation to the leaf-bearing axis.
In the young condition, however, it has the normal position, and it is by subsequent
twisting of the leaf stalk that the changed position is brought about. In the course
of this twisting process, stomata are developed on the upper surface, in addition to
those already existing on the under surface, and palisade parenchyma is developed
beneath the lower epidermis as well.* But it appears that the bundle of the leaf
* Bower and Vines, Practical Botany, Part I., 2nd Ed., p. 148.
AS AIDS IN THE DETERMINATION OF SPECIES. 13
stalk ig unaffected by the change, or at least not rendered symmetrical, so that in
every example investigated it does not form a complete circle, but a curve open
upwards, with the two ends approaching, but never meeting.
A section of the base of the midrib of the young sessile leaf (Fig. 8) lkewise
shows the curve open upwards, whereas a section of the young square stem of the
saine plant (Fig. 9) exhibits a symmetrical quadrangle, corresponding with the
outline of the stem, and continuous right round.
An attempt to account for the generally different distribution of the wood in the
stem and in the leaf-stalk is made by Herbert Spencer in his ‘‘Principles of Biology.’’*
After noticing that there is a direct relation between mechanical stress and the
formation of wood, and that the general arrangement in stems is that of a cylinder,
he says :—‘‘ While axes are on the average exposed to equal strains on all sides,
most leaves, spreading out their surfaces horizontally, have their petioles
subjected to strains that are not alike in all directions ; and in them the hard tissue
is differently arranged. Its transverse section is not ring-shaped, but crescent-
shaped ; the two horns being directed towards the upper surface of the petiole.
That this arrangement is one which answers to the mechanical conditions, is not
easy to demonstrate; we must satisfy ourselves by noting that here, where the
distribution of forces is different, the distribution of resisting tissue is different.
And then, showing conclusively the connection between these differences, we have
the fact that in petioles growing vertically and supporting peltate leaves—petioles
which are therefore subject to equal transverse strains on all sides—the vascular
bundles are arranged cylindrically, as in axes.”
That this is only a partial explanation is shown by the fact that in
many instances the cylindrical arrangement of the wood occurs where the
distribution of forces is different from that of the stem, as in the leaves of
fig and walnut, already mentioned. It is true, however, that in all the
petioles of Eucalypts examined the crescentic form prevailed, and was not
appreciably interfered with by the twisting of the leaf-stalk. In some cases, such as
that of E. ficifolia, where the horns of the crescent may approach so very close as to
make the wood practically a cylinder, the outline was not cylindrical, and there was
less twisting of the leaf-stalk than in many other species. Indeed, in one petiole
of this species the wood formed a compressed hemisphere, the two horns abutting,
but as four central canals were present, this petiole was evidently somewhat
abnormal.
The bundle will now be considered under three separate divisions, viz., hard
bast, soft bast, and wood.
* Vol. II., p. 261.
14 THE TRANSVERSE SECTIONS OF PETIOLES OF EUCALYPTS
3. Hard Bast.—This tissue is very easily recognised in transverse section from
its position and structure. It forms an irregular, interrupted ring of exceedingly
thick-walled cells. It passes not only round the outside of the bundle on its lower
surface, but curves round the upper surface, and after dipping towards the centre of
the bundle, and forming a thickened mass, it continues in a graceful curve.
Treated with Schulze’s solution, the bright, yellow girdle is made to stand out in
bold relief.
The hard bast, which has received various other names, such as sclerenchyma,
bast fibres, prosenchyma, &c., is one kind of mechanical or supporting tissue—a
skeleton in fact. This particular kind of tissue may occur elsewhere than in this
region, wherever, indeed, a strengthening of the parts becomes necessary, but in
order to prevent confusion the name of hard bast or bast-fibres will invariably mean
with us the skeleton accompanying, supporting, and protecting the soft bast.
4, Soft Bast.—The soft bast, all but completely enclosed by the hard bast, is a
ring of tissue immediately adjoining and surrounding the wood. It consists of cells
of much smaller diameter than those of the hard bast, and relatively thin-walled.
In a section treated with Schulze’s solution, this tissue is strikingly shown as a
purple patch, between the bright yellow of the hard bast on the one hand, and that
of the woody tissue on the other, which it completely envelopes as with a mantle.
The thin-walled cells, many of them with large cavities, and all with cellulose
walls, have evidently a different function from that of the hard bast. ‘The soft bast is
composed of two main elements—the cambiform cells and the sieve-tubes, both of
which have already been dealt with in considering the longitudinal section.
5. Woody Tissue.—The wood forms the central figure of the section, and with
its graceful infolding at each side gives rise to characteristic patterns. It sometimes
resembles the inverted volute of an Ionic capital, or simply a capital © lying flat, or
to change the figure, when, with Schulze’s solution, the radiating lines are made
to stand out dark upon the yellow back-ground, it resembles a caterpillar coiled up at
either end, with its body divided into numerous segments by beaded rings: When
the two horns curve round the central canals, as in E. maculata, there is something
of an owl-like appearance about it, when viewed upside down.
In a section treated with Schulze’s solution it is easy to resolve the woody tissue
into three elements—
(a.) Wood fibres.
(b.) Wood parenchyma.
(c.) Vessels.
AS AIDS IN THE DETERMINATION OF SPECIES. 15
The wood fibres, forming mainly the outer half of the woody curve, are
recognised by their thickened walls and comparatively small cavity. They are on an
average 1-3000in. in diameter.
The wood parenchyma consists of small elongated cells, arranged in radiating
lines so as to resemble medullary rays. They stain of a dark colour, and hence stand
out clearly from the surrounding yellow. The radiating lines are usually just
sufficiently equidistant to allow room between for a radiating row of vessels. ‘The
average leneth of each cell is about 1-2000in.
The vessels, conspicuous by reason of their large cavities, are arranged in radial
rows, and form mainly the inner half of the woody curve. A larger or smaller number
of vessels generally appear on the outer border, at the central and thickest portion of
the wood. The diameter of the vessels varies considerably, the largest being about
1-750 in., and the average diameter about half that. In a longitudinal section the
vessels are seen to be spiral and annular.
From the foregoing brief description of the parts of the F. V. B. in the petiole
of E. globulus, it will be seen that the bundle here is different from the normal
bundle. Usually the phloem and xylem follow each other in the same radius, the
phloem being outside and the xylem inside, so that when a bundle is traced into the
leaf stalk, the phloem portion is towards the outer or lower surface, and the xylem
portion towards the inner or upper surface. In this instance, however, there is
phloem not only on the outer but also on the inner side of the wood, and continued
round, so that the wood is in the centre completely surrounded by the phloem. In
most cases the phloem forms a narrow band, where it curves round to become
continuous. Such a bundle, with the xylem completely surrounded by the phloem, is
called concentric. With the normal arrangement of phloem and xylem, the form of
the bundle is collateral, and when there is phloem outside and inside the xylem it is
distinguished as bicollateral. According to De Bary,* the latter is the form of the
bundle in all investigated species of Eucalyptus, and ‘“‘ Eucalyptus globulus decidedly
belongs to this series.”” As far as the petiole is concerned in this, and we may add,
in the other species investigated, the bundle belongs to the concentric type.
Having seen the mode of arrangement of the parts of the F. V. B., we will now
endeavour to explain the use of this arrangement and structure.
The hard bast all round is evidently, from its very nature and position, a
mechanical support and protection.
The wood fibres in the centre likewise play the part of mechanical support, and
the soft bast is safely placed between the hard bast and the inflexible wood.
The cambiform cells of the soft bast have thin walls, to allow of the circulation of
diffusible substances, but as the thin walls might collapse and thick walls would pre-
* Comp. Anat. of Phanerogams and Ferns, p. 338.
16 THE TRANSVERSE SECTIONS OF PETIOLES OF EUCALYPTS
vent diffusion, a compromise has to be effected. The walls are thickened here and
there, and diffusion occurs in the thin places.
The sieve tubes, with their sieve plates, allow of the transport of indiffusible
substances, such as albuminoids. Just as the blood in our blood vessels is carried
bodily along, so is there a direct bodily transport of these indiffusible substances
through the meshes of the sieve.
The next step is to vessels proper in the wood tissue, where there is no longer
even a sieve plate, but the transverse partitions are removed completely. In that
case the walls want special strengthening to prevent collapse, and so there are all the
admirable contrivances of annular and spiral vessels. The vessels technically belong
to the wood, but they are circulating tissue. They are for quick transport, the
quickest of all; the cambiform cells the slowest, working by the slow process of
diffusion; and the sieve tubes intermediate. The fibrous cells, or wood fibres,
especially the younger, have probably the task of conveying water, for they readily
take it in and readily give it off, so that the I. V. B., with its varied cells, resolves
itself into a contrivance for mechanical support on the one hand, and a means of
circulation on the other.
To show the importance of the soft bast, it may finally be mentioned that it is
probably there albuminoids are manufactured, as well as conveyed to their destina-
tion. It contains more albuminoid matter than other parts, and it is thickly
bestrewn with crystals of oxalate of lime. The albuminoid matter thus produced is
not living protoplasm, but it is a necessary preliminary to it. The exact composition
of this raw material of life occurring in the soft bast is not known. It is quite
possible, however, that any living cell favourably situated may manufacture living
protoplasm when the proper material is supplied to it, for the final touch is the
touch of vitality to convert the lifeless mass into the living protoplasm.
Before glancing at the other transverse sections in a comparative way, the
effects of the application of Schulze’s solution on the different constituents of the
transverse section of EK. globulus may be briefly summarised. The cuticle of the
epidermis is generally stained yellow and orange; the cuticularised layer of a ruby-
red ; the cortical portion purple; the lignified cells of the hard bast, orange; the
soft bast, purple; and the woody tissue, orange, with a tint of brown. ‘The contents
of the cortical cavities seem to be unaffected, and the crystals are dissolved.
In the other transverse sections, the epidermis is generally a well-defined layer of
cells, with more or less thickened outer walls. In some this thickness is excessive,
specially in E. tetraptera, E. obcordata, E. pachypoda, and E. uncinata. It is worthy
of remark that these four are shrubs, or at most small trees, that the two former are
confined to Western Australia, while the two latter belong to desert country, and
constitute a portion of the Mallee scrub.
There is a kino-like exudation often met with in some, such as /eucoxylon,
lehmanni, occidentalis, and alpina, being quite a feature in the last form. The
AS AIDS IN THE DETERMINATION OF SPECIES. 17
hypodermal layer, noted in globulus, is decidedly absent in many, but indications of
it in some, suchas grossa. In /icifolia there is a decided and well-marked hypoderma,
usually consisting of fwo layers of cells. The cortex may be compared as to the
number and size of cortical cavities in it. This is shown in Table I., so that it is
unnecessary to dwell further upon it. In one case at least (E. tetraptera), the
cortical cells are continued into the organic centre of the section by means of two
narrow and adjoining lines of cells.
The hard bast is more or less dense in different sections, forming usually an
interrupted or broken ring.
The soft bast, on the other hand, usually surrounds the wood completely, but it
is sometimes discontinuous.
The wood-curve may be continuous or discontinuous, with a thinner or thicker
body, and longer or shorter horns, but, on the whole, so varied in character as to
serve in many cases as one of the characteristic features of the section.
Broken curves occur in all the species—in some they are the rule, as in /ehmanni,
in many they are exceptional, as in globulus, which was the most constant in its
features of all the types investigated.
This will be the most appropriate place to compare the essential features in the
transverse section of a young stem and a young leaf of E. globulus. The leaf being
sessile, it was necessary to take the base of the midrib instead (Figs. 8 and 9).
Youne Stem. Youne Lear. Oxp Lear.
Base of Midrib. Petiole.
Epidermis— 1 1 1
Thickness. . — = eit
1500 in. 1150 in. 570 in.
Cortical Cavities—
Number 14 at least 4 7 or 8
Shape round round round
i 1 1
Size (largest) — — ae
140 in. 210 in. 80 in.
Hard Bast— Dense and continuous on out- | Very sparingly developed in both | Well developed and dense.
side of F.V.B. In patches in- layers.
side, and absent from corners.
Soft Bast— Continuous outside and inside | Continuous round wood and well Continuous round wood.
of wood, but thinnest at the | developed, particularly on inside
corners. of crescent.
Wood— Forming a square, with slight | Crescentic, with horns well | Crescentic, with horns curving
inward depression on each side curved in. inward.
of square.
Vessels— Distributed mostly towards inner | Numerous, and distributed| Numerous, and distributed
surface of wood, and moderately | towards inner part of wood. | mostly in upper half of wood.
numerous. Largest towards ex-
terior of wood.
1 1 1
Size (widest) = —— —
830 in., an elongated oval. 830 in., elongated ellipse 700 in.
1 1 1
Average = about—— about ——
2300 in. 2300 in. 1400 in.
18 THE TRANSVERSE SECTIONS OF PETIOLES OF EUCALYPTS
The above table shows a contrast in various respects between the young square
stem and the midrib of the young sessvle leaf.
In the stem—Ist. The epidermis is thinner.
2nd. The number and size of cortical cavities is greater.
3rd. The hard bast is much denser.
4th. The wood forms a square, and not a crescent.
The width of the vessels is about the same in both. The upright growing stem
and the lateral growing leaf are necessarily unequally acted on by various forces,
hence the symmetrical and stronger F. V. B. of the stem, as compared with that of
the leaf.
If the comparison is made between the young and the o/d leaf, with its twisted
petiole, there are a few striking differences along with general agreement; but it
will be seen that the differences are simply those between a young and tender organ,
and the same arrived at maturity. In the old leaf, the epidermis is about twice as
thick, the cortical cavities are increased and immensely enlarged, the hard bast is
now strongly developed, and the vessels are a trifle larger.
The twisting of the petiole seems to have produced no noticeable difference in the
general arrangement of the tissues, particularly the vascular. The twisting
of a leaf-stalk round a support, in the case of leaf-climbers, has been shown by
Darwin to increase the growth. The pressure here has acted as a stimulus, but it is
worthy of note that while in the free petiole the bundles form an open curve, in the
clasping petiole they form a closed cylinder.
The mere twisting of the petiole in Eucalypts has not produced any curving of
the bundles greater than that met with in the young and sessile leaf, and even in
those cases where the horns of the crescent most closely approach, as in ficifolia and
calophylla, the leaf-stalk is less twisted than in ordinary cases.
Further, the statement made by Mr. Darwin, on the authority of Dr. Masters,
that ‘‘ the semi-lunar band of vessels is confined to petioles channelled along their
upper surface,” is not countenanced by these researches, for it is in the most
channelled leaf-stalks that the wood curve most approaches the complete ring.
’* Darwin, ‘‘ On the Movements and Habits of Climbing Plants,” p. 43.
AS AIDS IN THE DETERMINATION OF SPECIES. 19
V.— CHARACTERISTICS OF HacH Species AS DERIVED FROM THE TRANSVERSE
SECTIONS.
Baron von Mueller, in his ‘ Eucalyptographia,” as well as in his humbler
‘‘ Botanic Teachings,” has adopted two principal expedients for helping to determine
the numerous species of Eucalypts, viz., the nature of the bark and the nature of
the anthers,
We now add a third, and although it has only been rigorously applied to thirty
different kinds, and to a few more casually, there is every reason to believe that the
principle will hold good throughout that extensive genus of plants. It is based upon
the form and arrangement of the structures revealed in a transverse section of a
well-developed and fully formed leaf stalk, taken in conjunction with external
characters of the leaf.
While the learned Baron has given a leading place in his classification to the
two principles just mentioned, his fertile mind and wide knowledge have suggested a
number of others which he has partially applied. Thus the size of the pollen-grains
has been found to vary in different species, but to be more or less constant in each.
Again, the number and distribution of the stomata have enabled him to form
three series—Ist, according to the presence of stomata on the under surface only
(hypogenous); 2nd, their presence on both surfaces, but less numerous above than
below (heterogenous); or 3rd, their presence on both surfaces, but approximately equal
in number above and below (isogenous). Then again the fruit has been suggested as
a basis of classification, and each of these methods has its own special advantages,
and its suitability for different purposes.
The Carpologic system has this advantage, ‘“‘that any species might thus be
defined from fruiting specimens alone, which latter, through the long persistence of
the fruit, are always obtainable in collecting journeys, whereas flowering specimens
can be got only at some period of the year, subject even to fluctuations and uncer-
tainties.” (Huc. VIII.)
The Cortical system is adapted ‘to the technic requirements of woodmen, who
could not be expected to enter on a discrimination of the various species from such
purely scientific differences, on which descriptive botany would rely.” (Huc. I.) Of
course this system likewise suits the general seeker after a knowledge of these plants.
20 THE TRANSVERSE SECTIONS OF PETIOLES OF EUCALYPTS
The Anthereal system ‘‘ has proved the most convenient for easy-working with
museum material, so long as it was the main object to ascertain the name of any
species.” (Kuc. I. ‘“ Introduction.”)
There is another system recently propounded which ought to be noticed here,
viz., the ‘‘ Kino system ”’ of Maiden.* It is a chemical check, which he merely offers
as a supplement to other systems.
The three groups at present recognised are :-—
1. Ruby group, with ruby-coloured kinos, soluble either in cold water or spirit ;
E. macrorrhyncha, obliqua, and amygdalina, investigated by us, belong to this group.
2. Gummy group, containing gum, and therefore imperfectly soluble in spirit,
but soluble in cold water. £. leucoxylon and saligna belong to this.
3. Turbid group, soluble in hot water or alcohol, but the solution becomes turbid
on cooling.
With regard to the present system, we consider that it has many advantages and
few disadvantages. The Eucalypts being evergreens their leaves are readily obtainable,
and where the leaf happens to be sessile the section of the base of the midrib may
serve the purpose. The resemblance in habit of many different Kucalypts is often
misleading and always confusing, and the basing of characters upon the internal
anatomy, which is not so subject to fluctuation as the external, is an advantage.
Again, the most persistent part of the petiole—the wood—1is often sufficient for
purposes of discrimination, and it may be used, therefore, for the determination of
museum material.
It has, moreover, this great advantage, that it enables us in doubtful and difficult
cases to decide whether the balance of evidence is in favour of specific identity or
merely varietal distinction.
It may be an objection that the above are not naked-eye characters, and that the
system is suited rather to the laboratory than the field, but while less minute
characters may be used for rough discrimination, the final test for specific distinction
must be based upon essential characters that do not always lie upon the surface.
Moreover, the internal structure has been hitherto too much neglected by systematists,
just as anatomists have in like manner paid too little attention to the characters
recognised by the former. If the whole truth is to be expressed by our classification,
then all the features, external and internal, must be taken into account.
* Pharm. Jour. XX., p. 221.
AS AIDS IN THE DETERMINATION OF SPECIES. 21
The accompanying drawings almost render description unnecessary, but as this
is the main portion of the paper, the determining characters of each form examined
will be briefly given.
Before doing so, however, it will be necessary to fix upon some definite arrange-
ment, and a geographical one may be followed, in order to see how far anatomical and
ceographical relations agree.
Table I. exhibits the thirty different forms investigated, variously arranged as far
as the data will permit. It is hardly needful to state that the necessary information
has been obtained from the ‘‘ Eucalyptographia,”’ and we have simply thrown it into
tabular form. The only exception is the arrangement according to cortical cavities,
central canals, and wood curve, which has been deduced from our own researches.
Table II. starts with the forms arranged according to their geographical distri-
bution, and places opposite each the characters respectively deduced from the
transverse section of petiole, the leaves, stomates, fruit, anthers, pollen, and bark.
The very different affinities shown by these various systems of arrangement
only emphasises the fact that it is not by solitary characteristics affinities are deter-
mined, but by the complex of characters, which go to the making up of an individual
whole.
The transverse sections will likewise show that along with a certain amount of
variability there is sufficient constancy to give them a characteristic appearance for
the different species. The most striking features are: Size and shape of section,
relative thickness of epidermis, size and number of cortical cavities, frequency or
scarcity of crystals, density and continuity, or the reverse, of hard bast, pattern of
the wood, and size and distribution of the vessels.
Since all these details are only necessary for complete description, the charac-
teristic features ave given at the end as a sort of summary of characters, serving for
all practical purposes.
As regards the central canals, they occur in only a few of the investigated
species, viz., E. calophylla, ficifolia, and maculata, with its variety citriodora.
Two seems to be the normal number, but since each may divide in a forked
manner, the number is sometimes increased to four, or even six (maculata). (See
remarks in description of each species.)
In stating the characteristic features of the various sections, it is often found
necessary to speak of the relative sizes of different parts, such as the section of the
.
a THE TRANSVERSE SECTIONS OF PETIOLES OF BHUCALYPTS
bo
petiole, the epidermis, cortical cavities, central canals, and vessels, and in using the
expressions “large,” ‘“ small,” or ‘‘ moderate in size,” it all depends upon what is
considered an average size.
While it is not advisable to lay down a hard and fast line, it is well to have a
general understanding of the meaning of the relative terms.
It may be stated generally that the average section is reckoned as being about 1
line X 1 line or 2 line, and above or below that will be large or small respectively.
The size of the section seems to be rather a constant and trustworthy feature.
The absolute size is first given, then the relative breadth and thickness.
The thickness of the epidermis only varies within certain limits, and cannot
often be used as a characteristic, but when it is thicker than about 5 in., or thinner
than ;1. in., it may be regarded as above or below the average. The cortical cavities
1
and central canals may be considered large up to ; in., moderately large up to
1
200
given.
in., and small beyond that. The size of the /argest in each species only is
The vessels have a wide range of variation. In settling the average size it is
largely a matter of judgment, and the largest size given in each case has to be taken
into account. Roughly, when the largest size is ;, Im. or more, and the average not
too low, the vessels might be described as “large,” while at ;3; in., or less, and a
corresponding average, they would be “small,” and the intermediate would be
‘moderately large,” or medium-sized.
Of course the standard of measurement is confined to the petioles of Kucalypts,
and has no reference to the absolute sizes of the parts mentioned in any other
plants.
The size of the cavities and canals varies in different parts of the petiole, and
likewise the width of the vessels, so that too much importance must not be attached
to the actual measurements given. The general results of these measurements may
be given here.
Size of section varies in breadth from } to 1% lines (¢efraptera), and in thickness
from } line (séwartiana) to 13 lines (¢etraptera).
Epidermis varies in thickness from ; in. (viminalis and stuartiana) to x in.
1
oe kas me se :
(tetraptera). A very common thickness is 3; — gp mm.
Cortical cavities vary from in. (diversicolor) to ~ in. (globulus), reckoning only
the largest in each species.
AS AIDS IN THE DETERMINATION OF SPECIES. 23
ponte! canals vary among the four forms possessing them, from 5 in. ({ici-
folia) to + in. (calophylla), taking the largest in each case for comparison.
The vessels varied in width from 4, in. (/ehmanni) toy im. (gomphocephala),
comparing only the largest in each species.
Thus the largest section and thickest epidermis was found in ¢etraptera; the
widest cortical cavity in globulus, although it is closely approached by cornuta ( {; n-),
and the greatest width of vessel in MT gg but globulus, tetraptera, and
citriodora almost equal it, with a width each of 5, in.
The characteristic nature of the wood-curve has already been insisted on, and it
may be added that the extent of development of the hard bast in the different species
seems to be a very constant feature.
I.—Forms ConFINED TO WESTERN AUSTRALIA.
The following descriptions of the various sections are not merely based upon the
type-sections photographed, but are drawn in each case from a considerable number
of independent sections, of which camera lucida drawings were made. The usual
method followed was to make outlines of several sections from the same petiole as
that which supplied the type section, then from different petioles, and lastly from
herbarium specimens, at least three years old, to test the capability of the system for
determining museum material.
1. E. occidentalis, Endlicher ; flat-topped Yate.
' Shrub or tree, rising to 120ft.
Leaves.—Of thick consistence, equally green on both sides.
Size of Section.—1 line broad by ? line thick, or about 1} times broader than
thick.
Outline of Section.—Round to oval.
Epidermis.—Thickness, 4,in.; breadth, ;4, — jy In.
Cortical Cavities.— ;;1n., sieisal numbering from 8 to none, and therefore
variable.
Crystals.—Numerous.
Hard Bast.—Interrupted ring, sometimes only 1 cell thick.
Wood-curve (see Fig. 11).—Body thickened at centre, short horns, and sharply
or gently inturned.
Vessels.—Largest, ;4, im. in dia.
1
> 1500
Comparatively large and numerous, and distributed pretty regularly.
Average in.
2.
3.
THE fRANSVERSE SECTIONS OF PETIOLES OF EUCALYPTS
Characteristic features :
Section.—Average size.
Cortical cavities.—Moderately large, generally few.
Hard Bast.—Interrupted and thin.
Wood-curve.—Body elongated, and short horns sharply inturned.
Vessels.—Moderately large, numerous, and regularly distributed.
E. cornuta, Labillardiére ; the ordinary Yate.
Tree. —Of moderate size.
Leaves.—Of thick consistence, and nearly equally green on both sides.
Size of Section.—# line broad by 3 line thick, or about 13 times broader than
thick.
Outline of Section.—Oval, and somewhat irregular, flattened on upper surface.
‘pidermi icknes 1 yh 2
Kpidermis.—Thickness, 2,— J in.
Breadth, ;*, in.
Cortical Cavities.— 7 in. oval, numbering from 9 to 3.
Crystals.— Moderately numerous.
Hard Bast.—Pretty generally continuous, and several cells thick.
Wood-curves (see Fig. 12).—Body slightly thickened at the centre, and short
horns gently or somewhat sharply incurved.
Vessels.—Largest, >, in.
Average, about half that.
Somewhat large, and most numerous towards upper portion of wood.
Characteristic features :
Section.—Relatively small.
Cortical Cavities.—Large, numerous, often quite close together, and almost
occupying breadth of cortex.
Hard Bast.—Continuous and thick.
Wood-curve.—Horns sharply incurved, and one or both often separated
from body-curves.
Vessels.—Moderately large, and aggregated towards middle and upper sur-
face of wood.
E. cornuta, var lehmann., Preiss.
Tree.—Of moderate size.
Leaves.—Of thick consistence, and equally green on both sides.
Size of Section.—About 1 line broad by 3 line thick, or twice as broad as thick.
Outline of Section.-—An irregular oval, elongated laterally.
Hpidermis.—Thickness, <, — 4) in.
. La
Breadth, 3, in.
4.
AS AIDS IN THE DETERMINATION OF SPECIES. 25
Cortical Cavities.— 4, in., oval to round, numbering from 11 to 3.
Hard Bast.—Generally in isolated contiguous patches, a few or several cells thick.
Wood-curve (see Fig. 13).—Slender elongated body, almost always broken,
and with sharply incurved horns.
Vessels.—Largest, 54, 1n-
Average about , i.
Small, few, and regularly distributed.
Characteristic features :
Size of Section.—Twice as broad as thick.
Cortical Cavities. —Moderately large and numerous.
Hard Bast.—Isolated contiguous patches.
Wood-curve.—Generally broken, with slender body, and sharply imceurved
horns.
Vessels.—Few and small.
E. obcordata, Turezaninow ; “‘ Maalok.”
Tall shrub or small tree.
Leaves.—Thick, broad, blunt, equally green on both sides.
Size of Section._—About 1 line broad and thick, or as broad as thick.
Outline of Section.—Round to oval, the greatest diameter sometimes dorso-ventral.
Epidermis.— Thickness, 3 — go 19-
Breadth, =, — wm: or half the thickness.
Cortical Cavities. — 4, in. roundish, numbering from 10 to 6.
Crystals.— Numerous.
Hard Bast. —Very feebly developed, often consisting merely of isolated cells at
distant intervals.
Wood-curve (see Fig. 14),—With somewhat thickened body, graceful incurving,
and moderate-sized horns.
Vessels.—Largest, 4, in.
Average, about half that.
Small, not very numerous, in radiating lines, and mostly towards upper
portion of wood.
Characteristic features :
Section.—Average size, as broad as thick.
Cortical Cavities. —Large, and comparatively numerous.
Hard Bast.—Feebly developed.
Wood-curve.—With thickened body and graceful incurving horns.
Vessels.—Comparatively small, not very numerous, and distributed mostly
towards upper portion of wood.
26
5.
6.
THE TRANSVERSE SECTIONS OF PETIOLES OF EUCALYPTS
E. marginata, Donn ; Jarrah.
Tree, averaging 100ft. in height.
Leaves.—Somewhat paler beneath.
Size of Section.—About 1 line broad by 3 line thick, or fully twice as broad as
thick.
Outline of Section. Laterally elongated, ovoid usually.
Epidermis.—Thickness, 7 — 7 in.
Breadth, 5,— im.
Cortical Cavities. —-f, in., almost round, numbering from 9 to 3.
Crystals.—Numerous.
Hard Bast.—More or less continuous, and generally one or a few cells thick.
Wood-curve (see Fig. 15).—Elongated laterally and slender, sometimes slightly
thickened about the middle, incurving somewhat sharply, and horns
usually relatively short.
Vessels.—Largest, ,4, in.
Average, about half that.
Medium-sized, moderately numerous, and distributed pretty regularly.
Characteristic features :
Section.—Fully twice as broad as thick.
Cortical Cavities.—Moderately large and numerous.
Hard Bast.—Continuous ; a few cells thick.
Wood-curve.—With slender, elongated body, and usually short slender
horns.
Vessels.—Medium-sized, moderately numerous, and regularly distributed.
E. diversicolor, F. v. M.; Karri.
Tree.—Very tall and straight.
Leaves.—Elongated, much paler beneath.
Size of Section.—About # line broad by 4 line thick, or about 14 times broader
than thick.
Outline of Section.—Oval to round, usually flattened on upper surface.
Epidermis.—Thickness, ¢, in.
Breadth, about half the thickness.
Cortical Cavities.— 4, in., roundish, numbering from 7 to 2, or sometimes none.
Crystals.—Few.
Hard Bast.—Poorly developed, very discontinuous, often consisting merely of
single isolated cells.
Wood-curve (see Fig. 16).—Body may be either short and stout, or elongated
and slender, with gentle incurving and somewhat stoutish horns.
AS AIDS IN THE DETERMINATION OF SPECIES. 27
Vessels.— Largest, 5 I.
Average, 3, i.
Comparatively small, not very numerous, generally distributed in
radiating lines, absent from lower portion of wood.
Characteristic features:
Section.—Comparatively small.
Cortical Cavities.— Small, and often few.
Hard Bast.—Poorly developed:
Vessels.—Medium-sized, not numerous, distributed in upper portion of
wood.
7. E. gomphocephala, De Candolle ; Tooart.
8.
A good-sized tree.
Leaves.—Thick, shining, slightly paler beneath.
Size of Section—About # line broad and thick, or of equal breadth and
thickness.
Outline of Section.—From round to oval.
Epidermis.—Thickness, 7 — 7 1.
Breadth, .4, in.
Cortical Cavities.—, in., elongated oval, numbering from 4 to none.
Crystals.—Very numerous.
Hard Bast. Forming a pretty continuous ring, generally several cells thick.
Wood-curve (see Fig. 17).—Body usually exceedingly thick, with large and
stout horns, often well curved inward and approaching each other.
Vessels.—Largest, 4, in. and several ;;; In.
Average, o5 — jy In.
Large and comparatively numerous, distributed mostly towards
middle and upper surface.
Characteristic features :
Section.—Of equal breadth and thickness.
Cortical Cavities.—Moderately large, but few.
Hard Bast.—Continuous and thick.
Wood-curve.—With exceedingly thickened body, and large and stout horns.
Vessels.—Large, numerous, and distributed largely in upper half of wood.
E. megacarpa, F. v. M.; West Australian Blue Gum.
Tree.—Medium-sized.
Leaves.—Of pleasant odour, and equally green on both sides.
Size of Section.—1 line broad by } line thick, or twice as broad as thick.
28
Sh
THE TRANSVERSE SECTIONS OF PETIOLES OF EUCALYPTS
Outline of Section Generally oval and laterally expanded, sometimes roundish
or angular, flattened on upper and under surface.
Epidermis.—Thickness, 4, — 3 1
Breadth, 4, — ym in., or half the thickness.
Cortical Cavities.—#, in. round, numbering from 7 to 2.
in.
Crystals.—Very few.
Hard Bast.—Generally continuous, but irregular in thickness, giving it a zig-zag
appearance.
Wood-curve (see Fig. 18).—Elongated, thickened at centre, with gentle
inturning, and horns of medium length.
Vessels.—Largest, 3
Average, ae half that.
Moderately numerous, wide variations in size, lower portion of wood
comparatively free from them.
Characteristic features :
Section.—Flattened, about twice as broad as thick, tending to angularity.
Cortical Cavities.— Moderately large.
Hard Bast.—Generally continuous, but irregular in thickness.
Wood-curve.— Slender, slightly thickened in centre, and short horns gently
incurved.
Vessels.—Moderately large and numerous.
E. tetraptera, Turczaninow ; four-winged-fruited Eucalypt.
Shrub, seldom exceeding 10ft. in height.
Leaves.—Very thick and equally green on both sides.
Size of Section.—1# line broad by 13 thick, or nearly 14 broader than thick.
Outline of Section.—Circular to oval.
ee eg ee wo in.
Pred um
Cortical Cavities.— j, in. round to oval, numbering from 4 to 2.
in. or about 4 thickness.
Crystals.—Numerous.
Hard Bast.—Continuous except where wood-curve is broken, and usually several
cells thick.
Wood-curve (see Fig. 19).— Body thickened in middle, and gently incurved horns
of medium size.
Vessels.—Largest
Average
: rn in. and several ., in.
Ae
’ 300 nee
Large, numerous, and distributed pretty regularly.
AS AIDS IN THE DETERMINATION OF SPECIES. 29
Characteristic features :
Section.—Very large.
Epidermis. —Very thick.
Cortical Cavities.—Moderately large and few.
Hard Bast.—Continuous, and usually several cells thick.
Vessels.—Large, numerous, and regularly distributed.
10. E. rudis, Endlicher ; flooded or swamp Gum of Western Australia.
Tree.—Usually not very tall.
Leaves.—Generally thin, and equally green on both sides.
Size of Section.—About $line broad by 4 line thick, or 14 times broader than
thick.
Outline of Section.—Roundish, and flattened on top.
EPS aay Tckneas, zp mM.
Br Sea am in., or about half the thickness.
Cortical Cavities.— ~ in., roundish, numbering from 9 to 2.
Crystals.—Few.
Hard Bast.—Very poorly developed, often just represented by straggling cells.
Wood-curve (see Fig. 20), —Illongated, and with short or medium-sized horns.
Vessels.— Largest, 0 in.
Average, ay, In.
Comparatively small, moderately numerous, and mostly distributed about
middle.
Characteristic features :
Section.—Small.
Cortical Cavities.—Small.
Hard Bast.—Very poorly developed.
Wood-curve.—With body elongated and short horns.
Vessels.—Small, and mainly distributed in middle of wood.
11. E. pachypoda, F.v. M.
Shrub.
Leaves.—Equally green and pale on both sides.
Size of Section.—-About 1 lie broad and thick, or equally broad and thick.
Outline of Section.—Round, sometimes slightly elongated.
Epidermis.—Thickness, ¢,in.; oval, numbering from 4 to 2.
ue about 4, in.
Cortical Cavities.—i, in.
Crystals.—Few.
as
30 THE TRANSVERSE SECTIONS OF PETIOLES OF BUCALYPTS
Hard Bast.-—Ivregularly developed, discontinuous, many-celled, or only 1 cell thick.
Wood-curve (see Fig. 21).—Body thickened, with the two short horns gently
incurved, and approaching each other.
Vessels.— Largest, 5 1
Average, pe half that.
Medium-sized, regularly distributed, but not crowded.
Characteristic features :
Section.—Average size, equally broad and thick.
EKpidermis. — Relatively thick.
Cortical Cavities.—Moderately large, but few.
Hard Bast.—Weak and irregularly developed.
Vessels.—Medium-sized, and very regularly distributed.
12. E. calophylla, Brown; Calophyllum-hke Eucalypt.
Tall Tree.
Leaves.—Broad, acute, thick, more horizontal than vertical, and much paler
beneath.
Size of Section.—13 lines broad by 1} lines thick, or nearly 14 times broader
than thick.
Outline of Section.—Oval to round, and usually flattened on Sn surface.
Epidermis.—Thickness, variable in same section from J, in. to = in
Cortical Cavities.—Largest, ¢, in., round or oval, numbering from \4 to 3.
Central Canals.—Normally two, oval, one4 in., other J in., or sometimes half that.
Crystals.—Moderately numerous.
Hard Bast.—Very sparingly developed, only in small isolated patches.
Wood-curve (see Figs. 22 and 22a).—Generally somewhat slender, and of nearly
equal thickness throughout; the horns often approaching each other.
Vessels.—Largest, 2 a) in.
Average n.
’ mn i
Comparatively large and numerous, and distributed in radiating lines
mostly towards upper portion.
Characteristic features :
Section.—Large.
Cortical Cavities.—Generally large and numerous.
Central Canals.—Usually two ; very large, relatively to the others.
Hard Bast.—Very sparingly developed.
Wood-curve.—With moderately-thickened body and horns often
approaching.
Vessels.—Large, numerous, and distributed mostly towards upper portion
of wood.
AS AIDS IN THE DETERMINATION OF SPECIES. 31
Remarxs.—While two is the normal number of central canals, there may be
only one, or as many as four.
The sections from one petiole showed only a single canal near the base, but the
normal number (2) near the leaf-blade. The two main canals were also found in
several instances to divide into four on passing into the midrib. Probably this is a
normal occurrence in this species.
The contents of the canals were tested with various reagents. When treated
perfectly fresh they readily dissolved, but after long steeping in water, they became
hardened and insoluble in the ordinary reagents. Under these circumstances they
would not dissolve either when warmed in nitric acid or in potash and glycerine.
Nitric acid gave a red tint, and potash a brown, but that was all the change observed.
In the section of the young stem it is interesting to observe that central canals
are absent, while the cortical cavities are present. The latter are very numerous, just
as in some sections of the leaf stalk. The hard bast is not of equal thickness
throughout, nor is it always continuous, although it is shown thus in the drawing for
simplicity. Only the larger vessels of the wood are figured.
13. E. ficifolia, &. v. M.; scarlet-flowered Eucalypt.
Tree.—Seldom exceeding height of 50ft.
Leaves.— Generally resembling preceding, but not so broad in proportion to
length.
Size of Section.— 14 lines broad by 1 line thick, or 1} times broader than thick.
Outline of Section. Varying from round to oval to a much depressed semi-circle.
Epidermis.—Thickness, 5, — J in.
Breadth, 4 in
Hypoderma.—Equal to enderms in thickness, and usually a two-celled layer.
Cortical Cavities.—About 3, in., few in number, or absent.
Central Canals.— Normally 2, ‘a occasionally 3 or 4; roundish.
Largest, about 5, in.
Crystals.—Numerous.
Hard Bast.—In irregular patches, a few cells thick.
Wood-curve (see Figs. 23 and 23a).—Generally slender, and with horns curving
gently, and sometimes almost meeting.
Vessels.—Largest, 3 in
Average, abouts: mw}
Comparatively a Ae numerous, and distributed towards upper
and middle portions of wood.
32 THE TRANSVERSE SECTIONS OF PETIOLES OF EUCALYPTS
Characteristic features :
Section.—Large.
Hypoderma.—Generally two cells thick.
Cortical Cavities.—Small and few.
Central Canals.—Normally two, but liable to variation; about same size
as the largest of the cortical cavities.
Hard Bast.—In irregular patches.
Wood-curve.—Slender, and horns approaching each other, sometimes very
closely.
Vessels.—Comparatively large, numerous, absent from lower portion of
wood.
Remarxs.—Cortical cavities are certainly present in this species, but they are
few in number, and many sections of a series show none. ‘They are likewise small,
indeed the smallest of any species examined.
The normal number of central canals is two, but occasionally there may be
three or even four, as in one of the sections photographed. The occurrence of four
canals is rather interesting in view of the fact that they were found in sections taken
near the base of the leaf-blade. It would appear that while the normal number of
two usually occurs in the body of the petiole, as the lamina is approached each
divides into two before entering the mid-rib.
Il.—Form ConFINED To VICTORIA.
14. E. alpina, Lindley; alpine gum.
Shrub.
Leaves.—Very thick, broad, blunt, equally green on both sides.
Size of Section.—About 1} lines broad and thick, or as broad as thick.
Outline of Section.—Usually rounded,
Epidermis.—Thickness, J, in. below, 7 in. above.
Cortical Cavities. —4, in. roundish, numbering from 16 to 6.
Crystals.—Very few.
Hard Bast.—Well-developed, generally continuous, and several cells thick.
Wood-curve (see Fig. 24).—With thickened body and gently incurved short horns.
Vessels.—Largest, 4 in.
Average, za in.
Medium-sized and moderately numerous, principally occupying middle
tract of wood.
AS AIDS IN THE DETERMINATION OF SPECIES. 30
Characteristic features :
Section.—Large, as broad as thick.
Cortical Cavities. —Moderately large, and numerous.
Hard Bast.—Well-developed, and generally continuous.
Wood-curve.—With thickened body and gently incurved short horns.
Vessels.--Medium-sized, principally distributed in middle tract of wood.
Tll.—Form Conrinep to New SoutH WaAtgs.
15. E. punctata, De Candolle ; leather-jacket.
Tree.—Attaining a height of 100ft. or more.
Leaves.—Paler beneath.
Size of Section.—About } line broad and thick, or as broad as thick.
Outline of Section.—Round, with slight flattening on upper surface.
Epidermis.—Thickness, 3; — jg 1.
Breadth, about —,, in.
Cortical Cavities.— 4, in., elongated oval, numbering from 8 to 1.
Crystals.—Very few.
Hard Bast.—Very sparsely developed.
Wood-curve (see Fig. 25).—Nearly always found entire; body thickened, and
horns very gently incurved and short.
Vessels.— Largest, 4; in.
Average, gy i.
Relatively few, small and scattered.
Characteristic features :
Section.—Small, as broad as thick.
Cortical Cavities.— Moderately large.
Hard Bast.—Very poorly developed.
Wood-curve.—Nearly always found entire, with thickened body and short
horns.
Vessels.—Few, small, and scattered.
IV.—Forms ConFINED TO QUEENSLAND.
16. E. maculata, var. citriodora, Hooker ; lemon-scented Eucalypt.
Tree.—Only differing from maculata in the lemon-scented foliage.
Size of Section.—1} line broad by 1 line thick, or 1} times broader than thick.
Outline of Section.—Oval, somewhat irregular in outline, and flattened or
depressed on top.
34 THE TRANSVERSE SECTIONS OF PETIOLES OF EUCALYPTS
Epidermis.—Thickness, 5, — gp I.
Breadth, ;4, 1.
Cortical Cavities.— Largest, +, in., oval to round, numbering from 8 to 0.
Central Canals.—Normally 2, oval and round; largest about the same size as
cortical cavity.
Crystals.—Moderately numerous.
Hard Bast.—Very poorly developed, just represented at intervals by a few
straggling cells.
Wood-curve (see Fig. 26).—Usually of equal thickness throughout and slender,
with very gently incurved horns, each about one-third body-length.
Vessels.—Largest, =, in.
Average, about 5, im.
Numerous, large (about twice the size of those of maculata), and
generally distributed.
Characteristic features :
Section.—Large.
Cortical Cavities.—Moderately large, variable in number.
Central Canals.— Normally two, and same size as cortical cavities.
Hard Bast.—Very poorly developed.
Wood-curve.—Of equal thickness throughout, with very gently incurved
horns.
Vessels.— Large, numerous, and generally distributed.
Remarxs.—A petiole was noticed showing only one central canal near the base
or point of attachment to the stem, but the normal number (two) towards the leaf-
blade. Another petiole showed one only in all the sections examined.
V.—Form Bertonerna to Soutn AUSTRALIA AND VICTORIA.
17. E. corynocalyx, F. v. M.; sugary Eucalypt; sugar gum.
Tree.—Finally tall, said to be the largest tree growing in South Australia.
Leaves.—Shining, somewhat paler beneath.
Size of Section.—About # line broad by } line thick, or 1} times broader than
thick.
Outline of Section.—Round to oval, usually flattened on upper surface.
Epidermis.—Thickness, 5 in.
Breadth, ;4, in.
Cortical Cavities.-- 4, in., round or slightly oval, numbering from 4 to 18.
AS AIDS IN THE DETERMINATION OF SPECIES. 35
Crystals.—Few.
Hard Bast.— Sparingly developed, sometimes consisting of isolated cells.
Wood-curve (see Fig. 27).— Usually with slender, elevated horns, dipping inward.
Vessels.---Largest, 5 in.
Average, about 4, in.
Rather small on the whole, comparatively few, somewhat scattered,
and mostly absent from lower portion of wood.
Characteristic features :
Section.— Small.
Cortical Cavities —Numerous, but comparatively small.
Hard Bast.—Sparingly developed.
Wood-curve.— Usually symmetrical and entire, with slender and very gently
incurved horns.
Vessels.—Comparatively small.
VI.—Forms Brtoneine to Victoria, New SoutH WALEs, AND QUEENSLAND.
18. E. melliodora, Cunningham ; honey-scented Eucalypt, or yellow box tree.
Tree.—Middle-sized.
Leaves.—Equally dull-green on both sides.
Size of Section.—About 4 line broad and thick, or as broad as thick.
Outline of Section.—Round or oval, and flattened on upper surface.
Epidermis.—Thickness, 5 — a In.
Breadth, about ;4, in.
Cortical Cavities.—3, in., round, numbering from 4 to 1 or none.
Crystals.—Very numerous, and frequently in groups.
Hard Bast.—Continuous, and generally two cells thick.
Wood-curve (see Fig. 28).—Always found unbroken, with thickened body and
sharply inturned short horns.
Vessels.—Largest, jj, in.
Average, about =, in.
Medium-sized, numerous, and equally distributed.
Characteristic features :
Section.—Small; as broad as thick.
Cortical Cavities.—Small and few.
Hard Bast. - Continuous, and generally a two-celled layer.
Wood-curve.—Always found unbroken, with thickened body and short horns.
Vessels. —Medium-sized, numerous, and equally distributed.
36
THE TRANSVERSE SECTIONS OF PETIOLES OF EUCALYPTS
19. E. tereticornis, Smith; flooded gum-tree.
Tree,—Tall, handsome, seldom exceeding 100ft. in height.
Leaves.—Kqually green on both sides.
Size of Section.—About ? line broad by 3 line thick, or 14 times broader than
thick.
Outline of Section. eee aes round to oval.
EKpidermis.—Thickness, ~, in.
> 760
ee
Breadth, 5, — yj im.
Cortical Cavities. —*, in., round to oval, numbering from 10 to none, but on an
average ‘belat 5.
Crystals.—Very numerous.
Hard Bast.—Sparinely developed, discontinuous, in many places 1 cell thick.
Wood-curve (see Fig. 29).—Body moderately thickened and horns gently
incurved ; es found entire.
Vessels—Largest, 4, 1
Average, a half that.
Comparatively small, moderately numerous, distributed loosely towards
upper portion, and generally absent from lower portion of wood.
Characteristic features :
Section.— Small.
Cortical Cavities. —Small.
Hard Bast.— Sparingly developed.
Wood-curve.—Symmetrical, and always found entire.
Vessels.—Comparatively small, moderately numerous, and distributed
loosely towards upper portion of wood.
VII.—Forms Benoneine to New South Wates anp QUEENSLAND.
20. E. saligna, Smith; grey gum-tree.
‘Tree.—Tall.
Leaves.—Tapering, much paler beneath.
Size of Section.—% line broad by 4 line thick, or 14 times as broad as thick.
Outline of Section.— Rounded, with flattened upper surface.
Epidermis.—Thickness, 4, m.
Breadth, about 5 1
Cortical Cavities.— < in., round to eae numbering from 7 to 6.
Crystals.—Numerous.
Hard Bast.—Very poorly developed, and in many places entirely absent.
AS AIDS IN THE DETERMINATION OF SPECIES. : 37
Wood-curve (see Fig. 30).—Body thickened and elevated, horns generally not
much incurved.
Vessels.—Largest, 5, in.
Average, about half that.
Moderately large, numerous, and generally distributed.
Characteristic features :
Section.—Small.
Cortical Cavities —Moderately large and numerous.
Hard Bast.—Feebly developed.
Wood-curve.-—With thickened body, and generally elevated horns.
Vessels.—Moderately large, numerous, and generally distributed.
21. E. maculata, Hooker; spotted gum.
Tree.—Tall, handsome.
Leaves. —Often very large and coarse, and equally green on both sides.
Size of Section.—1 line broad by ? line thick, or about 1} times broader than
thick.
Outline of section.—Oval, flattened on upper surface.
Epidermis.—Thickness, about ,5, in.
Breadth, about half that.
Cortical Cavities.—,5 in. oval to round, numbering from 6 to 2.
Central Canals.—Normally 2, oval to round; largest, about 4, in., or generally
the size of the cortical cavities.
Crystals.---Numerous.
Hard Bast.—In closely adjoining patches, generally a few cells thick.
Wood-curve (see Fig.31).—Usually of equal thickness throughout, with very
gently incurved horns, each about one-third of body-length.
Vessels.—Largest, about 7, in.
Average, 4, — gr im.
Moderately large and numerous, confined almost exclusively to upper
and middle portion of wood.
Characteristic features:
Section.—Of average size.
Cortical Cavities.—Medium-sized, not very numerous.
Central Canals.—Normally 2, about same size as cortical canals.
Hard Bast.—In contiguous patches, a few cells thick.
Wood-curve.—Of equal thickness throughout, with very gently incurved
horns.
Vessels.—Moderate in size and number, and absent from lower portion of
wood.
38 . THE TRANSVERSE SECTIONS OF PETIOLES OF EUCALYPTS
Remarxs.—The number of central canals varies occasionally, due probably to the
splitting up of the normal number (two). Thus, in nine successive sections taken from
about the middle of the leaf stalk, there were four, two on each side; and in another
petiole, the sections showed five, three on one side and two on another. And, as if
to complete the series, a third petiole exhibited no less than six canals—three on
either side, one of each being about the normal size, and the other two rather
smaller.
VIII.—Form Betoneine to Sours Austrauia, Victoria, anD New Sout WALEs.
22. E. macrorrhyncha, F. v. M.; Victorian stringy-bark.
Tree.— Tall.
Leaves.—EHlongated, equally green on both sides.
Size of Section. —1 line broad by % line thick, or about 14 times broader than
thick.
Outline of Section.—Oval, or nearly round.
Epidermis.— Thickness, 5, in.
Breadth, 7, — gq in.
Cortical Cavities. — in., oval and round, numbering from 8 to 2.
Crystals.—Few.
Hard Bast.—Well-defined layer, several cells thick.
Wood-curve (see Fig. 32).—Body exceedingly thick and short; horns stout,
upright, or sharply inturned ; always found entire.
Vessels.— Largest
ae
> qqg WD.
Average
tele
Large and numerous; pretty equally distributed, but most thickly
towards upper portion of wood.
Characteristic features :
Section.—Average size.
Cortical Cavities.— Moderately large.
Hard Bast.—Well-defined, and several cells thick.
Wood-curve.—Always found unbroken, with much-thickened body, and
short stout horns.
Vessels.—Large, numerous, distributed mostly towards upper portion of
wood.
AS AIDS IN THE DETERMINATION OF SPECIES. 39
IX.—Form Bertonerne to Tasmania, Victorta, AND New SoutH WALES.
23. E. globulus, Labillardiere; blue gum.
Tree.—Very tall.
Leaves.—Thick, elongated, equally green on both sides.
Size of Section.—A little over 1 line broad and 1 line thick, or slightly broader
than thick.
Outline of Section.—Oval, slightly flattened on upper surface.
Epidermis.—Thickness, + in.
Breadth, 5,— gq in.
Cortical Cavities.— in., oval to round, numbering usually 7 or 8.
Crystals.— Very numerous.
Hard Bast.—Strongly developed, continuous, and usually dense.
Wood-curve (see Figs. 32 and 33a).—Body thickened, particularly at the centre,
and gently incurved horns, each about one-third length of body,
and directed towards it at the tip.
Vessels.—Largest, 4, in.
Average, about half that.
Comparatively large and numerous, radially arranged, and distributed
mainly in upper half. A number of large vessels usually scattered
about lower portions of wood.
Characteristic features :
Section.—Average size, slightly broader than thick.
Cortical Cavities—Large, and numbering about 7 or 8.
Crystals.—Very numerous.
Hard Bast.—Well-developed and dense.
Wood-curve.—-With thickened body, and horns curving gently inward.
Vessels.—Large, numerous, and distributed mostly in upper half, but
sparingly along lower margin of wood.
Remarx.—In one petiole examined, all the sections showed an isolated bundle at
one side of the ordinary curve.
This species, however, shows as little tendency to broken and isolated wood-
curve as any other examined.
X.—Formus Betoncine to SoutrH AusTrRALiA, TasMANIA, VICTORIA, AND
New Soutn Watgs.
24. E. obliqua, L’ Heritier ; messmate.
Tree.—Lofty.
Leaves.—Very inequilateral at base, equally green on both sides.
40 THE TRANSVERSE SECTIONS OF PETIOLES OF EUCALYPTS
Size of Section.— 1} lines broad by 1 line thick, or 14 times broader than thick.
Outline of Section.—Oval to circular, sometimes flat on upper surface.
Epidermis.—Thickness, ,-— 7, in.
Breadth, about ;%, in.
Cortical Cavities.—-;, n., roundish, numbering from 7 to 2.
Crystals.—Few.
Hard Bast.—Well developed, continuous, and many cells thick.
Wood-curve (see Fig. 84).—Comparatively slender body, with short horns
elevated or gently incurved.
Vessels.— Largest, 4, In.
Average, about half that.
Large, numerous, and pretty regularly distributed, but most
numerous towards upper portion of wood.
Characteristic features :
Section.—Large.
Epidermis.—Thickish.
Cortical Cavities.—Large.
Hard Bast.—Well developed and dense, especially between the horns.
Wood-curve.—With slender body and short horns.
Vessels.—Large, numerous, and pretty regularly distributed.
25. E. amygdalina, Labillardiére ; giant Eucalypt, Dandenong peppermint, mountain
ash.
Reputed to be the tallest of trees.
Leaves.—Thin, equally green on both sides, with copious and transparent oil-
dots.
Size of Section.—About 4 line broad and thick, or equally broad and thick.
Outline of Section.—Oval to round, with occasionally a flattened upper surface.
Epidermis.—Thickness, = —
=
ag in.
‘ 1 if =
Breadth, 3, — win.
aj I., oval or round, numbering from 7 to 2.
Cortical Cavities.— =,
Crystals.—Numerous.
Hard Bast.—Sparingly developed, one or a few cells thick.
Wood-curve (see Fig. 35).— Body excessively thick, with short stout horns elevated
or slightly incurved; always found entire.
Vessels.—Largest, |4, In.
Average, about half that.
Small, moderately numerous, on radiating lines, generally equally
distributed, but scanty towards lower margin.
AS AIDS IN THE DETERMINATION OF SPECIES. 41
Characteristic features :
Section.—EHxceptionally small, equally broad and thick.
Cortical Cavities. —Small.
Hard Bast.—Composed of few cells, but strongly thickened.
Wood-curve.—Always found entire, with excessively stout body and short,
stout horns.
Vessels.—Small, moderately numerous, and pretty regularly distributed.
Remarks.—In proportion to the size of the section, the wood-curve is the
thickest of any yet investigated. This is of interest in connection with such a very
tall tree, since the water must be raised so great a distance to reach the topmost
boughs. It was also observed that the leaves from this tree did not, in the least,
discolour the water in which they were kept for some months, although all other
species examined did so, more or less.
26. E. viminalis, Labillardiére ; Manna-Kucalypt.
Tree.—Tall, and sometimes even gigantic.
Leaves.—Thin, equally green on both sides.
Size of Section.—Hither 1 line broad by ? line thick, or of equal breadth and
thickness.
Outline of Section.—Round to egg-shaped, or flattened on top, so as to become
somewhat hemispherical.
Kpidermis.— Thickness, 5, in.
Breadth, as broad as thick, and sometimes broader.
Cortical Cavities—,, in. in some sections, and only half that in others;
roundish to oval ; numbering from 7, often 2, and sometimes absent.
Crystals.—Numerous, and sometimes large.
Hard Bast.—Moderately developed; slightly discontinuous; 1 sometimes,
but mostly several cells thick.
Wood-curve (see Figs. 36 and 36a).—Body straight and thickened, with short,
sharply incurved, and usually broken horns. Occasionally a distinct
isolated bundle at side of one of the horns.
a 1 ees
Vessels.— Largest, from ,5, — ji, I.
. We os
Average.—,4, in.
Small, comparatively few or moderately numerous, mostly distributed
towards upper portion of wood, and generally absent from horns.
Characteristic features :
Section.—Average size.
EKpidermis.—Relatively thin, composed of cells at least as broad as thick.
Cortical Cavities.—Medium size, often few.
42,
THE TRANSVERSE SECTIONS OF PETIOLES OF EUCALYPTS
Wood-curve.—Straight and thickened body, with short, sharply incurved
horns, often in detached parts.
Vessels.—Small, and distributed on upper portion of wood, generally absent
from horns.
27. E. gunni, J. Hooker; swamp gum or cider Eucalypt.
Tree.—Usually not tall.
Leaves.—Equally dark green on both sides.
Size of Section.—1} lines broad by 1 line thick, or 1} lines broader than thick.
Outline of Section.—Oval to round, sometimes flattened on top.
Epidermis.—Thickness, 5 — 7 i.
Breadth, about ;*, in.
Cortical Cavities.—,,, in., round or oval, numbering from 9 to 2.
Crystals.—Few.
Hard Bast.—Well developed, nearly continuous, and several layers of cells thick.
Wood-curve (see Fig. 37).—Body thickened (thickness sometimes ;, in.), with
short and sharply incurved horns directed towards body.
Vessels.—Largest, 3, in.
Average, ,2,n., or about half the largest.
Moderately large and numerous, and pretty equally distributed with
large ones at lower portion of wood as in globulus.
Characteristic features :
Section.—Relatively large.
Cortical Cavities.—Large and moderately numerous.
Hard Bast.—Somewhat dense.
Wood-curve.—With thickened body and short horns directed towards it.
Vessels.—Moderately large, numerous, and pretty generally distributed.
XI.—Form Betoneinc to Sourn Avustrauia, Tasmania, Victoria, New SoutH
WALES, AND QUEENSLAND.
28. E. stuartiana, F. v. M.; apple-scented Eucalypt.
Tree.—Moderate-sized.
Leaves.—Kqually dark-green on both sides.
Size of Section.—About 4 line broad by } line thick, or about 14 times broader
than thick.
Outline of Section.—Oval.
AS AIDS IN THE DETERMINATION OF SPECIES. 43
epi eres, Lnlgneees am in.
ee
Cortical Cavities.—
i= — im IN., or sometimes as broad as thick.
= in., oval to oe cis round, numbering about 6.
Crystals.—Moderately numerous.
Hard Bast.—Almost continuous, regular, generally two or three cells thick.
Wood-curve (see Fig. a): —Body curved and moderately thickened (thickness
Samieunpe zy in.), with short and sharply incurved horns.
Vessels.—Largest, = in.
Average, about half that.
Small, numerous, and generally distributed towards upper and middle
portion of wood.
Characteristic features :
Section.—Small.
lpidermis.—Relatively thin, component cells often as broad as thick.
Cortical Cavities.—Small.
Hard Bast.—Regular, but not very dense.
Wood-curve.—Moderately thick, and horns somewhat sharply incurved.
Vessels Small numerous, and distributed towards upper and middle
portion of wood.
XIJ.— Form Betoneine to Sourn Avustrauia, Victorta, New SourH WALES, AND
QUEENSLAND.
29. E. leucoxylon, F. v. M.; Victorian ironbark.
‘Tree.—Moderate size.
Leaves.—Equally dull-green on both sides, with copious, pellucid oil-dots.
Size of Section.—13 lines broad by # line thick, or twice as broad as thick.
Outline of Section.—Roundish, flattened, or somewhat quadrangular.
PL eae ppg aq in.
Breadth =o Vide
’ nS 1500
Cortical Cavities.— ;, in. oval, numbering from 9 to 1, or none.
Crystals.—Moderately numerous.
Hard Bast.—Poorly developed; in many places entirely absent.
Wood-curve (see Figs. 89 and 89a, b, c)—Body variable in thickness, and longer
or shorter horns, gently incurved.
Vessels.—Largest, 4 io in.
Average n.
1
; =
Medium-sized and numerous, radially arranged, and regularly distributed.
44 THE TRANSVERSE SECTIONS OF PETIOLES OF EUCALYPTS
Characteristic features :
Section.—Large, twice as broad as thick.
Cortical Cavities.—Large.
Hard Bast.—Scantily developed.
Vessels.—Medium-sized, numerous, and regularly distributed.
XIII.—Form Betoneinc To Western Australia, SoutH AUSTRALIA, VICTORIA,
New South Wass, QuEENSLAND, AND NortH AUSTRALIA.
30. EH. rostrata, Schlechtendal ; red gum.
Tree.—Of moderate size, often attaining a height of over 100ft.
Leaves.—Kqually green on both sides.
Size of Section.—About 4 line broad and thick, or equally broad as thick.
Outline of Section.—From round to irregular oval, generally with somewhat
flattened upper surface.
Epidermis.—Thickness, = in.
Breadth, about ;4, in.
Cortical Cavities.— ,,1n., roundish, numbering from 8 to none.
Crystals. —Moderately numerous.
Hard Bast.—Sparingly developed, discontinuous; one or a few cells thick.
Wood-curve (see Fig. 40).—Thickened body, and stout short horns, usually
gently incurved.
Vessels.—Largest, ,4, 1.
Average, ja, 1.
Moderately large and numerous, mainly distributed in upper portion.
Characteristic features :
Section.— Small, equally broad and thick.
Cortical Cavities. —Small and variable in number.
Hard Bast.—Scanty.
Wood-curve.—Moderately thickened, and short, stout horns.
Vessels.—Medium-sized, numerous, and mainly distributed in upper
portion of wood.
XIV.—Forms Betoneine to West Austraia, SoutH AUSTRALIA, VICTORIA, AND
New SoutH Watgs.
We are indebted to Mr. Bosisto for specimens of the leaves of E. uncinata
(Turezaninow) and EL. oleosa (F. v. M.), from the district between Dimboola and Lake
AS AIDS IN THE DETERMINATION OF SPECIES, 45
Hindmarsh. They constitute large portions of the ‘‘ Mallee Scrub,” and along with
E. incrassata, Labillardiére, and its smaller variety, H. dumosa, are the only species
of Eucalypt with the above Australian distribution, according to present knowledge.
They are all shrubby, or at most somewhat arboreous, with leaves equally green
on both sides.
Only a few sections were made of wncinata and oleosa, but there is a decided
difference between the wood-curve of each, so that they are thereby easily
distinguished. (Figs. 41 and 42.)
It is believed that the foregoing descriptions, with the accompanying drawings,
when taken in conjunction with external leaf characters, will materially aid in the
discrimination of the investigated species. Various specimens have been taken at
random and tested, and referred to their respective types.
As a very instructive example of this, no better could be given than the
following :—One of us had a few leaves sent without any information as to their
nature or locality, and sections were made in due course, and the following notes :—
“The leaf looks like that of citriodora, but the characteristic odour is wanting.
The first section shows two canals, and fairly numerous cortical cavities ; outline
somewhat like citriodora. A second section from another petiole has very irrecular
outline, but otherwise agrees with the first. A third section was cut from another
petiole, which was still more like citriodora, a third canal tube appearing as in one
of the type specimens of that species. I conclude that this specimen is either a
variety of citriodora, or another very closely allied species.” The leaf belonged to
E. maculata, thus justifying, from quite an independent source, Baron von Mueller’s
determination of citriodora as a variety of that species.
And, in the practical test to be presently described, the utility of the method
will be more fully shown. But, apart altogether from the practical utility of this
investigation, there is new light thrown on the histology of this important genus.
M. L. Petit’ has described the structure of the petiole of Dicotyledons in 48
families, including 300 genera and nearly 500 species, one of the families being the
Myrtacez, to which the Eucalypts belong.
He found the form of the petiole to be always convex below and concave on the
upper face, whereas our observations show, in the twisted petioles of Kucalypts, that
while this is often the case it is not invariably so. Thus, in obcordata and alpina the
upper surface of the section is more convex than the lower, and in grossa and rostrata
the section is round. The lower surface is generally convex and the upper flat.
* Mem. Sci. Phys. and Nat. Bordeaux III. (1887).
46 THE TRANSEVRSE SECTIONS OF PETIOLES OF EUCALYPTS
As to the fibro-vascular system, it is observed that aggregation of the bundles
generally occurs in woody plants, as we found to be the case, and that bicollateral
bundles occur in the Myrtacez, among others. In the genus Eucalyptus, at any rate,
the bundles are concentric.
However, the importance of the structure of the petiole for taxonomic purposes is
clearly established in this paper, and he naturally insists on the importance of the
petiole for purposes of classification.
A few of the facts recorded in the various suggestive and instructive papers on the
Eucalypts by J. Bosisto, Esq., C.M.G., read at different periods before this Society,*
may be noticed in connection with our present subject. He is led to the important
conclusion that the volatile oil is the base of other products, such as resin-like
substances, inasmuch as those species, great in the production of the oil, vigorously
supply it to the atmosphere, and thus do not allow sufficient time for the formation
of resinoid bodies, which require the absorption of oxygen by the leaf; and,
conversely, those species less vigorous in oil production allow time for such a
purpose, and so become well stored.
This suggested to us that there might be some relation between the size and
number of cortical cavities, and the greater or less production of oil or resinoid
bodies respectively. Two of the chief oil-producing species are amygdalina and
globulus, and of kino-producing species, Jeucoxylon and rostrata. On referring to
the detailed description, amygdalina is found to have the smallest and fewest
cavities of any of these, being even surpassed by viminaiis, which yields the least oil.
Globulus has certainly large and moderately numerous cavities, but, on the other
hand, Jeucoxylon is not far behind it, which yields kino to the extent of even 22 per
cent. from fresh bark.
We have observed that as a rule the notable producers of oil do not show
numerous cortical cavities in their petioles, so that we may perhaps conclude that the
production of kino in the leaf-stalk is not immediately dependent on the production
of oil in the leaf-blade. It is in the leaf-blade alone that the oil is manufactured, as if
was proved to be absent from the petiole, so that, contradictory as it may appear, it is
in the leaf-blade, with its two surfaces equally exposed to sun and air, that the least
oxidised constituent—the volatile oil—is made. ‘To prove the absence of oil from
the leaf-stalk (at least in amygdalina, and presumably in the others), a simple
experiment was made, which may be briefly described.
Branchlets of fresh leaves were kindly supplied by Mr. Slater, of Mitcham
Grove, such as he is in the habit of using for distilling his well-known oil. The
largest and best leaves were selected, and 1000 petioles carefully snipped off and
* See Trans. and Proceedings of Roy. Soc. of Vict., Vol. XII., &e.
AS AIDS IN THE DETERMINATION OF SPECIES. AT
weighed, amounting to a little over 200 grains. They were distilled by passing a
current of steam through them for some time, but not a trace of oil, not even the
smell of it, was obtained.
Hence it would appear that the volatile oil is confined to the leaf blade, the
flower-buds, and the fruit-capsules.
The presence of central canals or reservoirs throughout the length of the petiole
in calophylla, and its close ally ficifolia, maculata, and its variety citriodora, it 1s not
easy to explain. When it is known whether they occur in any, and in what other
forms, one may perhaps venture on an explanation. It is noteworthy that in
calophylla the kino is a liquid of treacle-like consistence, and obtained in considerable
quantity by tapping the tree (Kuc. Dec. X.), and that of maculata is readily soluble
in hot water (Dec. III.)
While weighing the leaf-stalks for the above purpose the weight of the leaf-blades
belonging to them was likewise determined. The blades were sometimes imperfect
from the ravages of insects or decay, so that the relative weight errs on the side of
defect ; 1000 leaf-blades were found to weigh 48323¢rs., and the corresponding leaf-
stalks 212ers., so that, on an average, each stalk has to bear a weight at least 23 times
that of its own.
VI. A Practicat Trsr.
Tn order to settle how far the transverse section of the petiole alone was capable
of determining species, we decided to submit the matter to a practical and very
searching test. The names of the 30 kinds investigated by us were handed to Mr.
Guilfoyle, and he selected specimens of leaves of szx of them. These were sent to
us with numbers attached, while the names were retained by him. It will be seen
that the test was really very severe, even more so than would occur in actual practice.
In the first place the docality of the tree would be known, when met with in its native
state, and by referring to the Table of Geographical Distribution (Table I.), the area
of selection would be limited. In the next place, there are certain characters belong-
ing to the leaf as a whole, which would help to settle specific nature. These two
features are taken into account in the next section, dealing with a scheme for deter-
mining species, but we thought it well to settle how far anatomic structure of the
leaf-stalk, apart from other characters, could be depended on for specific distinction.
This trial justifies us in our belief, that while sections of the leaf-stalk may be
valuable aids in the discrimination of species, it would be unsafe to rely upon them
exclusively.
48 THE TRANSVERSE SECTIONS OF PETIOLES OF EUCALYPTS
The result of this test was that four were named correctly and two wrongly, as
shown in the following, sent by Mr. Guilfoyle :—
YOURS. MINE.
No. 1 E. cornuta Soe EK. cornuta
ana ,, corynocalyx ie 5, corynocalyx
ae ,, gomphocephala. .... ,», gomphocephala
foetal ,, mMegacarpa ath ,, megacarpa
LED », viminalis ae ,, occidentalis
is 6 ,, melliodora ies ,», punctata
It will not be necessary to say much about the first four, which were correctly
named, but the two latter incorrectly named deserve further notice. The following
remarks are meant to show the principal points which guided us in coming to a
decision.
No. 1. E. cornuta showed the characteristic features in the size and shape of the
section; the large and numerous cortical cavities quite close together ; crystals
moderately numerous; the wood-pattern continuous or often discontinuous at the
horns, which are sharply incurved (as in Fig. 12), and the vessels distributed towards
the middle and upper portion of wood.
No. 2. E. corynocalyx showed numerous but comparatively small cortical cavities ;
crystals rather few; relatively few hard bast cells ; symmetrical and generally entire
wood-pattern, with slender, short, and gently incurved horns. (Fig. 27.)
No. 3. E. gomphocephala showed few and moderately large cortical cavities ;
wood-pattern, with very stout body and horns well curved in, and approaching each
other; and the vessels large and numerous, and mostly in upper portion of wood.
The large proportion of wood is a feature of Gomphocephala. (Fig. 17.)
No. 4. E. megacarpa showed flattened section, tending to angularity, and nearly
twice as broad as thick; hard bast irregular, and cells arranged in patches ; wood-
pattern, with slender body, slightly thickened in middle, and short horns gently
incurved. (Fig. 18.)
No. 5. E. occidentalis (determined as E. viminalis). Since E. occidentalis is
confined to Western Australia, and £. viminalis does not occur there, it is evident
that the locality of the specimen would have prevented that mistake.
As this particular specimen was somewhat puzzling to determine, it received
special attention, and the following diagnosis of it was made by one of us :—
Outline of Section.—Oval ; flattened on upper surface.
Outer wall of Epidermis.—,,, in., relatively thin.
AS AIDS IN THE DETERMINATION OF SPECIES. 49
Cortex.—Consisting of rather loose tissue.
Crystals.— Very numerous and in clusters.
Cortical Cavities.—Five is a common number ; sometimes very large, about
a il.
Wood-pattern.—Always unbroken.
Vessels.—Moderately numerous; distributed towards upper portion and
middle of wood ; largest about {4 in.
Body of curve is thickened about middle, and the two short horns curve
gently and symmetrically upward and inward.
The thinness of the epidermis, the number and relative size of the cortical
cavities, the loose tissue of the cortex, the hard bast of few cells, but regularly
distributed, the wood-pattern, and the distribution of the large and numerous vessels,
seemed to point to viminalis. But the specimen turns out to belong to a tree which
is exceedingly variable; ‘‘so variable,” as Baron von Mueller expresses it, ‘‘ as to
change much of former ideas as regards the precincts of Hucalyptus species,” and
therefore it would have been a mere accident to have determined it accurately from
the construction of the leaf-stalk alone.
No. 6. E. punctata (determined as E. melliodora). There is a general agreement
between these two forms in the small size and round shape of the section, the
thickness of epidermis, the number and size of cortical cavities, the hard bast composed
of few cells, and the general features of the wood-curve. But in the former the
vessels of the wood are relatively few, small, and scattered ; and in the latter twice
the size, numerous, and equally distributed. Further, crystals are very few in
punctata, but very numerous in melliodora.
It is thus shown in a practical way how the minute structure of the leaf-stalk
may be employed in the acknowledged difficult task of determining Eucalyptus
species. As might be anticipated, this structure will vary according to differences in
soil and climate, heat and moisture, but there is still a certain amount of constancy
in the internal characters which renders their aid valuable. It is not the failure of
the system adopted, but the imperfection of it, which prevents its wider application.
Taking the characters of the leaf as a whole, and not merely sections of the petiole, it
is believed that specific distinctions might be decided thereby.
50 THE TRANSVERSE SECTIONS OF PETIOLES OF EUCALYPTS
V1II.—Scueme For DETERMINING SPECIES.
As our investigations proceeded it became evident that some scheme of arrange-
ment might be devised, whereby the different species could be more or less completely
separated, by taking all the characters derivable from the leaf. In framing this
scheme we took the external leaf-characters principally from Baron von Mueller’s
‘Kucalyptographia,’ where the most minute peculiarities are clearly noted, and
combined them with such characters as were peculiar to the section of the leaf-
stalk.
It will be understood that this scheme is by no means meant to be final, since it
is concerned with thirty species selected at random from the whole range of genera ;
but it is merely given to show iow such ascheme might be constructed for any definite
and restricted section of the Eucalyptus-species. Thus, when all the Victorian species
have been investigated as to the structure of their petioles, it will be possible so to
arrange them that the different species will be more or less clearly marked out.
With this qualification, the scheme applicable to the thirty species will now be
indicated, and may be generally represented as follows :—
Section I.—Central Canals present.
yolles Sy ,» absent.
Section I.
A. Cortical Cavities large —
1. Hard Bast well developed—
1. E. maculata.
2. Hard Bast feebly developed—
2. E. maculata, var. citriodora.
3. EH. calophylla.
B. Cortical Cavities small——
4. K. ficifolia.
Szcrion II.
A. Leaves equally green on both sides—
1. Cortical Cavities large—
a. Hard Bast well-developed—
AS AIDS IN THE DETERMINATION OF SPECIES, 51
A. Vessels of Wood relatively large—
5. E. cornuta.
6. Ki. tetraptera.
7. KE. obliqua.
8. E. gunnii.
9. EK. megacarpa.
10. E. macrorrhyncha.
11. KE. globulus.
B. Vessels of Wood small—
12. K. alpina.
13. E. viminalis.
b. Hard Bast feebly developed—
A. Vessels of Wood relatively large—
14. E. leucoxylon.
15. E. grossa.
16. E. occidentalis.
B. Vessels of Wood small—
17. E. cornuta var. lehmanni.
18. E. obcordata.
2. Cortical Cavities small—
a. Hard Bast well developed—
19. KE. stuartiana.
20. EK. melliodora.
b. Hard Bast feebly developed—
. 21. H. amygdalina.
22. EK. rostrata.
23. E. rudis.
24, EK. tereticornis.
B. Leaves unequally green on both sides—
1. Cortical Cavities large—
a. Hard Bast well developed—
25. KE. gomphocephala.
26. E. marginata.
b. Hard Bast feebly developed—
27. K. saligna.
28. H. punctata.
2. Cortical Cavities, small
Hard Bast feebly developed—
29, KE. corynocalyx.
30. I. diversicolor,
or
bo
THE TRANSVERSE SECTIONS OF PETIOLES OF EUCALYPTS
The first and most general division is based upon the presence or absence of
central canals.
Then such a limited Section as I. is comparatively easily disposed of. The
cortical cavities are either large or small.
Section I.
A. Cortical Cavities large—
1. Hard Bast well developed—
1. E. maculata.
2. Hard Bast poorly developed—
a. Leaves, lemon-scented, and equally green on both sides—
2. HK. maculata, var. citriodora.
b. Leaves unequally green on both sides—
3. H. calophylia.
B. Cortical Cavities small—
4. Et. ficifolia.
In Section II. the first division may be based upon the leaves being equally
or unequally green on both sides.
Section IT.
A. Leaves equally green on both sides—
1. Cortical Cavities large—
a. Hard Bast well-developed—
A. Vessels of Wood relatively large—
(1) Section small—
5. HE. cornuta.
(2) Section large—
(a) Epidermis very thick—
6. E. tetraptera.
(0) Epidermis thickish and wood-curve slender—
7. EH. obliqua.
(c) Epidermis thinner and wood-curve thicker—
8. EK. gunnii,
(3) Section of average size—
(a) Section twice as broad as thick—
9. EH. megacarpa
(6) Wood-curve always entire, and exceedingly
thickened—
10, E. macrorrhyncha,
AS AIDS IN THE DETERMINATION OF SPECIES. 53
(c) Wood-curve almost always entire, and ordi-
~ narily thick
11. E. globulus.
B. Vessels of Wood small—
(1) Section large and epidermis of average thickness—
12, K. alpina.
(2) Section of average size and epidermis thin—
13, E. viminalis.
6, Hard Bast poorly developed—
A. Vessels of Wood relatively large—
(1) Section large, twice as broad as thick—
14. EK. leucoxylon.
(2) Section of average size—
(a) Epidermis relatively thick—
15. E. grossa.
(6) Epidermis of average thickness—
16. E. occidentalis.
B. Vessels of wood small—
(1) Section twice as broad as thick—
17. EK. cornuta, var. lehmanni.
(2) Section as broad as thick—-
18. E. obcordata.
2. Cortical Cavities small—
a. Hard Bast well-developed—
A. Epidermis relatively thin—
19. EK, stuartiana.
B. Epidermis of average thickness—
20. HK. melliodora.
b. Hard Bast feebly developed—
A. Wood-curve excessively thick—
21. EH. amygdalina.
B. Vessels medium-sized—
22. HK. rostrata.
c, Vessels comparatively small—
(1) Epidermis thicker—
23. EH. rudis.
(2) Epidermis thinner—
24. H. tereticornis.
Section II, B. may be further subdivided according to the size of the cortical
cavities and the development of the hard bast.
54 THE TRANSVERSE SECTIONS OF PETIOLES OF EUGALYPTS
B, Leaves unequally green on both sides—
1, Cortical Cavities large—
a. Hard Bast well-developed—
A, Wood-curve exceedingly thickened—
25. EK. gomphocephala.
B, Wood-curve slender—
26. E. mareinata.
b. Hard Bast feebly developed—
A, Vessels moderately large and numerous—
27. EK. saligna,
B. Vessels small and few—
28. E. punctata.
2. Cortical Cavities small—
a, Comparatively numerous—
29. K. corynocalyx.
b, Relatively few and very small—
30. E. diversicolor.
VIII. Srectres ComMPARED AND CoNTRASTED.
Before instituting comparisons between different species, it seemed advisable to
give the characters for all the species as derived from the transverse sections, and
then one was in a position to say where resemblances might be traced. Of course it
must be borne in mind that out of 134 recorded species, only 30 different kinds are
here considered, so that the range is rather limited for making perfectly just
comparisons ; still, as far as it goes, it will be useful to take a comparative view, at
least for some of the species.
In the ‘“ Eucalyptographia,” the affinity between certain species (and even
probable identity) is often pointed out, and we will now select some of the more
important of these, in order to see how far we agree with, and wherein we differ from,
Baron von Mueller, in his views of affinity, views which are based upon other
characters, and are the result of prolonged, and careful, and exhaustive study of this
special group. It will facilitate this comparison if we give a brief abstract of the
allied species. In the following table (III.) the principal affinities are classified, and
short reference is made to each :—
Or
On
AS AIDS IN THE DETERMINATION OF SPECIES.
TABLE III.
SPECIES AND THEIR ALLIES (Accorpine ro Baron von MUELLER).
SPECIES. ALLIED SPECIES. REMARKS.
1. Globulus oo oe Alpina
Megacarpa
| 2. Rostrata 50 O° Viminalis “Instances occur when Rostrata merges almost into
Tereticornis Viminalis, and completely into Tereticornis. It is also
Rudis almost linked by exceptional transit-forms with E.
rudis.” (Dec. 4.)
3. Viminalis cc On| Rostrata ‘EK. viminalis is closely allied as well to E. rostrata as to
Stuartiana E. stuartiana.” (Dec. 10).
4. Stuartiana .. a Gunnii
Viminalis
|
5. Ficifolia ae SI Calophylla
6. Maculata ae Sl Citriodora “Citriodora can only be considered a variety differing
merely in the exquisite lemon-scent of its leaves.”
(Dec. 3.)
7. Cornuta 20 O06 Lehmani ‘Specifically inseparable.”’ (Dec. 9.)
8. Occidentalis .. an Cornuta ‘«The lines of demarcation between these three are not
Obcordata always very clear.” (Dec. 6.)
1. As regards E. globulus and E. alpina, it is stated E. alpina ‘stands to that
species in nearest systematic affinity’ (Decade 2), and E. megacarpa likewise ‘‘ bears in
some respects alliance to E. globulus” (Dec. 6). If the sections are compared there is
a suggestion of resemblance between these three species, but in alpina the smaller
vessels and more numerous cortical cavities distinguish it, while in megacarpa the
cortical cavities are generally smaller. In both the comparatively few crystals contrast
with the large number in E. globulus.
2. Again, it is said of E. rostrata and E. tereticornis that “both might be regarded
as forms of one species” (Dec. 9), and of E. rostrata and E. rudis that they are “also
almost linked by exceptional transit-forms” (Dec. 4). It is further stated, ‘‘ On the
whole this is one of the most easily recognised of all species; still instances occur
when it merges almost into E. viminalis, and completely into E. tereticornis”
(Dec. 4).
On referring to the preceding scheme of arrangement it will be seen that rostrata,
rudis, and tereticornis are grouped together, but viminalis does not seem to be very
closely allied. On comparing the sections it is found that they exhibit resemblance
in the following order :—Rostrata, tereticornis, rudis.
It is likewise interesting to note that both E. rostrata and E. tereticornis have
their leaves sometimes devoured by the same insect (Dec. 9).
When rostrata and tereticornis are compared as regards the minute structure of
the leaf-stalk (and it may be mentioned that the characters of each were drawn out
quite independently), itis found that their close affinity, determined on other grounds,
is borne out.
56 THE TRANSVERSE SECTIONS OF PETIOLES OF EUCALYPTS
Summarising the principal differences between them, as given in the ‘‘ Hucalypto-
graphia,” and in the more recent ‘‘ Key to the System of Victorian Plants,” as well
as those deduced from the transverse section of the petiole, they are as follows :—
Rostrata. TERETICORNIS.
Calyx-tube .. --| Nearly hemispherical .. go ..| Almost semi-ovate.
Calyz-lid .. --| Hemispheric and sharp-pointed at top..| Elongate-conical—longer and blunter
at top.
Stamens G0 --| Outer inflexed before expansion ..; Outer straight before expansion (room
for straightening in the longer
calyx-lid).
Section of Petiole—
Crystals 60 -.| Moderately numerous .. 5¢ -.| Very numerous.
Vessels bo --| Moderately large .. oe ce -.| Comparatively small.
Such small differences completely justify teveticornis being regarded as merely
a variety of rostrata, but the latter, being the most cosmopolitan of all the Eucalypts
as far as Australia is concerned, might be expected to have undergone some
modification of structure in order to adapt it to its wider environment.
3. E. vinunalis is said to be closely allied to E. stuartiana as well as to E.
rostrata (Dec. 10). The thicker epidermis of LE. rostrata, the generally smaller
cortical cavities, the feebly developed hard bast, and the relatively larger vessels
distinguish it. Stwartiana agrees, on the other hand, generally in the thinness of
the epidermis, the development of the hard bast, and the small size of the vessels,
but the cortical cavities are much smaller.
4. When stuartiana and gunn are compared, the differences are clearly
marked. In gunn the thicker epidermis, the larger cortical cavities, and the wider
vessels form a contrast, but in the development of the hard bast, and the thickness
and curving of the wood, there is general agreement.
5. Ficifolia and calophylla are nearly allied, and it is said of the latter that it is
‘the only species to which LE. ficifolia bears very close alliance” (Dec. 7).
The sections reveal a very close general agreement indeed, the presence of two
internal canals normally in each, showing this. Calophylla has a much thicker wood-
pattern, and large and numerous cortical cavities, while in ficifolia they are very few
and very small, but the greater size of the former tree, as compared with the latter,
might account for this difference.
6. Maculata and citriodora may be considered here, since both possess the
internal canals. It is stated that ‘“citriodora can only be considered a variety of
maculata, differing merely in the exquisite lemon-scent of its leaves” (Dec. 3). And
AS AIDS IN THE DETERMINATION OF SPECIES. 57
it is a striking corroboration of this that a study of the two sections leads to the
same conclusion. Further, there is a remarkable resemblance between the relations of
the sections of maculata and citriodora, and calophylla and ficifolia. There is the
similar stout and slender wood-pattern respectively in both, although the cortical
cavities are about equally numerous and equally large in maculata and citriodora. In
ficifolia it would appear that the stunted habit of the tree had rendered large and
numerous cortical cavities less necessary, and so they have dwindled down in size and
numbers, whereas the variety of maculata has lost nothing of its stateliness. Such
are some of the affinities which have been noted by Baron von Mueller, and which
are borne out as well by the transverse sections of petioles. Let us turn now to some
others which do not agree with our sections.
The affinities of E. viminalis have already been discussed.
7. E. lehmanni is said to be “ specifically inseparable from E. cornuta” (Dec.
9), but a glance at the two sections would not suggest it. There is resemblance in
the large and numerous cortical cavities, and in the much broken wood-pattern, but the
vessels in cornuta are much larger and more numerous, and the wood-curve is consider-
ably thicker. Altogether, the resemblances in the sections would not justify their being
classed together as the same species, or the one as a variety of the other.
How are we to regard a discrepancy of this sort? Are we to regard the deep-
seated characters of the leaf-stalk as delusive ? External resemblances and allied
internal anatomy do not always go together, for there may be fundamental difference,
accompanied by superficial resemblance, just as there may be fundamental resemblance
with superficial difference. We consider that the relationship may still hold between
E. cornuta and E. lehmanni, the latter being a variety of the former, even in spite of
seeming contradiction from the section, when it is remembered what aberrant and
extreme forms are assumed by some species, and the extent of their variability.
That this applies here will now be shown. In arranging our sections independently of
external characters in the first instance, E. occidentalis was placed next to E.
lehmannt, which naturally formed the last term of the present series. The section
of the petiole of E. occidentalis exhibits a wood-pattern of such an irregular nature
that it is difficult to say what it resembles exactly, and yet it is suggestive of
resemblance to a number of forms. It is, in short, just such a form of section as
might easily develop into several of the others, and so we were unable to correlate it
with any. In considering the affinities of E. occidentalis in the “ Kucalyptographia ”’
(Dec. 6), much light is thrown on the present difficulty.
First of all, we are informed that E. cornuta, E. obcordata, and E. occidentalis
““seem to be the only three entitled to specific rank in the series of cornuta or
orthostemoneae, and even the lines of demarcation between these three are not always
58 THE TRANSVERSE SECTIONS OF PETIOLES OF EUCALYPTS
very clear.” So this brings E. occidentalis into some sort of affinity with E. cornuta,
and, by implication, with its variety, E. lehmanni. When the section of the latter is
compared with E. occidentalis, there is undoubtedly a certain resemblance, but not
directly, with E. cornuta.
Again, it is found from extended observation and increased material, that E.
occidentalis runs more or less into at least six named species (not reckoning two
varieties of E. cornuta), viz. :—
E. spathulata, Hook.
E. macrandra, F.v. M.
E.. cornuta, Labillardiére.
E. obcordata, Turez.
E. redunca, Schauer.
. pachypoda, F.v. M.
ty
And the inference is that E. occidentalis is capable of and subject to a vast
amount of variability.
Nay, further, in the concluding sentence of the notice of this form, it is stated
“ . occidentalis, in its scope as here considered, seems so variable as to change much
of former ideas as regards the precincts of Eucalyptus-species, a similar playfulness
of forms having been observed by me in FE. stricta and E. incrassata, the characters of
shrubby Eucalypts proving generally less constant than those of the tall timber-trees
of this genus” (Dee. 6).
In such a sentence as that there is contained the essence of a volume on the
“Origin of Species.’’ And when we consider that at least three species—E. glauca,
E. pulverulenta, and E. perfoliata—have been based upon the young state of E.
globulus (Dec. 6), it is not to be wondered at, but rather to be expected, that in such
a variable genus of plants there would be striking resemblance in internal structure,
as well as a certain amount of diversity.
IX.—Conciupine REMARKS.
If transverse sections of the petioles can be used as an important aid in the
determination of species, then it is evident that this test may be applied where there
is doubt as to specific identity.
AS AIDS IN THE DETERMINATION OF SPECIES. 59
Thus there is a remarkable resemblance, for instance, between the sections of
the petioles of E. calophylla and E. ficifolia, and the question arises—Are they to be
regarded as distinct species or not ?
The principal differences between LF. ficifolia and £. calophylla, as given by
Baron von Mueller in his “‘ Eucalyptographia”’ (Dec. 7), are as follows :—
E. ficifolia.
Tree.—Of less height.
Bark.—Somewhat more deeply furrowed.
Leaves.—Proportionately not quite so broad, but longer.
Flowers.—Mostly larger.
Calyces.—Assume a reddish hue.
Fruits.—Less turgid.
Seeds.—Much paler in colour, have a smaller kernel, and are provided with con-
Spicuous appendicular membranes.
If the leading specific differences of the same authority be compared, it will be
found that the crimson filament and the pale seeds, with a long membrane, of E. fici-
folia ave the principal offsets against the pale-yellowish, rarely pink filaments, and
dark membraneless seeds of E. calophylla.
Here the differences are comparatively small, but the characters, such as they
are, being permanent, are considered sufficient to fix this as a distinct species.
If we turn to the characters revealed by the petiole, the same remark applies.
The broad differences separating ficifolia from calophylla are :—
Slightly thicker epidermis, with a strong development of hypoderm, although
there are indications of something similar to the latter in calophylla.
Cortical Cavities.—Much fewer and smaller.
Central Canals.—Smaller.
Vessels.—About the same size, slightly larger, but weaker development of
wood.
Such permanent differences probably entitle E. ficifolia to specific rank, but it
will be observed that the differences are just such as might arise from the same kind
of tree, from any cause, becoming dwarfed and stunted in growth.
Both belong to the furrowed bark series (Schizophloiz), have anthers opening by
parallel slits (Parallelantherz), and pollen-grains of the same size. The fruits in both
are urn-shaped, and their habitat is confined to Western Australia.
60 THE TRANSVERSE SECTIONS OF PETIOLES OF EUCALYPTS
To take just another striking example. £. citriodora is regarded on general
erounds as a variety of E. maculata, and from the transverse sections of the petioles
of both the same conclusion might be drawn. It is a suggestive fact that the
resemblances and differences are mainly of a parallel kind with those of E. ficifolia
and E. calophylla. But the differences are evidently less in degree, or rather the
resemblances are closer, for they generally agree in thickness of epidermis, size, and
number of cortical cavities, and size of central canals. But the vessels are about
twice the size of those of maculata, while the wood itself is relatively much more
feebly developed.
On the whole the study of the transverse sections of petioles of well-developed
and fully-formed leaves of Eucalypts has led us to the following conclusions :—
1. Fresh species may be identified by means of such sections, combined with
external characters of the leaf. The aggregate characters of the section are employed
but chiefly those derived from the following parts:—Epidermis, hard bast, wood with
its vessels, cortical cavities, and central canals.
2. Herbarium material, by appropriate treatment, may likewise be used.
3. The size and shape of the transverse section is often characteristic.
4. The relative thinness or thickness of the epidermis is a more or less
constant feature, and it is often indicative of the habitat of a plant, say of a desert
species.
5. The number, size, and arrangement of cortical cavities are very variable, but
their relatively large size in some species is such a striking feature that it may be
used for purposes of discrimination.
6. The central canals are so decided and distinct, and occur in such a limited
number (at least of thirty-two species and varieties examined), that they might con-
veniently be used as a diagnostic character for a section of the Kucalypts.
7. Crystals are always present, more or less, but they form too precarious a
feature to be relied upon.
8. The hard bast, whether dense or scanty, seems to be generally characteristic.
9. The wood-curve is likewise a characteristic feature, with the relative number,
size, and arrangement of vessels; but in some few instances it is too variable to be
much relied on.
10. The broken nature of the wood-curve, as found in some species, cannot be
used as a distinctive character, like the invariably unbroken curve in others, because on
fuller investigation unbroken curves are found in all.
AS AIDS IN THE DETERMINATION OF SPECIES. 61
11. The closeness of resemblance in the sections of two allied reputed species
may lead to their being recognised merely as varieties; or, on the other hand, the
differences between reputed varieties may be sufficiently great to suggest their
distinction as species.
12. Lastly, the general agreement among the various sections, and the absence of
those clear distinctions which would mark off each species if quite independent, tend
to the conclusion that the varied species of Eucalypts may have arisen through a
process of evolution. After allowing for the changes which may have taken place
through possible hybridisation, as hinted at in the “ Kucalyptographia”’ (Dec. 9),
‘extreme forms,” ‘“ aberrant forms,’ and “transition forms,” so frequently referred
to in that work, are not thereby accounted for. While the mere existence of
transition forms does not prove that the transit has been accomplished through
descent with modification, still permanent varieties or species, with such forms
flanking them on every side as would be regarded as distinct species if isolated, would
seem to have arisen thus. Hence the proper study of the Eucalypts, like that of the
Platypus and Ceratodus, may add another page to Australia’s contribution to the
evolution theory. But perhaps it is hardly necessary to formally state this conclusion
in the face of the recent utterance at the Newcastle meeting of the British
Association, 1889, by the distinguished President (Prof. Flower) :—‘ I think I may
safely premise that few, if any, original workers at any branch of biology appear
now to entertain serious doubt about the general truth of the doctrine that all
existing forms of life have been derived from other forms, by a natural process of
descent with modification.”
Since this investigation is strictly comparative, we have endeavoured to preserve
as much uniformity as possible, both in the sections made and the language used,
Besides, we felt that an important principle was being tested here, and one
which might receive a wider application, viz., that in many families of plants the
intimate structure of the leaf-stalk might be made a means or an aid in the
identification of species. Therefore it was necessary to select a sufficiently varied
number of examples, as well as a fair proportion of the entire number of known
species on which to base our conclusions. It will be conceded, we think, that the
Eucalyptus formed a sufficiently critical genus to test, and that the number of species
(30), forming nearly one-fourth of the whole, was fairly extensive. According to the
latest ‘‘Census” by Baron von Mueller, the number of Australian species of Eucalypts
is 184. Whatever conclusions may be drawn, the photographed sections, and the
accurate outlines, particularly of the wood, ought to be a decided addition to our
knowledge of this characteristic and peculiar form of Australian vegetation.
62 THE TRANSVERSE SECTIONS OF PETIOLES OF EUCALYPTS
EXPLANATION OF PLATES.
REFERENCE LETTERS TO ALL THE FIGURES.
b. bast.
ce. cuticle.
cc. central canals.
ce. cavity of epidermal cell.
cp. cortical parenchyma.
er. crystals.
ct. cortex.
ctc. cortical cavity.
cw. cuticularised cell-wall.
e. epidermis.
h. hypoderma.
hb. hard bast.
k. kinoid substance.
p. pith.
sb. soft bast.
sp. sieve plates.
st. sieve tubes.
vw. vessels of wood.
w. wood.
we. wood curve.
PLATE 1.
Fig. 1. Epidermal cells of 2. globulus, shortly after treatment with Schulze’s solution, showing also hypoderma (h)
(x 114).
Fig. la. Single epidermal cell of samc, with underlying cells (x 600).
Fig. 2. Cortex of EZ. globulus, showing a cortical cavity (x 114).
Fig. 2a. Cortical cells adjoining hard bast (x 114).
3. Crystal in cortical cells of L. ficifolia (x 140).
4, Cortical parenchyma bordering on hard bast, in vertical section (x 600)
Fig. 5. Hard bast and adjoining tissue in vertical section (x 600).
6. Cambiform cells of soft bast, in vertical section—comparatively elongated and thin-walled (x 600).
Fig. 6a. Cambiform cells, from round to brick-shaped—the commonest form (x 600).
7. Soft bast in vertical section, showing sieve-plates (sp) on radial side-walls (x 600).
Fig. 8. Transverse section of base of midrib of young sessile leaf, to show not only structure of midrib, as far as the
wood-curve is concerned, but mainly for comparison with section of petiole of mature leaf.
Fig. 8a. Natural size of section.
Fig. 9. Transverse section of quadrangular stem of young H. globulus for comparison with that of petiole.
Fig. 9a. Natural size of section represented in Fig. 9.
Fig. 10. Transverse section of stem of young EZ. calophylla for comparison with that of F. globulus.
Fig. 10a. Natural size of section represented in Fig. 10.
Norr.—All the preceding drawings belong to H. globulus, except Figs. 3 and 10.
In Plates 6, Ga inclusive, the natural size of the section of each petiole is shown beside the enlarged section, and thus
the amount of enlargement throughout is seen at a glance.
Fig. 22a,
Fig. 23.
Fig. 23a.
Fig. 24.
Fig. 25.
Fig. 26.
AS AIDS IN THE DETERMINATION OF SPECIES.
PLATE 2.
E. occidentalis—Transverse section of petiole.
(Glycerine jelly).
E. cornuta— do. do.
(Balsam).
FE. cornita, var. lehmanni—Transverse section of petiole.
(Glycerine jelly).
E. obcordata— do. do.
(Glycerine jelly).
FE. marginata— do. do.
(Glycerine jelly).
E. diversicolor— do. do.
(Glycerine jelly).
PLATE 3.
E. gomphocephala—Transverse section of petiole.
(Glycerine jelly).
E. megacarpa— do. do.
(Glycerine jelly).
EE. tetraptera— do. do.
(Balsam).
EL. rudis— do. do.
(Glycerine jelly).
E. grossa-— do. do.
(Glycerine jelly).
EB. calophylla— do. do.
(Glycerine jelly and iron stain).
PLATE 4.
E. calophylla—Transverse section of petiole.
(Balsam).
E. ficifolia— do. do.
(Glycerine jelly, nitric acid, and iron stain).
E. ficifolia—Transverse section of petiole.
E. alpina— do. do.
(Glycerine jelly).
E. punctata— do. do.
(Glycerine jelly).
E. maculata, var. citriodora—Transyerse section of the petiole.
(Glycerine jelly and iron stain).
PLATE 5.
63
E. maculata, var. citriodora—Transyerse section of another petiole, showing abnormal number of Central Canals
(Glycerine jelly).
I. corynocalyx—Transyerse section of petiole.
(Glycerine jelly).
E. melliodora— do. do.
(Glycerine jelly).
E. tereticornis— do. do
(Glycerine jelly).
E, saligna— do. do
(Glycerine jelly).
E. maculata— do. do.
(Glycerine jelly
64 THE TRANSVERSE SECTIONS OF PETIOLES OF EUCALYPTS.
PLATE 6.
Fig. 32. EH. macrorrhyncha—Transverse section of petiole.
(Balsam).
Fig. 33. E. globulus— do. do.
(Glycerine jelly and iron stain).
Fig. 33a. E. globulus—Transverse section of petiole.
(Glycerine jelly).
Fig. 34. EH. obliqua— do. do.
(Balsam),
Fig. 35. E. amygdalina— do. do.
(Glycerine jelly).
Fig. 36. E. viminalis— do. do.
(Glycerine jelly).
PLATE 6a.
Fig. 36a. E. viminalis—Transverse section of petiole.
(Glycerine jelly).
Fig. 37. EH. gunnii— do. do.
(Balsam).
Fig. 38. LE. stuartiana— do. do.
(Glycerine jelly).
Fig. 39. E. leucoxylon— do. do.
(Glycerine jelly).
Fig. 39a, b,c. E. lewcoxrylon—Outlines of three sections from different petioles, invariably showing broken wood-curve.
Fig. 40. E. rostrata—Transverse section of petiole.
(Glycerine jelly).
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On tHE Victor1aAN Lanp Puanarians, spy AntHuR Denpy, M.Sc., F.L.S., Demon-
STRATOR AND AssisTANT LicturER In Bronogy aNnD FELLOW oF QUEEN’S
CoLLEGE IN THE University or MELBouRNE.
(With Plate 7.)
(Read May 8th, 1890.)
INTRODUCTION.
The present paper may be regarded as a sequel to my memoir on the Anatomy
of an Australian Land Planarian, already published in the Transactions of this
Society.* Having fully described, in the memoir referred to, the anatomy of one
species, which may be taken as a type of the whole group, it remains to name,
briefly describe, and where possible figure the other Victorian species.
There are, as I have pointed out in my previous memoir, two genera of
Australian land Planarians, Geoflana, with many small eyes, and Rhynchodemus,
with only two larger ones. Both of these genera are found in Victoria and it would
doubtless have been desirable to have described in detail the anatomy of a type from
each ; as, however, I have only seen a single specimen of Rhynchodemus, and that
only recently, I have been unable as yet to do this. The gap thus caused is, however,
to a great extent filled by the researches principally of Moseley and von Kennel, to
which frequent reference is made in my earlier paper.
In their ‘‘ Notes on Australian Land Planarians, with Descriptions of Some New
Species ” (Part I.),+ Messrs. Fletcher and Hamilton enumerate eighteen Australian
and Tasmanian species. Of these, seventeen were found in New South Wales, one
in New South Wales and Queensland, one in New South Wales and Victoria, and one
in Tasmania only. In the present paper I propose to describe fifteen Victorian
species, Viz.:
. Geoplana cerulea, Moseley sp.
. quinquelineata, Fletcher and Hamilton.
. munda, Fletcher and Hamilton.
. Spenceri, Dendy.
. ad@, X. sp.
. lucasi, n. sp.
SR YE
QAAHRAO
* Transactions of the Royal Society of Victoria, 1889.
+ Proceedings of the Linnean Society of New South Wales, Vol. II. (Series 2), p. 349.
66 ON THE VICTORIAN LAND PLANARIANS.
7. G. m‘mahont, n. sp.
8. G. alba, n. sp.
9. G. hoggi, n. sp.
10. G. sugdeni, n. sp.
11. G. mediolineata, n. sp.
12. G. quadrangulata, n, sp.
13. G. walhalle, n. sp.
14. G. fletcheri, n. sp.
15. Rhynchodemus victoria, un. sp.
Of these fifteen species the first three occur also in New South Wales and have
been already described by Moseley and Fletcher and Hamilton. The twelve
remaining species are, so far as is yet known, exclusively Victorian, but it is more
than probable that some of them will sooner or later be discovered in one or other
of the adjacent colonies. Still it is evident, when we consider that out of twenty-
nine known Australian species, nearly equally divided between the colonies of
Victoria and New South Wales, only three have been found in both colonies, that
the land Planarians, however widely they may be distributed as a class, do not enjoy
wide specific areas of distribution; or, in other words, that in different districts of
the same country we may expect to find different species of Planarians. In spite of
the occurrence of one or two comparatively widely ranging species, such as Geoplana
alba, which I have had from M‘Mahon’s Creek, Warragul, Macedon, and Croajin-
golong, this expectation is justified by my own observations.
With regard to the question of specific distinctions I have come to the conclusion
that we may safely rely on a combination of the following characters (1) the colour
and pattern, (2) the position of the external apertures, (3) the general shape of the
body. Although I have examined considerable numbers of the same species, as, for
example, in the case of Geoplana spenceri and G. hoggii, I have not found any
important variations in these respects. Perhaps the most variable species are the
common yellow Planarians which I have named Geoplana sugdeni and G. mediolineata.
These two species agree fairly well in the general shape of the body and in the
position of the external apertures, but the former has none of the stripes present in
the latter. ‘The two species are, however, connected by an intermediate variety with
imperfect stripes. Under these circumstances I have assigned all specimens with
any stripes at all to G. mediolineata and all those without any to G. sugdent. It
must, however, be remembered that although in G. mediolineata the number of
stripes may vary yet the arrangement is always identical, and this is the important
point. In the most perfectly striped examples which I have seen we find a single,
fine, dark median stripe, a lighter stripe at some distance on each side of it, and
outside this again on each side an indication of a stripe at the anterior extremity of
ON THE VICTORIAN LAND PLANARIANS. 67
the body only (Figs. 3, 8a), All these stripes may be more or less suppressed, the
first to disappear being those on the outside and the last the one in the middle; or
we may with equal justice assume that the parent form had no stripes and that the
first to appear was the median one and the last the outside ones. The observations
of Fletcher and Hamilton on the markings of very young specimens of Geoplana
quinquelineata, which will be found quoted in the description of that species, seem to
indicate that the latter assumption is the correct one. So it is in most of the species;
the markings, if there are any, may be more or less strongly emphasised in different
individual specimens, but fundamentally the pattern is the same in all. I have
pointed out* the same fact in the case of Peripatus leuckartii.
We now know a good deal about the general habits of land Planarians. They
are What I have elsewhere+ termed ‘‘ Cryptozoic” animals, living for the most part
under logs and stones, or the dead bark of trees. They prefer moderately damp
situations and appear to be much more abundant in the autumn, after rains, than in
the drought of summer. When at rest in their hiding places they commonly lie
coiled up with the anterior extremity in the centre of the coil (Fig. 12). Asa rule
they seem to venture abroad in search of food at night only, or in very damp weather,
but I have more than once found Geoplana sugdeni crawling about in broad daylight,
once on a stone near the top of Mount Macedon, in a decidedly dry situation.
Geoplana lucasi, again, was obtained by Professor Spencer at a height of 4000 feet
on the top of the coast ranges in the Croajingolong district.
Still there can be no doubt that land Planarians require a very considerable
amount of moisture for their existence. Thus I once found a considerable number
of specimens which had escaped from the collecting box dried up on the floor of a
room, having apparently exhausted all their supply of moisture in the production of
their slimy track.
In this connection I may mention that when a living land Planarian is placed
in loose dry earth, it forms a cyst for itself by cementing together the particles of
earth with its slimy secretion. Within this cyst the Planarian lies completely
hidden, but if the cyst be torn it will crawl out perfectly clean and free from earth.
This habit of forming cysts of earth may be a protection against drying up and may
perhaps account for the disappearance of land Planarians in the heat of summer.
Land Planarians crawl with an even, gliding motion, which I believe to be
largely due to the action of the numerous strong cilia on the ventral surface, for I
have seen a minute fragment of the worm, snipped off with a pair of scissors,
gliding away over a smooth surface by itself, much as in the case of detached parts
* Proceedings of the Royal Society of Victoria ; 1889, p. 50, et seq.
} Victorian Naturalist; December, 1889.
68 ON THE VICTORIAN LAND PLANARIANS,
of the common mussel described by McAlpine.* Cilia are probably present on the
dorsal surface also, but very small and much obscured by slime. In crawling the
horseshoe-shaped anterior extremity of the body, on which the eyes are mostly
situate and which has usually more or less of a reddish tinge, is uplifted to gain a
more extended view, and the path of the animal is marked, like that of a snail, by a
slimy track.
They live upon the juices of insects and other small animals, which they extract
by suction, the sucker-like pharynx being inserted in some soft part of the victim
and the latter held fast by the intensely sticky slime secreted around it by its
captor. Mr. C. Frost informs me that on one occasion when out collecting he placed
a living land Planarian in a box with a live Cicada and that when he opened the box
again to show the Cicada to a friend he found the insect quite flat and empty, all the
inside having been sucked out by the Planarian. Hence the land Planarians are
certainly carnivorous and they must be able to find an abundant supply of food
amongst the innumerable cockroaches, beetles and other small animals which make
up so large a proportion of the cryptozoic fauna.
Land Planarians are found breeding in the autumn and winter months and I
have been able to observe the method of copulation in Geoplana mediolineata, at
Upper Macedon on April 7th of the present year. The two individuals were
precisely similar in colour and markings and of about the same size.
They were lying beneath a log in such a manner that the posterior portions of
théir ventral surfaces were applied together, the tail of the one pointing in the
opposite direction to that of the other. The genital atrium in each case was
expanded to form a sucker and the two suckers being applied together held the
worms in position. The orifices of both male and female copulatory organs were in
each case somewhat protruded and it follows from the position of the animals that
the male opening of the one must come into close contact with the female opening of
the other and vice versé. The most important factor in bringing this about is the
use of the common genital atrium as a sucker, in the hollow of which lie the male
and female openings. . Thus the method of action of the copulatory organs,
whose minute anatomy I have described in the case of Geoplana spenceri, 1s
made clear.
The eggs are laid in cocoons, several in each. I was fortunate enough to obtain
recently a specimen of Geopflana hoggii with a cocoon in the uterus, which proves
clearly that the cocoon is an internal structure and not, as in the case of the earth-
worm, an external structure formed by the skin. In the case observed the body of
* Transactions and Proceedings of the Royal Society of Victoria, Vol. XXIV., Part 2, p. 139.
ON THE VICTORIAN LAND PLANARIANS, 69
the worm was seen to be greatly distended just behind the genital opening, and
sections showed the presence of a fully formed cocoon occupying the whole of the
ereatly dilated uterus, and squeezing the other internal organs to one side. The
cocoon itself was crowded with closely packed yolk-cells, whose nuclei and outlines
were still visible. In older cocoons the yolk disappears as the embryos increase in
size until at last the young worms come to occupy the whole of the cavity. Finally
the young escape through simple rupture of the wall of the cocoon or through a
definite circular opening. After having been laid for some time the cocoon appears
as an oval or nearly spherical body of a shining black appearance. In the specimen
which contained the cocoon in utero the yolk glands were, as might have been
expected, very largely developed, and there was some evidence in support of the
suggestion which I made in my previous memoir to the effect that the shell of the
cocoon is secreted by certain glands opening into the small chamber which receives
the contents of the united oviducts before they enter the uterus. To this chamber I
therefore propose to apply the name “ shell-gland chamber.”
The brilliancy of colour and markings in the land Planarians are so well known
that I need scarcely do more than allude to them. Blue, green, yellow, brown, black
and red may be found in various combinations and generally arranged in longitudinal
bands or stripes symmetrically disposed on the two sides of the body. It is difficult
to account for these varied and brilliant markings, which seem, as I have already
pointed out, to be tolerably constant for each species. Wallace proposes* to divide
the colours of animals into four groups which he terms Protective, Warning, Sexual
and Typical. I would suggest that the brilliant colours of land Planarians may be
“warning” colours. There are many butterflies which have been shown by Wallace
to be brilliantly coloured and at the same time inedible by birds and there is good
reason for believing that the brilliant colours serve to warn the birds of the
objectionable character of the butterflies and thus protect the latter from being
pecked to death. In the same way I think it possible, though not as yet by any
means proved, that the colours of Planarians may serve to protect them from being
eaten by birds by rendering them readily recognisable.
But it requires to be shown first that land Planarians are inedible by or
distasteful to birds. I have made two experiments which seem to indicate that this
is the case ; (1) I tasted two species myself and found that the mere application of
the tongue to the slimy surface of the animal was sufficient to produce an exceedingly
unpleasant sensation, something like that caused by putting a piece of velvet ora
lump of alum in the mouth. (2) I threw a living specimen of Geoplana spenceri to a
number of hens. The hens, not being native birds, would, of course, not recognise
the worm, and they at once attacked it, broke it up and took it in their mouths.
* «Tropical Nature, and other Essays,” p. 172.
70 ON THE VICTORIAN LAND PLANARIANS.
Instead, however, of swallowing the pieces they dropped them again. This
experiment, though but a solitary one and of course in need of confirmation, tends
to show that land Planarians are unpalatable to birds as well as to human beings.
In my paper on the anatomy of Geoplana spenceri I have already suggested that
the unpalatable character of these worms may be due to the presence of the rod-like
bodies in the slime with which they are always covered.
In concluding my introductory remarks I have much pleasure in expressing my
indebtedness to my wife for her valuable assistance in collecting specimens; to
Professor W. Baldwin Spencer for numerous Planarians from various localities and
especially for an exceptionally interesting collection from Croajingolong, including a
species of Rhynchodemus, which genus is now for the first time recorded from
Victoria; to Mr. H. R. Hogg, for many specimens of Planarians from Macedon and
for his kind hospitality and assistance whilst I was collecting in that district; to
Mr. J. Bracebridge Wilson, for Planarians from the Otway Forest, and to Mr. Henry
Dendy, of Walhalla, for assistance in collecting in that locality.
DESCRIPTION OF VICTORIAN SPECIES.
Genus Geoplana.
This genus, as I have already had occasion to point out, is distinguished from
the only other known genus o¢ Australian land Planarians by the presence of
very numerous minute, unicellular eyes, which occur principally on the lateral
margins of the anterior end of the body and are usually, if not always, continued
round the edge of the horseshoe-shaped anterior extremity. Frequently, also, the
eyes occur more sparsely scattered along the sides of the body to the extreme
posterior end.
1. Geoplana cerulea, Moseley sp.
1877. Ccnoplana cerulea, Moseley, Quarterly Journal of Microscopical
Science, Vol. XVII., N.S., p. 285.
1887. Geoplana cerulea, Fletcher and Hamilton, Proceedings of the
Linnean Society of New South Wales. Series II., Vol. 2, p. 361,
Pls, Higa ds
“Entire body of a dark Prussian blue colour, somewhat lighter on the under
surface of the body and with a single, narrow, mesial, dorsal, longitudinal stripe of
ON THE VICTORIAN LAND PLANARIANS. (fal
white. Length 5 em., extreme breadth 4 mm., mouth central; generative aperture
8mm., posterior to the mouth. Parramatta, near Sydney. Under the bark of a
species of Eucalyptus.” (Moseley, /oc. cit.)
A number of specimens brought in spirits by Professor Spencer from Croajin-
golong appear to belong to this species. According to Professor Spencer the colour
of the living worms was cobalt on the ventral and dark olive green on the dorsal
surface, exactly as in G. spenceri but with a median, dorsal, yellow line. The spirit
specimens still show very plainly the narrow median dorsal line, and they also show
indications of a narrow, median, ventral light line in the posterior part of the body,
apparently not observed in the living animals. They also still exhibit traces of a
reddish anterior tip.
According to Messrs. Fletcher and Hamilton (Joc. cit.) there appears to be a
certain amount of variation in the colouration of this species. Thus these authors
have found specimens without any red tip and with the dorsal median stripe varying
from a dirty white to a distinct yellow, changing to white in spirit. These obser-
vations make it probable that the Croajingolong specimens are correctly identified as
G. cerulea, though the opening into the peripharyngeal chamber appears to be a
little further back than Moseley places it. In shape this Planarian closely resembles
Geoplana spencert.
Localities —Sydney, Parramatta, Ryde, Springwood, Mount Wilson, Hunter
River (New South Wales); Cairns (N. Queensland) ; Croajingolong (Victoria).
2. Geoplana quinquelineata, Fletcher and Hamilton.
1887. Geoplana quinquelineata, Fletcher and Hamilton, Proceedings of the
Linnean Society of New South Wales. Series II., Vol. 2, p. 366,
Pl. V., Figs. 4, 5, 15, 16.
‘Under surface whitish. Ground colour above presents considerable variations,
pale yellow or nearly orange, dull olive-green, ochreous-brown, reddish-brown,
sometimes almost brick-red. The dorsal surface divided into six longitudinal bands
by five longitudinal lines, also varying in colour, sometimes a darker and more
intense tint of the ground-colour, from dark brown almost black to warm brown or
red, their margins irregular when viewed with a lens, arranged as follows: usually a
very fine dark line occupies the median line, external to which on each side is a
narrow band of ground colour ; outside of which again on either side is a line of
brown or red usually slightly broader and better defined than the mesial line; each
72 ON THE VICTORIAN LAND PLANARIANS.
of these again is bordered by a band of ground colour one and a-half times or twice
as wide as the inner stripe on each side; beyond each of which is the outermost
brown or red line of the same width as the first on each side but sometimes narrower,
and each of these is followed by a narrow band of ground colour extending outwards
to the lateral margin of the body. At the anterior extremity the lines blend in the
red tip. The ground colour, and the reddish tint of the anterior extremity usually
disappear more or less completely in spirit, while the bands become brown or
sometimes black.
Largest living specimen 10 cm. long. In two contracted spirit specimens 42
and 23 mm. long respectively, the apertures of the mouth are 20 and 12 mm.
respectively behind the anterior extremity; in a third specimen 26 mm. long the
genital orifice is 4 mm. anterior to the hinder extremity. In none of our specimens
are both apertures visible.
Young specimens on emerging from the cocoon are 2.5 to 4 mm. long. In
these and sometimes in larger ones the colour of the anterior portion of the body is
more intense. In very young specimens also the lines are brighter, but the
outermost one on each side is only faintly indicated, or absent.’’ (Fletcher and
Hamilton, /oc. cit.)
Messrs. Fletcher and Hamilton state that this is one of the commonest species
in New South Wales, and they also record it from Sandhurst, Victoria. I myself
collected four specimens near Sandhurst, all of which were found under stones on an
old gold-field near the Back Creek Cemetery. The ground colour of the dorsal
surface was, in the living worms, brownish yellow. The five dark lines were placed
nearly equidistant from one another, running all down the body. All five were
rather thin, the middle one black and the others chestnut brown. The ventral
surface was greyish cream coloured.
I have also received two specimens from the Otway Forest, sent to me in spirit
by Mr. J. Bracebridge Wilson. The latter are very large, the largest measuring
60 mm. in length and 5 mm. in breadth even in spirit.
Localtties.—Near Parramatta, near Springwood, near Capertee, Guntawang,
Beaudesert Hills, Biraganbil Hills (New South Wales); Sandhurst, Otway Forest
(Victoria).
ON THE VICTORIAN LAND PLANARIANS. 73
3. Geoplana munda, Fletcher and Hamilton.
1887. Geoplana munda, Fletcher and Hamilton, Proceedings of the Linnean
Society of New South Wales. Series II., Vol. 2, p. 369, Pl. V., Fig. 8.
‘‘Undersurface greyish in centre, yellowish towards the margins. Above
there is a narrow median dorsal line of pale olive brown, bounded on either side by
a very fine dark line, external to which is a broader band of a slightly darker brown,
and this is bordered externally by a very dark brown line which gradually merges
into a rather broad band of very dark brown which fades gradually towards its outer
margin.
“‘This pretty little Planarian retains its colours in spirit very well but the under-
surface becomes quite white. The single specimen obtained measured when alive
and crawling, 2°5 cm. long, and 3mm. broad. In spirit it measures 15 mm. long,
4mm. broad, the mouth 6 mm. behind the anterior extremity, and the generative
aperture 2 mm. behind the mouth.” (Fletcher and Hamilton, Joc. cit.)
As I have not seen this species in the living condition I quote the above
description of Fletcher and Hamilton. Mr. J. Bracebridge Wilson sent me sixteen
specimens in spirits, from the Otway Forest, which agree excellently with the above
account, but I could not find the genital aperture. The species was described by
Fletcher and Hamilton from a single specimen.
Localities —Hartley Vale (New South Wales) ; Otway Forest (Victoria).
4. Geoplana spenceri, Dendy.
1889. Geoplana spenceri, Dendy, Transactions of the Royal Society of
Victoria. Volo. Part, p50; Piss 7, 8; 9) 40:
For the description and figures of this species I must refer the reader to my
former paper.
Localities —M‘Mahon’s Creek, Warburton, Walhalla (Victoria).
5. Geoplana ade, n. sp. (Fig. 7).
The body is nearly oval in section, flattened ventrally, tapering rather more
oradually in front than behind. ‘The opening into the peripharyngeal chamber is
74 ON THE VICTORIAN LAND PLANARIANS.
situate in about the middle or a little behind the middle of the ventral surface. The
eround colour of the dorsal surface is pale yellowish brown. ‘There is one very fine,
median, dark brown stripe, and a very broad lateral dark brown stripe on each side of
it and separated from it by a fairly wide interval of ground colour. The lateral
stripes are rather ill-defined at their outer edges, where they touch a band of the
ground colour splotched with darker brown. The ventral surface is pale yellowish
brown, slightly mottled, or white. The largest of the three specimens which I have
seen was about 40 mm. in length and 3 mm. in greatest breadth when crawling.
I have called this beautiful Planarian after my wife, who has greatly assisted me
in my search for cryptozoic animals.
Localities —Macedon, Warburton (Victoria).
6. Geoplana lucasi, n. sp.
The body (in spirit) is very broad and much flattened, especially on the ventral
surface ; very blunt behind and tapering more gradually in front. The opening into
the peripharyngeal chamber is rather behind the middle of the ventral surface and
the genital aperture about halfway between it and the posterior end of the body.
The dorsal surface, in the living animal, was of a creamy white colour, with a narrow
dark brown median stripe and on each side of the stripe numerous close-set, narrow,
discontinuous, wavy longitudinal streaks of brown over all the remainder of the dorsal
surface. There is no record of any markings on the ventral surface of the
living animal.
I received three specimens of this worm in spirit from Professor Spencer, from
Croajingolong. The largest measures (in spirit) about 40 mm. in length and 8 mm.
in greatest breadth, and came from a height of 4000 feet on the top of the coast
ranges. I have named the species after Mr. A. H. S. Lucas, M.A., B.Sc.
Locality.—Croajingolong (Victoria).
7. Geoplana m‘mahont, n. sp.
Body (in spirit) strongly convex on the dorsal surface, flattened on the ventral,
tapering gradually towards the anterior end and much more suddenly towards the
posterior. Opening into the peripharyngeal chamber situate at about the junction of
the middle and posterior thirds of the body. Genital aperture nearer to the posterior
ON THE VICTORIAN LAND PLANARIANS. 75
end than to the opening of the peripharyngeal chamber. The ground colour of the
dorsal surface is yellow (pale or bright) and there are two strong brown stripes, one
on each side of the middle line, separated by a considerable interval and meeting at
the anterior and posterior ends. The ventral surface is cream-coloured. A specimen
in spirit measured 23 mm. in length and 4:5 mm. in greatest breadth.
Locality —M‘Mahon’s Creek, on the Upper Yarra (Victoria).
8. Geoplana alba, n. sp. (Figs. 10, 11).
Body broad and much flattened, strap-shaped; when at rest often crenated at the
edges, the crenations disappearing when the animal is fully extended; sub-triangular
in transverse section, the apex of the triangle with a very wide angle; tapering
very gradually in front and unusually suddenly behind, so that the hinder end of
the body is blunt, especially when the animal is at rest or in spirit. The opening of
the peripharyngeal chamber is near the junction of the middle and posterior thirds of
the body and the genital opening nearer to it than to the hinder end of the body.
There are no stripes and except for the pinkish anterior tip the entire body is of a
very pale and nearly uniform tint. Ihave noted specimens from M‘Mahon’s Creek
as “cream-coloured or white’ and specimens from Macedon as ‘ peach-coloured or
yellow flesh”’ and “brownish flesh-coloured all over, dorsally and ventrally, with
beautiful peach-coloured tip.’ Specimens from Macedon measured when crawling
about 65 mm. in length and 4 mm. in greatest breadth, but I have larger specimens
from M‘Mahon’s Creek and from Warragul.
Localities—M‘Mahon’s Creek, Warragul, Macedon, Croajingolong (Victoria).
9. Geoplana hoggit, n. sp. (Figs. 4, 5.)
Body almost elliptical in section but flattened beneath, long and narrow, tapering
gradually towards the anterior extremity and more suddenly towards the posterior.
Opening into the peripharyngeal chamber nearly in the middle of the ventral surface
and genital aperture about one-third of the distance between it and the posterior end
of the body. The ground colour of the dorsal surface is rather pale, translucent
yellow, or greenish yellow. There are four stripes, two on each side of the middle
line and extending throughout the entire length of the body. The two stripes
nearest the middle line are separated from one another by only a very narrow band
of ground colour and are usually of a green colour with a tinge of indigo; they may,
76 ON THE VICTORIAN LAND PLANARIANS.
however, be grey or greenish grey. The two remaining stripes, one on each side,
are separated by wider intervals from those nearest to the middle line and are of a
very dark brown, almost black colour. The relative breadth of the stripes appears to
vary a good deal. The anterior extremity of the body is of the usual brownish pink
colour, into which the stripes merge. The ventral surface is of a uniform yellow
colour, a little paler than the ground colour of the dorsal surface. The largest
specimen which I have seen alive was about 125 mm. long and 5 mm. in greatest
breadth when crawling (Fig. 5). Usually, however, specimens are only about 70 or
80 mm. long when crawling.
I have seen great numbers of this species under logs and stones at Macedon and
I have great pleasure in calling it after Mr. H. R. Hoge, on whose property it is
particularly abundant and who first brought it to me in a living condition.
Locality.—Macedon (Victoria).
10. Geoplana sugdent, n. sp. (Figs. 12, 13, 14).
Body very narrow and much elongated; approximately oval in section but
somewhat flattened ventrally, more nearly cylindrical than in any other species of
the genus with which I am acquainted; tapering gradually to each extremity.
Opening into the peripharyngeal chamber a little in front of the middle of the
ventral surface. Genital opening only a little behind the middle, separated from the
opening into the peripharyngeal chamber by an interval of about 10 mm. (in spirit)
and from the posterior end of the body by an interval of about 18mm. The colour
of the dorsal surface is bright canary yellow all over except the reddish brown
anterior tip; the ventral surface is of a paler yellow colour. There are no stripes
at all. Length when crawling about 70 mm., greatest breadth only about 2 mm.
This species is rather remarkable for its habit of wandering about in broad
daylight, to which I have already referred in my introductory remarks. I have much
pleasure in naming it after the Rev. E. H. Sugden, B.A., B.Sc., the Master of
Queen’s College in the University of Melbourne.
Locality.—Macedon (Victoria).
11. Geoplana mediolineata, un. sp. (Figs. 1, 2, 3, 3a).
Body long and narrow, oval in section but not very much flattened; tapering
very gradually towards the anterior and more suddenly towards the posterior end.
ON THE VICTORIAN LAND PLANARIANS. 77
Opening into the peripharyngeal chamber somewhat in front of the middle of the
ventral surface. Genital opening at about the junction of the middle and posterior
thirds of the body. The ground colour of the dorsal surface is bright canary yellow.
The stripes vary considerably in the extent to which they are developed. The most
typical condition appears to be that represented in Figs. 1 and 2. Here there is a
single, narrow, very dark brown or almost black, median dorsal stripe. At the
anterior end the brownish-pink tip is continued into two short, dusky brown stripes
on each side of the middle line; the one nearest the middle line being the longest.
At the posterior end also an ill-defined dusky stripe seems to be always present on
each side of the middle line. Specimens with stripes answering to the above
description are very abundant at Macedon. In a slight variety from Warburton and
Walhalla, of which I obtained six or seven specimens, the stripes are less developed,
even the median one being present only at the anterior end of the body, though
longer than the others. In a third variety, on the other hand, from Macedon, there
are three continuous stripes (Figs. 3, 3a), the innermost of the two lateral stripes on
each side extending all the way down the body and becoming continuous at the
posterior end with the rudimentary lateral stripes mentioned as existing there in the
typical form. The ventral surface is pale, nearly white. When crawling, examples
of this species are about 90 mm. in length and hardly 3 mm. in greatest breadth.
Localities —Macedon, Warburton, Walhalla (Victoria).
12. Geoplana quadrangulata, n. sp. (Figs. 6, 6a).
Body almost quadrangular in section but rounded off at the angles. Dorsal
surface a good deal broader than the ventral and connected therewith by inwardly
sloping lateral surfaces. Both the dorsal and ventral surfaces are flattened. The
opening into the peripharyngeal chamber is situate a little behind the middle of
the ventral surface, and the genital opening is about half way between it and the
posterior end. The dorsal surface is of a dark reddish brown colour, slightly mottled,
with a narrow median stripe of very dark brown, and a dark brown anterior tip. At
the junction of the dorsal with the lateral surfaces there is, in two out of three of my
specimens, a single row of small, pale spots, very distinct in spirit specimens. The
lateral surfaces are also reddish brown, but much paler than the dorsal surface. The
ventral surface is white, but owing to the slope of the sides of the body the latter
appear as a brown stripe on each side of a median white band (Fig. 6a). Length
when crawling about 27 mm., greatest breadth about 1.5 mm.
Locality.—Macedon (Victoria).
78 ON THE VICTORIAN LAND PLANARIANS.
13. Geoplana walhalle, n. sp.
This species agrees with Geoplana spenceri in shape, and in the uniform dark
olive green colour of the dorsal surface. 'The ventral surface, however, instead of
being blue, is of a light, speckled brown colour. The anterior tip is dark brown.
The opening of the peripharyngeal chamber is situate near the middle of the body,
but slightly in front, and the genital opening is about half way between it and the
posterior end, if anything a little nearer to the posterior end. Length of two
specimens when crawling about 37 and 50 mm. respectively.
I found two specimens of this Planarian at Walhalla, Gippsland, Victoria, one
crawling on the top of a stone (in the day-time), and one under a log.
Locality.—Walhalla (Victoria).
14. Geoplana fletcheri, n. sp. (Figs. 8, 9).
Body much flattened ventrally, convex dorsally ; a good deal broader behind than
in front ; tapering gradually to the anterior, and much more suddenly to the posterior
extremity. Opening into the peripharyngeal chamber a little behind the junction of
the iniddle and posterior thirds of the body. Genital opening somewhat nearer to the
posterior extremity of the body than to the opening into the peripharyngeal chamber,
and hence very near the posterior end. The ground colour of the dorsal surface is
canary yellow, not quite so bright as in G. sugdeni, with a tendency towards the
formation of two brown lateral stripes, continued backwards for some distance from
the brownish pink anterior tip. In one specimen (Fig. 9), the ground colour was
slightly mottled with brown, and, in addition to the two partial stripes already
mentioned at the anterior end, there was a faint, narrow, median, brownish stripe
running along the posterior half of the dorsal surface. The ventral surface is pale
yellow. Length when crawling about 60 mm., greatest breadth about 3 mm.
This species at first sight resembles G. sugdeni, but may be readily distinguished
by the general shape of the body, and the position of the apertures. I have named
the species after Mr. J. J. Fletcher, B.A., as a slight recognition of his valuable work
on the Australian land Planarians.
Locality.—Macedon (Victoria).
ON THE VICTORIAN LAND PLANARIANS. 79
Genus Rhynchodemus.
This genus is distinguished from the preceding principally by the possession
of only two, multicellular eyes, situate near the anterior extremity, and on the dorsal
surface. Although apparently not uncommon in New South Wales, five species being
described by Messrs. Fletcher and Hamilton (Joc. cit.), only a single specimen has as
yet been found in Victoria, and that in the Croajingolong district, not very far from
the New South Wales border. The species is, however, a new one.
15. Rhynchodemus victoria, nu. sp. (Figs. 15, 15a).
Body (in spirit) much flattened, especially on the ventral aspect, not tapering
evenly to the anterior extremity but with a slightly developed neck, which seems,
indeed, to be characteristic of the genus (vide Figs. 15, 15a). The opening into the
peripharyngeal chamber is very distinctly margined and situate a little behind the
middle of the ventral surface ; the genital opening is about half way between it and
the posterior end of the body. The eyes are very near the anterior extremity (Fig.
15, 2). Professor Spencer informs me that the principal colour of the dorsal surface
in the living animal was French grey or “‘elephant’s breath.’ At the anterior end, on
the dorsal surface, the following stripes are visible :—A narrow dark median stripe,
dying out about half way down the body and edged on each side by a creamy white
stripe of about the same width and dying out a little before the dark one. Just
above each lateral margin of the body is another narrow creamy white stripe, soon
dying out. The two white stripes of each side unite near the anterior extremity and
at their point of union the eye of that side is situated (Fig 15, ¢). The ventral
surface in the living animal was lighter grey, with a single, median, creamy white
stripe dying out just behind the genital aperture, and a similar stripe very near the
margin on each side also dying out towards the posterior end. Length in spirit
about 26 mm., greatest breadth about 3 mm.
I received a single spirit specimen of this interesting Planarian from Professor
Spencer, who collected it at Croajingolong. Professor Spencer supplied me with the
description of the colours of the living animal, but the arrangement of the stripes I
was obliged to make out in the preserved specimen.
Locality.—Croajingolong (Victoria).
80 ON THE VICTORIAN LAND PLANARIANS.
Derscrietion oF Puate VII.
Figures 1, 2.—Geoplana mediolineata, n. sp. Dorsal aspect. Fully extended.
Typical form. Painted from life; natural size. Macedon.
Figure 8.—Geoplana mediolineata, nu. sp. Dorsal aspect. Fully extended.
Variety with three complete stripes. Painted from life; slightly enlarged. Macedon.
Figure 8a.—Geoplana mediolineata, n. sp. Dorsal aspect. Anterior extremity
of the same specimen as that from which Figure 3 is taken; enlarged to show the
arrangement of the stripes. Painted from life.
Figures 4, 5.—Geoplana hoggii,n. sp. Dorsal aspect. Fully extended. Painted
from life; natural size. Macedon.
Figure 6.—Geoplana quadrangulata, nu. sp. Dorsal aspect. Fully extended.
Painted from life ; twice the natural size. Macedon.
Figure 6a.—Geoplana quadrangulata, n. sp. Ventral aspect. Not quite fully
extended. O, opening of peripharyngeal chamber ; g.a., genital aperture. Painted
from life ; twice the natural size (same specimen as in Fig. 6),
Figure 7.—Geoplana ade, n. sp. Dorsal aspect. Fully extended. Painted from
life; natural size. Macedon.
Figure 8.—Geoplana fletcheri, n. sp. Dorsal aspect. Pretty fully extended.
Painted from life ; slightly enlarged. Macedon.
Figure 9.—Geoplana fletcheri, n. sp. Slight variety. Dorsal aspect. Fully
extended. Painted from life; one and a half times the natural size. Macedon.
Figures 10, 11.—Geoplana alba, n. sp. Two specimens. Dorsal aspect. Pretty
fully extended. Painted from life. Natural size. Macedon.
Fieures 12, 18, 14.—Geoplana sugdeni, n. sp. Dorsal aspect. Figure 12
represents the worm at rest, coiled up with its anterior end in the middle, slightly
enlarged. Figure 13, partly extended, natural size. Figure 14, fully extended,
natural size. Painted from life from three specimens. Macedon.
Figure 15.—Rhynchodemus victorie, n. sp. Dorsal aspect; e, eye. Drawn from
a spirit specimen, twice the natural size. Croajingolong.
Figure 15a.—Rhynchodemus victorie, nu. sp. Ventral aspect of the same
specimen as represented in Figure 15. O, opening into the peripharyngeal chamber;
g.a., genital aperture.
Fig |
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Trans. Royal Society, lictorta 1890 Plate 7
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ARTICLE III.—Tur Evcatyrets or Grppsuanp, sy A. W. Howrrt, F.G.S., F.L.S.
(With Plates 8, 9, 10, 11, 12, 18, 14, 15, K(S)\s
(Read Thursday, July 10th, 1890).
CONTENTS.
I.—Inrropuction .. a ee Be ec . ao 83
Il.—Vanrertes or Typrs—GENERA Ee Be a 35 D0 84
III.-—Distrinution or Typzs .. ie 26 Ss a ac 104
4A.—As To AREAS... Se aa Ase se aie 104
B.—In Autirupr Apove Sea Lrven es on ee np 107
C.—As to GronocicaL Formation ee da 50 a 108
TV.—Inrnvence or Crimate anp GEoLoGicAL ForMATIOoN ON THE DisrripuTion or Typrs 108
V.—InFuvence or SerrnemMmNnt oN THE Eucanyprus Forrsts .. He oc 109
LIST OF EUCALYPTS OF GIPPSLAND.
RENANTHERA— PoRANTHERA— ORTHANTHERE—
EK. pauciflora. EK. leucoxylon. K. pulverulenta.
K. stellulata. K. melliodora. K. stuartiana.
K. amygdalina. EK. polyanthema. EK. viminalis.
E. piperita. KK. odorata. K. tereticornis.
K. eugenioides. EK. hemiphloia (albens). _E. gunnii.
EK. muelleriana. K. botryoides.
E. capitellata. EK. goniocalyx.
K. macrorhyncha. K. globulus.
EK. obliqua.
K. stricta.
I
sieberiana.
D
blo
THE EUCALYPTS OF GIPPSLAND.
TABLE OF BOTANICAL, ABORIGINAL* AND LOCAL NAMES OF THE GIPPSLAND RUCALYPTS.
| E. paucitlora 30 | Bundagra aD 36 | Mountain White Gum.
| E. stellulata ue | Yimbit | Black Sally, Muzzlewood.
Ei. amygdalina Chunchuka | Peppermint Gum.
3 (Gi) Katakatak or Yertchuk ..
» (e) Wang-ngarat
E. piperita | Yangura White Stringy- bark. |
E. obliqua Katakatak | Messmate.
Ii. capitellata Dumung Mountain or Red Stringy-bark.
i. muelleriana Yangura Yellow Stringy-bark.
E. macrorhyncha Katakatak or Yurokat Mountain Stringy-bark.
TPES) {White ron-batl, Wooly But, |
E. leucoxylon Yirik or Bwurawi Tron-bark. |
KE. melliodora Dargan | Yellow Box. |
| E. polyanthema Den or Dern | Red Box.
E. odorata. . Dargan | Yellow or Grey Box.
E. hemiphloia Den or Dern Grey Box.
KE. pulverulenta Bindirk Silver Leaf Stringy-bark.
E. stuartiana But-But | Apple Tree or Apple Box.
E, yviminalis (a) Binak | White Gum.
| * (vb) Cabbage Gum.
Ii. tereticornis Yuro.. Red Gum.
E. gunnii .. Gura-Binak Swamp Gum.
E. botryoides Binak Snowy R. Mahogany.
E. goniocalyx Baluk Spotted or Bastard Gum.
| E. globulus Baluk or Wang-ngarat} . Blue Gum.
I. eugenioides Yangura | White Stringy-bark.
* The majority of these native names are taken from the Muk-thang dialect. Muk-thang. or “excellent speech,”
was spoken by the Brabolung Kurnai, who lived on the Mitchell, Nicholson, and ‘'ambo Rivers.
+ Wang-ngara, in the Nulit dialect, spoken by the Brataua and Tatung, who dwelt between the Lakes and the
sea, and in South Gippsland; wang means “bark,” and ngara “‘a string” or ‘‘ tough,” hence ngarang ‘“‘a sinew.” The
application of such a term seems more appropriate to KE. amygdalina than to E. globulus, for the bark of the former is
extremely tough, and can be detached from the bole in long strips. The bark of E. globulus is not so tough, but at times
hangs from the tree in long strings, in reference to which, perhaps, the name is given.
+ Yuroka was the name given by the Krauatun Kurnai, who lived at the Snowy River and spoke Thang quai, or
“broad speech,” to the mountain stringy bark, E. macrorhyncha.
Tur Evucarypts or Grepsuanp, By A. W. Howitt, F.G.S., F.L.58.
INTRODUCTION,
So much has already been done by Baron von Mueller towards bringing into
order the numerous varieties of the recorded Eucalypts, that it only remains to work
out their local distributions, as well as the effects produced by soil and climate. This
attempt I have now made as regards the Gippsland Eucalypts.
The difficulties which have met me in this enquiry have been greater than I at
first anticipated, as it has necessitated a series of investigations, spread over a
number of years, and covering the greater part of Gippsland. Even now, after
examining the whole of the district more or less carefully, I find that in order to
complete my work with even a fair degree of accuracy, it would be necessary to again
visit certain localities, my earlier observations of which are wanting in that minute
examination which later investigations have shown to be absolutely necessary. I am
unable to revisit the localities referred to, and have therefore confined my remarks
more especially to Central and West Gippsland, touching the North-eastern part very
lightly, and scarcely referring to Kast Gippsland at all.
I regret that this should have been unavoidable, for it is in the country between
the Snowy River and Cape Howe, and south of the coast range, that one would most
probably find that the New South Wales species meet and mingle with those most
general in Gippsland,
It is my pleasing duty to thank Baron von Mueller most heartily for the unending
kindness with which he has, throughout the course of my enquiries, responded to
my constant requests for information, for the readiness with which he has
examined and named the collections which I forwarded to him, and thus resolved
difficulties which I met with, and for most kindly making available to me the
specimens in his museum. I am much indebted for the kindness with which Mr.
Luehmann has given me his valuable aid in comparing my samples with those of the
departmental herbarium, with which he is so fully acquainted.
To Dr. Wools I am under great obligations for valuable information as to New
South Wales species, and for samples of the same.
84 THE EUCALYPTS OF GIPPSLAND.
Also to My. J. E. Brown, F.L.S., of Adelaide, to whom I addressed myself for
information as to South Australian species.
Mr. John O’Rourke, of Woolgulmerang, Mr. David O’Rourke, of Buchan, and
Mr. 8. C. Holme, of Eagle Vale, have all most kindly made collections of the Eucalypts
srowing in their respective neighbourhoods.
Typrs oF KUCALYPTS GROWING IN GIPPSLAND.
RENANTHERE.—In the following notes on the Types of Hucalypts in Gippsland, I
have endeavoured to avoid repetition of the descriptions which have been
given by Baron von Mueller with such admirable clearness and precision in
the “ Eucalyptographia,” and desire only to add such particulars as have
presented themselves to me, or have special reference to the local occurrence
of these Eucalypts in Gippsland.
E. pauciflora.—This Euealypt is extremely constant in character, whether
found in small isolated colonies in the littoral tracts, as at Providence Ponds and
Morwell, or forming forests over large areas in the Gippsland Alps up to an elevation
of 5000ft., as on the Wonnongatta Plains, at Omeo, Woolgulmerang, and Delegate.
It appears to be essentially an Alpine species, yet able to maintain itself, to
some extent, in localities but little elevated above sea level.
E. stellulata—This is also an Alpine species, ascending almost if not quite to
the same elevation as E. pauciflora, but does not descend, according to my observa-
tions, lower than 7OOft. at Dargo and Ensay. No varieties occur, as far as my
observations go.
E. amygdalina.—This is one of the most variable, and at the same time,
naturally most widely spread of the Gippsland Eucalypts.
I have observed the following well-marked and constant varieties :—
(a). The ordinary narrow-leaved variety.—This is the type described and figured
in the ‘Eucalyptographia;” it generally grows throughout this district on all formations,
trom the sea level up to about 4500ft.
(b). The broad-leaved variety. In the mountains, and more especially in some
of the Plutonic and Metamorphic areas, as at Dargo and the Wentworth and Omeo,
there occurs a form of Amygdalina which is to some extent distinct from the typical
form referred to. I have not observed it at a lower elevation than 700ft. at Dargo,
THE EUCALYPTS OF GIPPSLAND. 85
and it grows upon Mount Livingstone at about 8000ft., which is, probably, near its
upper limit. According to my observation it does not exceed 100ft. in height, and is
more frequently under 50ft. Its bark is wrinkled, approaching to fibrous, and
persists up to the smaller branches.
The seedlings and young saplings have opposed sessile lanceolar leaves, which
are, however, much broader than the ordinary form, approaching at times pointed
ovate.
The leaves when scattered are broadly foliate and unequal-sided. The umbels,
buds, and flowers are those of the typical form, but the fruit is much larger, and
almost always ovate top-shaped, with a flat or slightly convex margin, and a brown
or brownish-red tint. The valves, as in the ordinary form, are sinall.
However much this tree resembles the ordinary form of this Eucalypt, it is
clearly to be distinguished from it, because the two varieties very commonly grow
together, each maintaining its own character.
In other places they may be found forming independent colonies. While the
common variety grows especially in the damp gullies and on the shady sides of the
ranges from the sea level up to about 4000ft., this form of Amygdalina (4) grows
preferably upon the sunny slopes from 7O0Uft. to 4000ft. (See Pl. 8.)
(c.) This variety is restricted to barren sand ridges of the littoral tracts, for
instance, in South Gippsland, between Merriman’s Creek and Warrigal Creek,
together with Banksia serrata, Acacia oxycedrus, and other sand-hill vegetation. It
does not exceed 30ft. in height, and has a marked drooping habit in its branches and
foliage, while the bark is wrinkled and persistent up to the small branchlets.
The form of the leaves distinguishes it from variety (a), for they are long and narrow
lanceolar or falcate, the venation is so little spreading as to resemble in some respects
that of KE. pauciflora or E. stellulata; the flowers and fruit are typical of E.
amygdalina, to which I have assigned the tree. (See Pl. 10, Figs. 6 to 12.)
(d.) This is the most widely-spread variety of E. amygdalina, and at the same
time that which departs most from the typical form. It grows most freely upon the
rather poor sandy and clay lands of the littoral tracts, but I have also observed it in
the mountains, for instance, where poor sandy tracts occur, as well as on the quartz
grits and conglomerates at Wild-horse Creek, Wentworth River, on the Upper Silurian
sediments, between Toongabbie and Walhalla, the Silurian sediment in the Tambo
Valley Road, the Upper Devonian formations of the Insolvent Track, the Devonian
porphyries at Gelantipy, and the Silurian formations at Delegate River.
It rarely grows more than 100ft. in height, but is generally a rather small tree,
often stunted. The bark classes it with the stringy barks, for it is fibrous and
86 THE EUCALYPTS OF GIPPSLAND.
persistent up to the smaller branches, somewhat resembling that of E. obliqua, but
thinner, more fissile, and lighter in colour. For roofing purposes the bark is worth-
less, and the timber of no value for splitting or sawing, having the soft, veiny
character of some types of Amygdalina.
The seedlings have opposed, narrow, lanceolar leaves, with, occasionally, tufts of
hairs and frequently wavy margins.
The opposed character of the leaves is not maintained beyond the first two or
three pairs, and the leaves then become ovate lanceolar, resembling in their pointed
and unequal-sided form those of E. obliqua, though rarely as large. They are thick
in consistence, of a dull green, and not shiny ; in the Jatter trait resembling those of
HK. sieberiana.
The umbels have numerous buds, with the typical form of E. amygdalina,
to which also the shape of the calyx tube, the short style, and depressed lid belong.
The fruit is ovate top-shaped, with a flat and slightly convex margin and small deltoid
valves.
The tree is found at a height of 100ft. above sea level at Merriman’s Creek and
the Bairnsdale to Buchan-road, to 2500ft. at the Upper Wentworth River. I have not
observed it on the mountain plateaux, even where they descend to the latter height,
and conclude that it is a littoral species which ascends the coast ranges. (See Pl. 9.)
(e.) This is the Wang-ngara* of the Gippsland blacks. It is found in the
eastern part of Gippsland, but, according to my observations, not so commonly as
the other varieties of this type. It grows along the rivers and streams, and in moist
valleys, where it takes the place of EK. viminalis (a). It has a smooth, tall, but
comparatively slender bole, with a scanty, often rather spreading, head, in which there
is frequently a marked absence of foliage. The bark is persistent, and wrinkled only
on the lower part of the bole, above which it becomes smooth and almost white. It
is of extreme toughness, whence the aboriginal name,
The seedlings and young saplings have sessile, rather long, lanceolar, opposed
leaves, resembling those of the normal Amygdalina, but which in the older trees
become narrow lanceolar-falcate, attenuated at the stalk and pointed. The venation
is rather indistinct, the marginal vein considerably removed, and the lateral veins
very longitudinal.
The umbels are on stalks as long or longer than the bud and stalklet, the stalklet
slender and longer than the bud, the lid small and depressed, with a slight point.
* This native name has a reference to the extreme toughness of the bark of this tree. Wang—a band; ngara is
connected with ngarang—a sinew.
THE EUCALYPTS OF GIPPSLAND. 87
Buds numerous, 3 to 20. Flowers normal. Fruit, ovate truncate, with slightly
contracted orifice, compressed rather narrow rim, and small weak valves.
Compared with samples, for which I am indebted to Dr. Wools, of Sydney, this
appears to be the White River gum of New South Wales, Eucalyptus radiata
(amygdalina). This seems to be one of those eastern forms of vegetation which are not
found any further to the westward than the Mitchell River, though this tree
individually does not extend beyond the Tambo River.
I have observed it at an elevation of 50ft. at Jimmie’s Point backwater, 200ft. at
the Tambo crossing, 300ft. at the Murrendel River, and at Wangrabell on the Genoa
in Kast Gippsland, at an elevation, judging from memory, of not more than 500ft.
(See Pl. 10, Figs. 1 to 5.)
(f) (or E. regnans).—This Eucalypt, though possessing a specific title, belongs to
K. amygdalina, and is less removed from the typical form than the varieties which I
have designated (d) and (e). Under its common name of black-butt it is found in
the western part of Gippsland on the Mesozoic carbonaceous formations, where it
especially flourishes. It reaches 300ft. in height, and according to the statements of
some observers* to 400ft. and above that height.
The young seedlings of this Eucalypt are at first like those of the typical
Amygdalina, but with somewhat broader, lanceolar, opposed leaves. These are soon
replaced by broadly lanceolar, scattered, unequal-sided, pointed leaves, very like those
of E. obliqua. The saplings so much resemble those of this Eucalypt in other
respects that at first sight they might be confused. The leaves of saplings of E.
regnans, however, are thinner in texture, rather lighter in tint, not so pointed nor
quite so unequal-sided as in E. obliqua.
The flowers and fruit connect this tree with E. amygdalina, and do not differ
from the typical form more than do those of the varieties (d) and (e).
It occurs over a wide area in South and Western Gippsland, chiefly on the
carbonaceous formations, together with E. obliqua, and E. globulus, from the sea
level up to about 1200ft. It is also found in the mountains, as at Walhalla, 1200ft.,
and at Tucker Creek, Wentworth River, 2500ft.
E. piperita and E. eugenioides.—In accordance with the list of typical Eucalypts,
given at p. 81, I should now describe E. piperita and eugenioides in sequence,
but it will be more convenient to speak of these together, since their near alliance
renders it necessary to point out the distinctions which may be drawn between them.
Eucalypts referable to these types grow more or less plentifully throughout the
whole of Gippsland, ascending from the sea level to 3000ft. above it.
* Amongst others Mr. J. Rollo, formerly a sawmiller at Yarragon.
88 THE EUCALYPTS OF GIPPSLAND.
In the “‘Kucalyptographia’’ Baron vonMueller says, in speaking of H. eugenioides,
that the distinctions between E. piperita and H. eugenioides are not yet clearly made
out, and that, perhaps, Bentham’s view, that both should be regarded as forms of one
species, may have to be adopted.* And, in referring to EH. Piperita he describes
its fruit as truncate or globular ovate, contracted at the narrow-edged orifice, with
valves perfectly enclosed.
This description accurately fits the sample from New South Wales, which, by the
courtesy of Baron von Muelier, I was enabled to examine in the collection of the
Department of Botany of Victoria. But it does not apply so aptly to the fruit of
E. piperita as Ihave found it in Gippsland. Occasionally I have met with examples
of this Eucalypt with truncate-ovate narrow-rimmed fruit, as, for instance, at the
Tambo River; but the general form of the fruit, especially in the western part of thé
district, is extremely variable, as will be seen from the following particulars.
In order to study the possible differences which might exist between E. piperita
and EK. eugenioides, I collected 26 samples from various parts of Gippsland, and
compared the fruit as one of the readiest means of reaching some definite conclusion ;
for I had already found that a diagnosis, based upon the general characteristics of the
aged trees, saplings, seedlings of the umbels, buds and flowers, led to no definite
result, except to raise very strong doubts as to the distinctions between the so-called
separate species being maintainable. Of the 26 samples, 3 had truncate-ovate
fruit, 16 truncate-globular, and 7 truncate-spheroidal.
In 10 examples the rim was sharp, and in 16 blunt, but the bluntuess of the rim
was not always associated with a much-truncated form of fruit and a wide orifice; and
in several cases I noticed on the same tree two forms of fruit, one truncate-ovate,
with a comparatively narrow sharp rim, and another which was much truncated and
more globular, and with a wide orifice.
The samples which I thus examined were, six from the country east of Bairns-
dale, seven from the neighbourhood of Toongabbie, and the remainder from isolated
places throughout the district, from the coast line to an elevation of 30U0ft., at the
sources of the Wentworth River.
If any conclusion is justified from these comparisons, I should say it is that the
greater number of samples having a truncate globular fruit, with a contracted aperture,
and a narrow sharp rim, are to be found in Eastern Gippsland, while those few, which,
in their much truncated fruit, with wider, blunt rims, and wider apertures, may be
assigned to Hugenioides, occur in the western parts, as, for instance, at Drouin and
the Agnes River.
* Eucalyptographia Tenth Decade.
THE EUCALYPTS OF GIPPSLAND. 89
It is in Western Gippsland, namely, near Drouin and at the Agnes River,
that I found those samples which were most near to the typical forms of KH. eugenioides,
and in which the valves were distinct, but yet only slightly exserted.
The conclusion to which I am led is that the suggestion thrown out by
Bentham, and noted by Baron von Mueller, is correct so far as concerns Gippsland,
namely, that E. piperita and HK. eugenioides are near varieties of the same type.
But when comparing the extreme forms found in South-western Gippsland with the
typical E. piperita from New South Wales, to which the examples from near the
Tambo River must be assigned, the conclusion is arrived at that the differences are
such as to have justified Dr. Wools in regarding E. piperita and HK. eugenioides,
as they occur in New South Wales, as distinct.
An examination also of numerous individual trees in different localities where
these Eucalypts flourish as one of the principal forest trees, for imstance, at
Merriman’s Creek, Toongabbie, the Macalister River, Budgee Budgee, the Went-
worth, and the Tambo, has shown me that in all these localities trees can be found
growing side by side, having fruit which is either truncate ovate-globular with a narrow
aperture, or compressed globular, much truncated, and with a wider aperture. The
seedlings and saplings growing in such localities, with the aged trees bearmg such
fruits as above referred to, are indistinguishable from each other, and I have come to
the conclusion that, so far as concerns Gippsland, possibly excepting perhaps its
extreme eastern part, there are no strong features to separate these EKucalypts into
Piperita and Eugenioides. If, however, samples are compared of the New South
Wales Piperita with the Gippsland samples, which agree with the definition of
Eugenioides, then it will be found that there are good and distinctive characteristics.
I have felt some difficulty in deciding to which type the Gippsland form should
be assigned, but on consideration I think that, in the majority of cases, the form of
the fruit and the seedings with ovate-crimped leaves, point to E. piperita rather than
E. eugenioides (*). I have, therefore, assigned them to that species, though there
are a few localities such as the Agnes River and at Drouin, where Kucalypts occur,
which can rightly be referred to H. eugenioides.
E. muelleriana.—This Eucalypt has an extensive range in the western half of
Gippsland. It is a littoral species, and is principally found between the Hoddle
Ranges and the sea coast. There it forms the bulk of the forest, growing upon sands
and sandy clays, from Monkey Creek, 20 miles from Sale, to Shady Creek, west of
Alberton, in an east and west direction, and from Carrajung southwards to the coast.
The area thus covered by this tree is about 300 square miles. It also occurs in lesser
* The seedlings of the New South Wales variety, for which I have to thank the courtesy of Dr. Wools, bear out
this view.
90 THE EUCALYPTS OF GIPPSLAND.
colonies on the ridges extending from the Tertiary tracts up to the high ranges forming
the spurs of the mountains. I have not observed it west of Toongabbie, where it ascends
the hills of Upper Silurian sediment for about 6 miles northwards to 1000ft.in elevation.
I have also seen it growing extensively on the hills across which the road, known as the
Insolvent Track, runs from the Stockyard to Cobannah Creek. ‘The formations here
are Upper Devonian, resting upon sediments which may be either Devonian or Upper
Silurian. Its range north and south in this locality is at least 25 miles, and its
highest elevation probably over 1200ft. I have noted a third locality where this tree
occurs under precisely similar conditions, extending northwards on the spurs of the
mountains, skirted by the Tambo Valley-road. ‘There it grows for several miles on
the Silurian sediments, northwards from the edge of the tertiary marine beds, and
reaches an elevation of at least 1000ft. I have little doubt that it will be found in the
intervening localities, and perhaps further to the eastward; but of this I have no
direct evidence.
It appears to grow to the largest size on the sands and sandy clays of South Gippsland,
where it forms most valuable forests. Its maximum height is 170ft. or thereabouts,
but more frequently 100ft. to 150ft. The bole is straight and rather massive, with mode-
rately spreading branches, and a fibrous and dark-grey bark, which is more deeply and
coarsely fissured than that of E. piperita, in fact, resembling the bark of E. capitellata,
where that species grows to a good size in favourable localities. The bark is
persistent up to the small boughs, which are more or less smooth. The leaves of the
aged trees are lanceolar, falcate, and more or less inequal-sided, rather dark green in
colour, equally shining on both sides, and usually three to five times as long as broad,
with a sharp apex.
The seedlings have narrow lanceolar opposed leaves of a dark green, shining but
paler on the under side. In the earlier stages they are frequently more or less beset
with small tufts of hairs. I have noticed that the leaves are still opposed in young
plants 2ft. to 3ft. in height. In young saplings, and those some feet in height, the
leaves are rather broad lanceolar, or ovate lanceolar in shape, less shiny on the lower
page, much dotted with transparent pores, and rather thin in substance. A marked
feature in the saplings of this Kucalypt, and one by which it can be distinguished
almost at a glance from those of other stringybarks, is that the broadly lanceolar and
pointed leaves have a tendency to assume a horizontal position, rather than a vertical
one, and this gives the saplings a shining appearance. ‘The stems of these saplings
and young trees are somewhat smoother than those of K. piperita, E. capitellata, or
K. macrorhyncha.
The umbels are usually solitary, and there is a marked tendency in this Eucalypt
for them to become strongly panniculated. The buds are from 3 to 12 in most
of the umbels. The stalk is frequently slightly flattened, and not much longer than
THE EUCALYPTS OF GIPPSLAND. 91
the buds, and the stalket nearly as long as the calyx tube, the lid semi-ovate to
hemispheric, smooth and occasionally slightly pointed, the stamens (rather sparse) are
large and reniform like those of EH. capitellata. Fruit almost hemispherical to
approaching semi-ovate, the rim flat or even slightly inverted, not wide, valves deltoid,
small, and inserted, or more rarely slightly prominent; four-valved, less frequently
three to five valved.
The timber of this tree is usually rather darker in tint than that of E. piperita.
It is fissile, free from gum veins or shakes, clear in the grain, and enjoying a great
reputation for durability. It is used for fencing and sawing, and, according to Mr.
Macalpine, of Tarraville, who has lived for 40 years in South Gippsland, fences are
still standing at Woranga with posts split from this Eucalypt, which have been from
30 to 40 years in the ground. I have myself observed posts of this timber standing
in fences at Woodside since 1859. The local name of this tree is ‘‘ yellow stringy-
bark.”
This Kuealypt, therefore, is to be placed between E. eugenioides and E. capitel-
lata. It resembles both, but the dissimilarities are more marked than the resemblances.
The characteristic distinctions are quite as constant as those which distinguish those
two species, and the occurrence of these species over so large an area, as well as in
independent lesser colonies, negatives the probability of its being a mere hybrid. The
distinctions which I have now noted as separating this Eucalypt from its nearest
congeners are such as to have led me early in my enquiry to regard it as a species
distinct from either. I hesitated, however, to definitely state this until I had an
opportunity of comparing my samples with those in the collection of the Government
Botanist. Having been enabled to do this through the courtesy of Baron von Mueller,
and having been most kindly aided in the comparison by Mr. Luehmann, I feel that,
being fortified by the opinion of our greatest authority, the venerable author of
_“« Euealyptographia,” I may establish this Hucalypt as an independent species under
the designation of Eucalyptus muelleriana.
E. capitellata.—This tree occurs in small colonies, scattered over a great part of
Gippsland. It cannot, strictly speaking, be called one of the littoral species, for I
have not seen it growing at a lower elevation than 500ft., as at Drouin West, 75Uft.
at Darlimurla, and above that elevation between Bruthen and Buchan. Thence it
ascends the mountains to 1200ft., near Walhalla, and 2000ft. on Mount Elizabeth,
near Noyang. It varies but little in character, yet in places the fruit approaches that
of E. macrorhyncha in the somewhat protruded vertex and to HK. muelleriana when
the margin is but slightly convex. I have referred to the peculiarity of its seedlings
in comparing those of the stringy barks generally, after speaking of those of E. obliqua.
E. macrorhyncha.—This is essentially a mountain species in Gippsland, and the
92, THE EUCALYPTS OF GIPPSLAND.
lowest elevation at which I have found it growing is at Glen Maggie, 200ft. above sea
level, upon Upper Silurian sediment. In the valley of the Macalister it grows
extensively, ascending the mountains near Mount Wellington to 3000ft. On the
Insolvent Track it appears at 1000ft., and thence extends through the mountains to
the sources of the Wentworth River, reaching an elevation of 3000ft. Similarly on
the Tambo River it commences at S00ft., near Noyang, and extends to 8000ft. at
Omeo. Further to the East at Turnback 1000ft., Jingalala 2500ft., and Bonang
3000ft; it is also found extensively on the Buchan, Snowy, and Deddick Rivers,
extending towards the high mountains in New South Wales. It grows especially
upon dry ranges, on Plutonic, Metamorphic, and Sedimentary formations of Silurian
and Devonian age. Ihave not observed it anywhere in the Tertiary tracts.
E. obliqua—This Eucalypt is principally found in the western and south-western
portions of Gippsland, where it, in many places, forms the whole of the forests, or is
in others mixed with E. goniocalyx, EH. viminalis, E. gunnii, and EH. globulus. It
appears to be essentially a littoral form, but ascends the mountains to considerable
elevations in the cool, shady, moist gullies on the southern slopes. For instance, in
the great Dividing Range, where the Nicholson River rises, E. obliqua follows up the
damp gullies on the south side, and forms part of the forest on the summit, together
with E. sieberiana (0), E. viminalis (a), and E. amygdalina(b). It occurs also in Kastern
Gippsland, as, for instance, at Buchan, Gelantipy, Bonang, and Bendoc. It varies but
little in character, although the form of the fruit is in some cases, as, for instance,
near Port Albert, in the sandy coast country, not quite so truncate-ovate as in the
typical forms, yet in all cases the peculiar unequal-sided ovate lanceolar or eyen-
cordate lanceolar and pointed form of the leaves always marks the saplings and large
seedlings from those of any other species.
Having now referred to the various types of the stringy-bark groups, it is con-
venient to mention the distinctions between their seedlings and saplings.
The seedlings of E. piperita, E. eugenioides, and EH. capitellata are beset on
stems and leaf-stalks with numerous tufts of hairs, which also line the edges of the
leaves.
The leaves themselves are more or less hairy, except in that form of H. capitellata
growing in the mountains, as at Osler’s Creek, where they are smooth. Those of E.
piperita and E. eugenioides are, at this stage, universally hairy. The seedling
leaves of both E. piperita, E. eugenioides, and EH. capitellata are at first ovate and
opposed, but in the former I have often observed them to be ovate-pointed, or even
lanceolar and smaller than those of H. capitellata, which are always ovate. In
neither species are the leaves shiny.
In E. macrorhyncha the seedlings are also more or less beset with tufts of hairs,
THE EUCALYPTS OF GIPPSLAND. 93
giving the stems a rough appearance, but ina less degree than the last-named species.
The leaves, at first opposed, are lanceolar in form, and slightly shiny. The
seedlings of E. muelleriana are as characteristic as those of any other species known
tome. ‘The stem and stalklets are slightly tufted with hairs, or are even smooth, the
leaves rather long, lanceolar, pointed, and opposed throughout, even in seedlings of a
foot or more in height, while their extremely shiny upper surface distinguishes this
form from all the other species of this group, being more marked even than in E.
obliqua, from which the persistent opposition of the leaves readily distinguishes it.
The seedlings of EB. obliqua are usually free from hairs, but are very commonly
warty, and the leaves are lanceolar, shining on one side, and thinner in texture than
those of E. macrorhyncha. They become scattered somewhat sooner than those of
E: macrorhyncha, and very much sooner than those of E. muelleriana, and soon
show the marked unequal-sidedness which is so characteristic of this tree.
The saplings of these Eucalypts may also readily be distinguished from each other.
Those of E. piperita remain rough up to 10ft. in height, the leaves then
become unequal-sided, ovate-lanceolar, or ovate-pointed, having the upper surface
slightly darker green, and more shining than the lower.
E. capitellata soon produces unequal-sided cordate leaves, fully twice the size of
those of E. piperita, and of a lighter shade of green, moreover, they hang more vertically,
and are consequently more equally tinted on both sides. In size, and the inequality of
the sides, they resemble the sapling leaves of E. obliqua, but are readily distin-
euished by not being attenuated as those of Obliqua are.
The saplings of E. muelleriana are distinguishable from all the others by having
opposed leaves, even up to two or three feet in height. The leaves are lanceolar and
unequal-sided, but in a less degree than others of the group. The upper page is
very shining, and the lower much duller and paler in hue. The apex is more or less
acute, and the lateral veins are more numerous and less spreading than in K.
capitellata. Even in saplings from 8ft. to 10ft. high, the leaves have a general
tendency to assume a horizontal position, thus producing a peculiar shining
appearance of their upper pages, which is characteristic of this tree when young.
The saplings of E. obliqua have somewhat large, very unequal-sided leaves,
broadly lanceolar, or even cordate, and always attenuated, thus being, as I have
pointed out, distinguished from EH. capitellata, whose sapling leaves are not
attenuated.
E. stricta.—The only locality in Gippsland in which this Eucalypt is found is,
as far as I am aware, on St. Pancras Peak, a rocky mountain between the Buchan and
Snowy Rivers, probably 4000ft. in height.
94 THE EUCALYPTS OF GIPPSLAND.
Baron von Mueller refers to it in the ‘‘ Eucalyptographia, Tenth Decade,” on
the authority of Dr. Wools and Rey. R. Collie, as growing on the elevated parts of
the Blue Mountains, particularly at some of the summits. In Gippsland it is locally
called ‘‘ Mallee,” from some fancied resemblance to the growth of that species,
growing in a number of slender saplings, from one thickened stump-like butt.
This variety agrees in almost all respects with that figured by Baron von Mueller,
and described by Sieber as E. rigida, except that the fruit, in my example, has no
prominent rim, and is rather hemispheric than truneate-ovate.
E. sieberiana.—There are two well-marked varieties of this Eucalypt which I
distinguish as (a) and (b). EH. sieberiana (a) occurs extensively on almost all
formations, in parts of Gippsland, up to an elevation of 3000ft., near Grant and the
sources of the Wentworth River, and at 4500ft. on Mount Wellington. It ‘is
characterised by a very rough deeply-fissured bark, which is persistent on the stem
and larger limbs only, hence its local names of ‘‘gum top,” or ‘silver top ;” its name,
‘white ironbark,” only refers to the colour of the wood. It grows especially on
the dry tops of the mountain ridges, and upon their sunny slopes.
E. sieberiana (b) occurs only in the mountains above the limit of E. sieberiana
(a), sometimes as low as 2500ft., but in other places, as on the summit of the Great
Dividing Range at the sources of the Livingstone Creek, where K. sieberiana (a)
ceases at 3000ft., and KE. sieberiana (6) commences at 3500ft. It extends on the
summits of the higher mountains, ¢.g., the Bowen Mountains, near Omeo, and the
Dargo High Plains to about 4500ft.
I estimate the height to which the tree attains as not exceeding 200ft. The bark
is fibrous, and rather like that of E. obliqua, but perhaps more flaky ; it is persistent
upon the bole, the upper part of which and the branches are smooth, but with much
detached bark, pendant from the forks and from the termination of the persistent
bark.
The seedlings of these two trees have much the same features, but that of the
(a) variety is much darker, and the stems more purple or reddish in tint. The fruit
also is usually smaller and darker, but otherwise no marked difference can be observed
between the seedlings of these varieties.
The timber of the (5) variety is of a light colour, long in grain, and remarkably
fissile, yet elastic. It is not a heavy wood, and it seems to me should be valuable for
many purposes. About 20 years ago, at Omeo, I made a set of swingle-bars for a
four-horse team from this timber, which stood work remarkably well, and one of
which is still in existence after much hard usage. I have also seen palings split from
this tree over 6ft. in length, which were so clean, that after being planed up, were
used successfully as weatherboards. Locally the tree is called ‘‘ Woolly-butt,” from
the character of the bark, and also ‘‘ Mountain Ash.”
THE EUCALYPTS OF GIPPSLAND. 95
E. odorata—This tree has in many respects a superficial resemblance to EK.
melliodora, with which it was for a long time locally confounded in Gippsland.
Of late it has received the local name of ‘‘ Grey-box”’ from the splitters and
sawmillers.
It grows principally on the Miocene limestones in the littoral tracts of North
Gippsland.
The difference between KE. odorata and E. melliodora was long apparent to me,
from a careful comparison of the trees growing in the Mitchell River district, and
especially from distinctions which have been apparent to the timber men there.
The wood of this Eucalypt is much browner in colour than that of E. melliodora,
and while the timber of the latter can very rarely be split into posts or rails, that
of the former, although it is difficult to split ‘‘ on the quarter,” is, when once the log
is opened, ‘‘ backed off” with great ease. The principal differences upon which a
rapid diagnosis may be made lie in the greatly superior height of E. odorata, in
its freer growth, the rhytiphloious bark, the smooth upper portion of the stem and
limbs, and the somewhat larger fruit, with a narrow compressed rim, and more deeply
sunk orifice. Finally, the outer stamens are all provided with fertile anthers, while
those of E. melliodora are anantherous.
The timber of this tree is most durable, and is one of the most serviceable of
the Eucalypts of Victoria, especially for work which is exposed to damp.
This tree grows to 200ft., or in exceptional cases to perhaps 250ft. in height.
I have observed a small colony of E. odorata growing in South Gippsland, near
Four-mile Creek. The occurrence of this tree in the Miocene limestones of North
Gippsland falls in with the statement made by Baron von Mueller that it occurs
upon limestone areas at St. Vincent’s Gulf.
E. leucoxylon.—This tree does not form forests in Gippsland, as in other parts of
Victoria, but occurs scattered over a wide extent of country, from sea level up to
2000ft. It grows upon various formations, as, for instance, at Toongabbie, on recent
Alluviums, Tertiary clays, and Upper Silurian. At Bairnsdale, upon Miocene and later
Tertiary beds; at Glen Maggie, upon Upper Silurian sandstone ; at Upper Freestone
Creek, upon Upper Devonian conglomerates; at Noyang, upon Paleozoic Plutonic
rocks ; and near Buchan, on Tertiary sands and clays.
I have not observed it further to the westward than Toongabbie, and it varies
but little, if at all, in character throughout Gippsland.
E. melliodora.—This is also of wide distribution, from about Traralgon, in South
Gippsland, to Jingalala, in North-east Gippsland, and at all elevations up to 2000ft.,
and on all formations. Its characteristics are also constant.
96 THE BUCALYPTS OF GIPPSLAND.
E. polvanthema.—Of this type there are two varieties, which, however, are not
sufficiently marked to justify me in separating them, as 1 have done in other cases.
Where it occurs in the littoral districts, as, for instance, at the Lakes’ Entrance, or
river flats at Heyfield or Bruthen, it has full foliage of a rather dark green colour,
and the leaves somewhat thin in texture. The tree grows to some size, but in many
cases, as Baron von Mueller has already pointed out, become so hollow as to form a
mere shell.
The second variety is found in the hill country, and ascends from about 100ft.
above sea level, as at Heyfield, to an elevation of 2000ft., as at the Wellington River and
at the Tambo River (Fainting Range). This mountain variety is a much smaller
tree than the lowland form ; the leaves are thicker in texture, frequently pruimous, or
even mealy. At first sight the tree resembles somewhat EH. hemiphloia (variety
Albens) in its bark, and ash coloured, and sometimes rather lengthened ovate leaves.
But it is readily distinguished by the form of the buds, by the outer filaments being
anantherous, and by the fruit. The seedlings and young saplings of both have much
in common.
E. hemiphloia—This species is extensively represented in Gippsland as a
mountain form. It occurs, for instance, in the valley of the Tambo River, north of
Fainting Range, where it forms the principal part of the forest, from about 750ft. at
Numlamungie to 2500ft. at Tongeo Gap. It is found at Turnback, at the Snowy
River, at Deddick, and, more rarely, at Tubbut.
Its characteristics accord entirely with the diagnosis given in the ‘‘ Kucalypto-
graphia,” with the exception that the umbels are formed by buds of comparatively
large size. The fruit is proportionately large. The bark, also, extends frequently far
up the branches, so that when the leaves are not markedly elongated, this tree
resembles, as I have already said, at first sight, the mountain form of H.
polyanthema. Yet, so far as I have observed, the two species are sharply marked off
from each other.
This form of E. hemiphloia appears to me to be that variety called E. albens.
E. pulverulenta is found in many places in Gippsland, though it nowhere forms
the bulk of the forests, or it is found scattered widely among other Eucalypts. In
South Gippsland, and near Boolarra, it grows to a maximum height of under 50ft., but
is of a smaller growth in other parts, as, for instance, at Osler’s Creek, Providence
Ponds, and Buchan. The peculiarity of this variety, as found in Gippsland, to which
Baron von Mueller has drawn attention in his ‘“ Kucalyptographia,” lies in the
lanceolar or falcate form of the opposed leaves of the aged trees, the ovate or
rounded leaves being confined to the saplings, or young trees.
THE EUCALYPTS OF GIPPSLAND. 97
This tree is worthless for timber, and I have never seen it used for any purpose
except for fuel, in default of better. The local name, “silver-leaf stringy-bark,”
refers to the pulverulent or ashy character of the foliage, and to its fibrous bark.
Tt occurs from near sea level up to an elevation of 700ft. at Darlimurla, nearly
the same at Buchan, and 1000ft. at Osler’s Creek. (See Pl. 15, Figs. 1 to 4, 9, 10,
13, 32, 33, 34.)
A few other remarks remain to be made when I compare E. pulverulenta,
HK. stuartiana, and the mountain form of E. viminalis.
E. stuartiana.—This species is well marked, and is one of the most persistent in
character of any of the Eucalypts of Gippsland. Wherever I haye seen it I have
found it to be a tree with somewhat large and spreading limbs, with a scaly, wrinkled
bark, which is persistent up to the small branches.
The wood of this tree is valueless for splitting, sawing, and even for fuel. Its
general appearance has caused it to be confused with E. hemiphloia, under the
designation of ‘ white box,’’ under which name I have known it to be cut for sale, at
least at one sawmill in Gippsland. It is found growing over almost the whole of the
district, from a little above sea Jevel up to a considerable elevation in the mountains,
as, for instance, up to 2300ft. on the track from Dargo to Omeo. It is found on all
formations, but I have observed it especially on the Tertiary clays of South Gippsland,
and on the Metamorphic and Plutonic areas of Tubbut, Dargo, and Jingallala.
FE. viminalis——My observation has shown me that there are at least three
Kuealypts, which may be assigned to the type of Eucalyptus viminalis.
(a.) The typical form of K. viminalis in Gippsland is the so-called ‘‘ white
gum,” or ‘‘river gum,” which grows along the immediate course of streams. It
may, for instance, be seen along the Glengarry, Thompson, Mitchell, and other rivers.
It accords well with the description of the species given in the ‘‘ Kuealypto-
graphia,’ hence I need not refer to it further here, more especially as I shall have
to draw attention to it in distinguishing the varieties.
The typical form grows from near sea level to at least 4000ft., as on some small
streams rising in Mount Livingstone. It is especially found following the river
courses, and ascends to its highest elevation without extending to the hills on either
side. (See Pl. 15, Figs. 23 to 31; Pl. 14, Figs. 7, 8, 9.)
(o.) This Eucalypt is sub-alpine, and appears some 7OOft. above sea level, as at
Dargo and Noyang, but somewhat higher on the Wellington River, and at Gelantipy.
It also grows about Morwell, where, together with EK. pauciflora, it forms part of the
98 THE EUCALYPTS OF GIPPSLAND.
forest, just as it does throughout the alpine and sub-alpine localities, which are its
special habitat. In the highest tracts, as at Dargo High Plains, it grows to a height
of over 100ft. It has a rather massive bole, wlth moderately spreading limbs, and
fairly full foliage. ‘The bark is smooth and very white, excepting near the ground,
where it more or less persists; it has frequently, when about to dehiss, a decided
‘‘coppery” tint upon the bole and limbs. The wood is somewhat soft, not very
fissile, and resembles that of the lowland form, except that it has a more reddish or
pinkish tint. The leaves are lanceolar, slightly faleate, and more or less attenuated
at the stalk, the marginal vein is usually but slightly removed, the lateral veins
numerous, not very marked, and inclined at about an angle of 40dee. with the midrib.
The umbels are axillary or solitary, and the stalklets about as long as the buds. The
buds are ovate, most commonly three in number, and arranged in the cruciform
manner so characteristic of E. viminalis. The lid is semi-ovate, smooth, and
occasionally pointed, but is not mamillated, as is the case with KE. stuartiana. Judging
by the examples which I have examined, I think that the buds of this variety are
more broadly ovate, have shorter stalks, and more rounded lids than those of the
jowland form. The fruit is semi-ovate, with a somewhat wide and more or less
convex margin, and with rather strong deltoid and protruding valves.
These characteristics are those of the typical E. viminalis, but the seedlings and
young saplings have peculiarities which raise doubts whether indeed this Eucalypt
should not rather be referred to HE. stuartiana.
The seedlings have round or ovate opposed leaves, which are closely sessile,
rarely they are ternary in verticels, the stems as well as the leaves are mealy, and
thus resemble very strongly, as also in other respects, the young plants of EK.
pulverulenta as it grows in Gippsland, rather than those of KE. stuartiana.
Even when as high as 8ft. or 10ft., the saplings still have pulverulent leaves of
an ovate form and opposed position, and the grown trees themselves occasionally
show a reversion to this structure at the ends of their pendant branches. In some
localities, as for instance at Dargo, this tree grows together with K. stuartiana, the
latter being in its typical form. When the seedlings and saplings of these trees are
thus compared, those of HE. stuartiana are found to be much less mealy, to be of
thicker consistence, and more pointed than those of this variety of E. viminalis, in
which the opposed condition of the leaves is continued much longer than in E.
stuartiana. There can be no doubt that E. viminalis and E. stuartiana ave nearly
allied, and it becomes necessary now to enquire to which of these this Eucalypt
stands nearest. EK. viminalis differs from E. stuartiana, as I have observed them
to be in Gippsland, by having much smaller limbs as compared to the bole. The
leaves of the former are smaller, as a rule, more attenuated at the stalk, of thinner
THE EUCALYPTS OF GIPPSLAND. 99
“consistence and lighter colour. The marginal vein is less distant, the lateral veins
more numerous, nearer together, and more obscure. The angle formed by the lateral
yeins, with the direction of the midrib, is greater in KE. viminalis than in E.
stuartiana.
The mean of a considerable number of measurements gave 10deg. more for
the former than the latter. In the greater number of cases the umbels of EK.
viminalis are three in number, arranged in a cruciform manner, while those of
E. stuartiana are more numerous and not so arranged. The buds of EK. viminalis
are more ovate, with a rounded, or at most, minutely pointed lid; those of
E. stuartiana are distinctly pointed, or even mammelated. The stalklets of E.
viminalis are, as a rule, shorter, particularly in the mountain form, although in
some cases those of the lowland forms are as long, or longer, than those of EK.
stuartiana.
The fruit of E. viminalis is more ovate than that of E. stuartiana, with a rim,
which is always more or less convex, or frequently strongly so.
Especially is this the case in the lowland form along the river courses, but is
less so in the mountain form.
In the mountain form the fruit is usually more hemispherical than in the
lowland variety, with a convex vertex, and the valves somewhat weaker, although
protruding. It is in those examples which grow at the highest altitudes, as, for
instance, the Dargo High Plains, 4500ft., that I have observed the form of the fruit to
resemble that of E. stuartiana, and in these the fruit is semi-ovate, the rim not very
wide, and the valves, although exserted on the whole, having their insertions placed
a little below the level of the rim. Yet, on examining the fruit from a number of
trees, I observed that there was a considerable variation in that of the same tree.
While some agree with this description, others have the protruding convex rim and
exserted valves of the typical E. viminalis. (See Pl. 15, Figs. 14 to 22; Pie
Figs. 1 to 6, 10 to 18.)
In the variety which I am now considering, it must be noted that the habit of
growth, and the character of the wood, is that of E. viminalis, and not of EK. stuartiana.
Although the cortical character is not of much value, the character of its bark places
it among the Leiophloiz.
This tree is the manna-producing Eucalypt of the mountain country. The
manna is produced as plentifully, in the same manner and of the same kind, as that
produced by the typical EK. viminalis.*
* When travelling through the Morwell district, where this tree forms part of the forest, some school children,
whom I requested to point out the ‘“ manna gum,” indicated this tree, saying that in December the ground under
the tree was white with manna.
100 THE EUCALYPTS OF GIPPSLAND.
I must note, in this connection, however, that I have found small quantities of
manna indistinguishable from that of E. viminalis, either by appearance or taste,
attached to slight injuries on the leaves of saplings of KH. stuartiana at Toongabbie.
The differences between this Eucalypt and the typical form of E. viminalis lie
almost entirely in the form of the leaves of seedlings and young plants, and in their
pulverulent character. In general appearance these certainly resemble the young
plants of Stuartiana, but even more those of EK. pulverulenta, as found in
Gippsland.
Having to choose between EH. viminalis and E. stuartiana, I have, after weighing
all these considerations, assigned it to the former, notwithstanding the strong
presumption which arises to the contrary from the extreme departure of its seedlings
from the typical form.
It seems not only to connect E. viminalis and EK. stuartiana, and thus to
strengthen the alliance which, as Baron von Mueller points out, exists between these
Kucalypts, but also to connect these two with E. pulverulenta, in some variety of
which I have observed the fruit to have a protruding margin and exserted valves. *
(c.) A somewhat peculiar form of E. viminalis grows between Toongabbie and
Walhalla, from about 1000ft. to 2500ft. above sea level. It does not exceed 100ft. in
height. The bark is somewhat rugged, and persists over the bole, but on the
branches is smooth and of a reddish brown tint, the foliage is plentiful, and of a
somewhat ashy-grey tint. The leaves are Janceolar to falcate, the veins rather
indistinct, but agree with those of EK. viminalis. The umbels, buds, blossoms, and
fruit are also of this type, but with this difference, that the umbels have numerous
buds, and only rarely three arranged in a cruciform manner.
The timber of this tree is of no use except as fuel. I have not observed it
erowing in any other locality.
E. tereticornis.—This tree, the well-known ‘“‘Red-gum” of Gippsland, is
essentially a littoral species.
It grows mainly on the recent alluviums, river flats, ancient lake basins,
and on the lower terraces of the Tertiary formations, up to an elevation of
150ft. or more, rarely 200ft., above sea level. The western limit is about Traralgon,
and it extends eastwards, almost to Buchan, and northwards to Glen Maggie,
Glenalladale, and Bruthen.
* The difference in the young plants of the lowland and mountain forms of this Eucalypt are little, if any, greater
than those which I observed exist in the young forms of the two varieties of E. leuacoxylon,which grow in the neighbourhood
of Heathcote.
The smooth-barked variety, locally known as spotted box has, in its young form ovate, opposed, somewhat mealy
leaves. The rough-barked form—the Ironbark—has opposed leaves only in very young seedlings.
THE EUCALYPTS OF GIPPSLAND. 101
In only one instance have I found it growmg in the mountains, viz., at
Glen Falloch, where it occupies a basin of soft shale of Upper Devonian age, from
700ft. up to 1500ft. above sea level. The soil of this basin, which is derived from
the soft shales, resembles that of the lower districts where E. tereticornis thrives
best.
The slight distinctions which separate it from EH. rostrata are constant throughout
Gippsland, although the terete form of the lid varies ; some forms being much more
attenuated than others. Yet in all the characteristic arrangement of the anthers is
constant.
E. gunniu.—tThis type is very widely spread over Gippsland, not only in horizontal
range, but also in elevation above the sea level. I have observed a lowland and a
highland form, and each of them has a dwarf variety.
Lowland form (a).—This form has been fully dealt with by Baron von Mueller,
and I have only to add that it occurs throughout the littoral country. In the damp
climate of West and South-Western Gippsland it grows to a considerable height, say
150ft., with a straight clear bole, and was there cut by saw-millers, in one instance
at any rate, as ‘‘blue-gum.’”’ In Central and Eastern Gippsland it does not grow to
so great a height, but maintains its other characteristics.
(6).—This dwarf variety grows in poor, boggy country in the low-lying tracts, but
also occurs in the drier hills at Foster. It usually does not grow higher than 4ft. to
5ft., but at Foster it is found from 18ft. to 20ft. in height. The bark is smooth in
texture and ashy grey in colour, which becomes lighter in the upper branches.
Generally, when in its dwarf form it has a large butt level with the ground of several
feet in diameter, from which rise numerous shoots.
In the dwarf form the leaves (excepting in the upper shoots) are somewhat
broadly ovate, and are opposed and sessile. Thetexture is thick and leathery, of a
dull, rather dark green colour. In the taller examples the leaves become scattered,
ovate lanceolar, somewhat attenuated at the stalk, and accuminate. They are
equilateral, slightly shining, and of a rather brighter tint than the sessile leaves, and
have the marginal vein distinctly removed, the lateral veins numerous and rather
spreading. Very often the terminal leaves are opposed.
This Eucalypt flowers and fruits when in a completely dwarf state. The umbels
are mostly axilliary, and of a bright yellow to orange colour, as are also the stalks and
young shoots, ‘The stalklet is angular and wrinkled, sometimes rounded, about
twice as long as the sessile buds, which are 3 to 7, and much crowded together. The
fruit sessile in clusters of 3 to 7, semi-ovate, margin slightly compressed, valves small,
not exserted, stalk slightly flattened.
102 THE EUCALYPTS OF GIPPSLAND.
The Tall Mountain Form (c).—This much resembles some of the lowland varieties,
which grow upon dry tracts of land; but the leaves are shorter, more ovate, smoother,
thicker in consistence, and rarely have the wavy margin which is characteristic of the
tall lowland form.
Its lower limit is probably about 600ft. above the sea level, but I think it possible
that the tall lowland form intermingles with it at that elevation, or less.
Dwarf Highland Form (d).—I have observed this form of EK. gunnii growing
extensively in the swampy flats at the source of the main branch of the Livingstone
Creek, at an elevation of about 3000ft.
The description given of the dwarf lowland form applies in many respects to this
also. It does not exceed 2O0ft. in height; the bark is smooth, persistent at the butt,
and smooth and greenish on the branches. The leaves are ovate, and at first
opposed and sessile, or nearly so, finally scattered, of a dull green. The umbels are
axillary, or solitary, of sessile crowded buds. Fruit, semi-ovate rim, rather broad
and slightly convex, the valves barely exserted. The young twigs and umbels are all
slightly mealy. This form, however, differs from the corresponding lowland one in
the leaves being shorter and broader when they become scattered, in the darker
green of the foliage, and the smaller size and mealy character of the buds. The fruit
also is of a smaller size.
E. botryoides.—This, compared with the other species of Gippsland, is one of
the eastern types of Eucalypts. It spreads along the coast line westward to the sandy
tracts known as Bole Bole, between the Gippsland Lakes and the sea, where it
becomes stunted and worthless as a timber tree, though on the northern shores of
the lakes, at Lake Tyers, and at the Snowy River, it grows to a large size, and is a
valuable timber. At the latter locality it is known as ‘‘ mahogany.”
On the northern shore of Lake King it is to be found growing to about 100ft. in
height, and I have observed a small colony at the very extreme end of the long narrow
delta through which the Mitchell River enters Lake King. ‘This is probably the most
westerly point at which this tree grows, and, with Rubus rosifolius, which is found
in one gully at Mount Taylor, is marked the western limit of the east coast flora in
Gippsland.
E. goniocalyx las a wide range in Gippsland, especially in the western parts.
It grows well in the deep shady gullies of the southern slopes of the mountains, where
it reaches some 2O00ft. to 250ft. in height, with a tall, massive bole. In its typical
form it occurs in the valley of the Thompson River, on the Upper Wellington, near
Grant, on the southern slope of Fainting Range, at Gelantipy, and elsewhere, up to
4000ft. above sea level in favourable localities.
THE EUCALYPTS OF GIPPSLAND. 103
It is very commonly termed ‘‘Blue-gum,” and as such, has, to my knowledge
been cut by sawmillers. At Walhalla it is used in preference to H. sieberiana or EK.
capitellata, as being the best procurable in the district for props m the mines, and, so
far as my experience goes, may be placed after EK. globulus as a useful timber tree
for work that is not placed in or on the ground—as framing or planking.
The typical form of E. goniocalyx seems not to be able to cross from the cool
southern slopes to the warmer and drier northern sides, but there is found in such
places a peculiar divergent form.
On the south side of Fainting Range HE. goniocalyx ascends to the summit at
about 2000ft., while on the northern face of the mountain, and at about 2500ft. above
sea level, there is a peculiar variety of this type. The seedlings and young plants
have opposed, ovate, sessile leaves, of a rather light tint of green, not shining, and
without the peculiar and characteristic rank odour of the leaves of the young plant of
the common form.
The tree is usually under 50ft. in height, often with a short bole, and scanty
limbs and top. The bark is distinctly wrinkled, and the branches only are smooth.
The leaves are finally scattered, long lanceolar or falcate Janceolar, and more attenuate
at the stalk than is usually the case in the typical form. The marginal veins are
somewhat removed, and the lateral ones slightly spreading. The umbels, flowers,
and fruit accord well with the general character of this Mucalypt.
E. globulus is much more widely distributed in Gippsland than might be
expected. It varies but little, and the variation is, 1 think, principally in the more
or less smoothness of the fruit. It grows especially well in the cool and moist
districts of the Strezlecki and Hoddle Ranges, more particularly on the southern
slopes, to the very shore of Corner Inlet. It is more rarely found on the northern
side of these mountains, as also scantily on the foot-hills of the main ranges—as, for
instance, about Toongabbie, Freestone Creek, and the Tambo River. It grows in the
gullies on the north-side of Lake King, near Jimmie’s Point, at Lake Tyers, and in
other places towards the eastern boundary of the colony. In the Gippsland
mountains, it is found here and there in isolated colonies, even ascending to 4000ft.
on the north-eastern slopes of Mount Livingstone.
104 THE EUCALYPTS OF GIPPSLAND.
DiIsTRIBUTION OF THE Eucatypts.
In deciding how the various types of Eucalypts are distributed throughout
Gippsland, the first point which attracts notice is that the greater number fall into
one or the other of two groups. One group essentially belonging to the mountains,
and the other to the lowlands. Besides these, there are a few which are spread
more or less over the whole district, whether littoral or alpine.
Certain types, peculiar to the littoral tracts, are found, which thence extend up
the river valleys, and moist, cool gullies at their sources on the southern slopes of
the mountains. They then cease, and are replaced on the warm, dry northern
slopes, and on the more elevated, cool, sub-alpine tracts, either by other types, or by
other varieties of the littoral forms.
A few instances will make my meaning more clear. I commence with the best-
marked westerly example, that of the valley of the Macalister. In it E. viminalis, E.
piperita, E. obliqua, and E. goniocalyx afford instances.
The lowland form of EK. viminalis, which is the ‘river-gum” of Gippsland,
follows up the Macalister River, growing only on the banks, or on the flooded flats,
to an elevation of 3000ft. at Lake Karng, and at an elevation of 1000ft. at the
Wellington River. The mountain form of E. viminalis (b) oceurs on the open, grassy
spurs flanking the river, and then is found here and there, until it becomes plentiful
on the table-land, through which flows the Caledonia River.
E. piperita, which forms extensive forests along the foot of the hills, follows up
the valley of the Macalister, growing on the flats above flood level, but gradually
ceases on approaching Glen Falloch.
EK. macrorhyncha commences at Glen Maggie, on the dry Silurian ridges, extends
all along the stony ranges flanking the valley, and reaches an altitude of about 3000ft.
on the track leading up from the Wellington to the Snowy Plains. In the mountain
gullies, having a southerly aspect, which descend from the mountain at Lake Karng
towards the Wellington River, E. piperita again appears, together with EH. viminalis
(a), and K. goniocalyx, which all reach, in this spot, an altitude of 3500ft. above
sea level.
The high mountain ranges to the westward of the Macalister, and the still
higher alpine tracts to the west of the Mitchell, above the Dargo junction, and again
those to the west of the Snowy River, above its junction with the Buchan,
precipitate the westerly rains, and divert the rainfall from the valleys on the eastern
side of the mountains.
THE EUCALYPTS OF GIPPSLAND. 105
Somewhat similar seems to be the effect produced by the coast ranges upon the
sub-alpine plateau of Omeo and the Dargo basin.
At the sources of Cobbanah Creek, and at the Budgee Budgee Mount, similar
observations can be made.
E. piperita and E. obliqua ascend the sources of Granite Creek to the saddle
separating it from Bulgaback Creek at 500ft.
They then cease, and on the dry north side are replaced by E. macrorhyncha,
which then extends as the sole stringy-bark to about 1200ft. in the mountains leading
to Grant. On following these mountains, however, HE. obliqua re-appears in the
south gullies of Mount Ewing, at an elevation of 4000ft., which is its limit in these
mountains.
In proceeding from Dargo to Omeo, by way of the Wentworth River sources,
instructive observations can be made as to the distribution of the different types of
Kucalypts.
In Dargo itself, which is essentially one of the mountain areas, E. viminalis (0)
occurs at its lowest limit of 700ft., growing plentifully over the open grassy hills of
Plutonic rocks. EH. viminalis (a) occurs in the river courses, following them up from
the littoral tracts. At the source of Teapot Creek, near Mount Steve, is to be found
an outlying colony of E. amygdalina (f) the Black-butt, of South Gippsland, together
with E. piperita and EK. obliqua at an elevation of 2500ft. The summit of the ridge
is crested by E. sieberiana (a), and in the creeks falling into the Wentworth River,
the Eucalypts are essentially lowland forms, with the examples of EK. amygdalinas
(a) and (6) which occur locally at Pheasant Creek. These lowland forms, for
instance, H. viminalis (a), E. amygdalina (a), E. piperita, KE. obliqua, K. melliodora,
E. polyanthema, E. stuartiana, follow up the sources of the Wentworth River towards
the Great Dividing Range.
On the range along which runs the Wentworth track to Omeo, I observed that
K. melliodora ceased at Wild Horse Creek at about 1500ft., EH. macrorhyncha at
2400It., KE. amygdalina (b), which here grows plentifully at one place, and at an unusual
elevation of 2400ft. HE. polyanthema gradually decreases in number, diminishes
in size, until at 2300ft. it ceases with a few dwarfed trees. At the same place EH.
stuartiana also ceases abruptly on the south side of the ridge, and with a diminution
of size.
At 3000ft. K. piperita disappears at the extreme source of Wild Horse Creek,
without any change of size. At the same point E. viminalis (a) also ceases. It
appears that the elevation above the sea level has less to do with the disappearance of
these trees than the change of climate from the cool moist gullies on the southern
slopes to the cold yet drier northern side of mountains.
106 THE EUCALYPTS OF GIPPSLAND.
On the great Divide itself HK. sieberiana (a) ends abruptly at 3000ft., and BE.
sieberiana (b) commences at 3500ft., here also grows E. obliqua to great size, but
ceases at 3100ft. at the descent into the Upper Livingtone Creek, where the mountain
types of Eucalypts commence, viz., E. gunnii (0), H. stellulata, E. pauciflora, E.
viminalis, and E. amygdalina.
In descending from the Omeo plateau towards Tongio, I observed that BH.
hemiphloia, variety albens, commences at 2500ft. This Eucalypt grows preferen-
tially upon the Plutonic and Metamorphic formations of the dry valleys of Tongio
and Ensay, the Upper Snowy (in Victoria), and of the Deddick and Tubbut Rivers.
In the descent from Omeo one finds E. melliodora at 2000ft. In passing from the
deep valleys of Ensay to the coast ranges, which may be regarded as commencing at
Mount Elizabeth, I observed that E. hemiphloia ceased at 1000ft. at Hnsay, but this
elevation must probably be decreased considerably, as I believe this type grows on the
Tambo at Numlamungie, approximately 700ft. above sea level. The same remarks
apply to E. stellulata, on the summit of Fainting Range. I have noticed that E.
polyanthema, E. goniocalyx, K. piperita, and E. obliqua re-appear at their upper limit
in this locality, and it is about here that the littoral types touch and intermingle
with the mountain forms. It is here to be noted, as at other places to which I have
referred, that it is the lowland types which slightly cross the limit set them by the
summit of the cool mountains, and descend to some little extent in the warmer slopes,
rather than the mountain types, which descend the moist southern side.
At Noyang, on the Tambo Valley road, there occurs a tract of Plutonic and
Metamorphic rocks, and here there is, to a certain extent, a re-appearance of the
sub-alpine types intermingling with a greater number of the lowland Eucalypts.
Further down the road E. muelleriana appears, growing on the Silurian for-
mations from the upper limit of the Tertiary marine beds at 7OOft. to 1300ft.
I observed only one or two scattered examples growing in the forest of E. piperita on
the Tertiary sands near Monkey Creek.
I might instance other examples in the mountain country, in the sources of the
Buchan River—at Murrendel, at Woolgulmerang, and at Bonang, but the above will
suffice, and my observations in these localities, although made with sufficient care to
satisfy me as to the general results, want the precision which I have endeavoured to
give to the cases which I have noted. My notes on the eastward of the Snowy River,
in Croajingolong, were made many years ago, and fall into line with those I have
given, but as they were less precise even than those made between the Tambo and
Snowy Rivers, I prefer not to rely upon them.
The observations which I have now recorded as to the distribution in the
THE EUCALYPTS OF GIPPSLAND. 107
mountainous and littoral tracts of Gippsland, of the several types of Eucalypts, seem
to me to show that elevation above sea level, rainfall, and aspect have had more to
do in their distribution than have the geological formations, or than have the different
soils produced therefrom.
The range of mountain and littoral types has, it seems, varied from time to time ;
that is to say, the mountain types have at times spread into or over the littoral tracts,
while at others the littoral types have spread into the mountains.
The manner in which E. piperita, E. obliqua, E. goniocalyx, E. polyanthema,
and others climb up the southern slopes, and there meet with the subalpine forms,
and others whose habitat is in the dry valleys within the coast range, shows that the
distribution of these forms is regulated by climate.
It would only require that the conditions which now prevail on the southern
slopes should extend over the drier mountain pleateaux; in other words, that the
rainfall should be greater there than it is now for those Kucalypts to extend still further
northwards from the lowlands.
I have been led to suspect, from the study of the Gippsland Eucalypts, that
varieties have arisen, not only through the impulse of the favouring conditions of
climatic change, but also that to the same cause is due an emigration from the
lowlands to the highlands, and vice versa.
The instance of E. goniocalyx is significant. It is at present almost entirely
confined to the cool, moist littoral regions, where the soil is good, and the cool,
humid gullies of the southern slopes ; it does not, in its typical form, extend across
the watersheds into the warmer and comparatively dry northern slopes. Its place is
there taken by a dwarf variety having marked differences.
Were the climate to become so much altered that the same conditions obtained
throughout the district as are now found, for instance, at Dargo, Ensay, or Glen
Falloch, 1 conclude that the lowland form of E. goniocalyx would disappear, and the
dwarfed mountain form might take its place. The species would no longer be
represented by a tree reaching 200ft. to 250ft., with a smooth-barked massive bole,
but by a straggling tree with, often, a short gnarled trunk, a wrinkled strongly
persistent bark, scanty foliage, and a young form, in which the leaves, although
opposed, are not pointed-ovate or broadly lanceolar, nor haying the powerful odour
which characterises the lowland form.
The existence of small colonies of mountain species in the lowlands, as, for
instance, the before-mentioned E. pauciflora and E. viminalis (6) points, I think, to
survivals from a time when the climate was much colder than it is now. ‘The
oscillations of level which have affected the coast line of the southern half of this
108 THE EUCALYPTS OF GIPPSLAND.
continent may be as well studied in Gippsland as in any part that I have seen from
Shoalhaven to beyond Adelaide. These must: certainly have produced variations of
climate, extending back beyond the Cainozoic period.
The Gippsland Alps have not been submerged below a contour line of some
800ft. to 1000ft. above sea level, not only during that period of time, but that land
surface must have been continuous backwards to the time when the Mesozoic coal
measures of Gippsland were formed.
The lauraceous and other plants which have been found in the Miocene drifts
of Gippsland, indicate, as does also the fauna of the marine limestones of that district,
a warmer climate than of the present day.
Lake Karng at Mount Wellington, if it be a moraine lake, points, on the
other hand, to an alpine climate, descending to within at least 3000ft. of the sea
level. Such changes of climate have evidently been attended by a corresponding
change in the Tertiary flora, in which that element, which is now characteristic
of Australia, has gradually predominated. Such changes of climate may, as it
seems to me, also account in part for the great number of recorded types of
Eucalypts and their varieties, and of which no less than 35 occur in Gippsland.
Geological formation, as producing variation of soil, has no doubt influenced
the present distribution of the Eucalypts, but its effects cannot be made out so
clearly as those produced by climate, but the broad features can be readily seen by
anyone travelling through Gippsland. LE. tereticornis grows almost entirely on lands
which have been at one time lake or estuary beds, or in the alluvial flats of rivers.
The stringy-bark Eucalypts preter the Tertiary sands and sandy clays. H. odorata
grows mainly on the Miocene limestone, but this partiality to particular formations
is not so apparent when all the Eucalypts are considered. Still, in looking over
the whole of Gippsland, I observed some marked cases which it would be well to note.
A good instance is afforded by E. amygdalina regnans (0), which, in Gippsland,
grows almost wholly upon the Mesozoic coal measures. E. hemiphloia appears to be
confined to the Plutonic and Metamorphic areas of the Tambo and Snowy Rivers. A final
instance may be taken from the Gelantipy tableland, to the west of the Snowy River,
which shows how certain Eucalypts grow preferentially upon certain formations.
This tableland is formed by a great thickness of Devonian and Plutonic rocks,
overlaid by more or less connected sheets of Tertiary basalt. 1 observed that on the
former grow especially KE. piperita, E. globulus, E. sieberiana, and E. amygdalina,
and on the latter formation E. stuartiana, E. melliodora, EK. polyanthema, and E.
macrorhyncha.
The annexed table has been compiled from observations which I have made in
almost all parts of Gippsland, and shows the distribution of the Eucalypts on the
various formations.
THE EUCALYPTS OF GIPPSLAND. 109
INFLUENCE OF SETTLEMENT ON THE Eucatyptus Forests.
The influence of settlement upon the Eucalyptus forests has not been confined
to the settlements upon lands devoted now to agriculture or pasturage, or by the
earlier occupation by a mining population.
It dates from the very day when the first hardy pioneers drove their flocks and
herds down the mountains from New South Wales into the rich pastures of
Gippsland.
Before this time the gramminivorous marsupials had been so few in comparative
number, that they could not materially affect the annual crop of grass which
covered the country, and which was more or less burnt off by the aborigines, either
accidentally or intentionally, when travelling, or for the purpose of hunting game.
These annual bush fires tended to keep the forests open, and to prevent the
open country from being overgrown, for they not only consumed much of the
standing or fallen timber, but in a great measure destroyed the seedlings which had
sprung up since former conflagrations.
The influence of these bush fires acted, however, in another direction, namely,
as a check upon insect life, destroying, among others, those insects which prey upon
the Eucalypts.
Granted these premises, it is easy to conclude that any cause which would
lessen the force of the annual bush fires, would very materially alter the balance of
nature, and thus produce new and unexpected results.
The increasing number of sheep and cattle in Gippsland, and the extended
settlement of the district, lessened the annual crop of grass, and it was to the interest
of the settlers to lessen and keep within bounds bush fires which might otherwise be
very destructive to their improvements.
The results were twofold. Young seedlings had now achance of life, and a severe
check was removed from imsect pests. The consequences of these and other
co-operating causes may be traced throughout the district, and a few instances will
illustrate my meaning.
The valley of the Snowy River, when the early settlers came down from Maneroo
to occupy it, as for instance, from Willis downwards to Mountain Creek, was very open
and free from forests. At Turnback and the Black Mountain, the mountains on the
western side of the river were, in many parts, clothed with grass, and with but a few
large scattered trees of Hi. hemiphloia.
110 THE EUCALYPTS OF GIPPSLAND.
The immediate valley was a series of grassy alluvial flats, through which the
river meandered. After some years of occupation, whole tracts of country became
covered with forests of young saplings of E. hemiphloia, pauciflora, viminalis,
amyedalina, and stellulata, and at the present time these have so much increased, and
grown so much, that it is difficult to ride over parts which one can see Ie the few
scattered old giants were at one time open grassy country.
Within the last twenty-five years many parts of the Tambo valley, from Ensay up
to Tongio, have likewise become overgrown by a young forest, principally of E.
hemiphloia and macrorhyncha, which extend up the mountains on either side of the
valley. This dates especially from the time when the country was fenced into large
sheep paddocks, when it became very important that bush fires should be prevented
as a source of danger to the fences, and even when fire occurred the shortness of the
pasturage checked the spread.
Similar observations may be made in the Omeo district, namely, that young forests
of various kinds of Eucalypts are growing where a quarter of a centuryago the hills were
open and park like. In the mountains, from Mount Wellington to Castle Hill, in
which the sources of the Avon River take their rise, the increase of the Eucalyptus
forests has been very marked. Since the settlement of the country, ranges, which
were then only covered by an open forest, are now grown up with saplings of E.
obliqua, E. sieberiana, and others, as well as dense growths of Acacia discolor, A.
verniciflua, and other arborescent shrubs. These mountains were, as a whole,
according to accounts given me by surviving aborigines, much more open than they
are now.
In the upper valley of the Moroka River, which takes its rise at Mount
Wellington, I have noticed that the forests are encroaching very greatly upon such
open plains as occur in the valley. I observed one range, upon which stood scattered
cigantic trees of E. sieberiana, now all dead, while a forest of young trees of the same
species, all of the same approximate age, which may probably be twelve years,
srowing so densely that it would not be easy to force a passage through on horseback.
Again, at the Caledonia River, as at the Moroka, the ranges are in many parts quite
overgrown with forests not more than twenty years old. The valleys of the Wellington
and Macalister Rivers also afford most instructive examples of the manner in which
the Eucalyptus forests have increased in the mountains of Gippsland since the country
was settled. The forest in these valleys, below 2000ft. above sea level, is principally
composed of Eucalyptus polyanthema, E. macrorhyncha, with occasional examples of
EK. melliodora and E. stuartiana ; while E. viminalis occupies the river banks and moist
flats. I noticed here that E. melliodora and E. macrorhyncha formed dense forests
of young trees, apparently not more than twenty-five years old. In some places,
moreover, one could see that the original forest had been composed, on the lower,
THE EUCALYPTS OF GIPPSLAND. 111
undulating hills and higher flats of a few very large EH. melhodora, with scattered trees
of E. polyanthema and HK. macrorhyneha. At the present time the two latter have
taken possession, almost to the exclusion of E. melliodora. In other places E. poly-
anthema or EK. macrorhyncha predominate ; but, on the whole, I think the latter will
ultimately triumph over its rivals, unless the hand of man again intervenes.
Such observations may also be made in Western and Southern Gippsland, but,
of course, with reference to different species of Kucalypts.
In the great forest of South Gippsland many places can be seen where there are
substantially only two existing generations of trees; one of a few very large old trees,
the other of very numerous trees which are, probably, not older than 30 to 40 years,
and, in most cases, certainly not half that period. The older trees of this second
erowth do not, I suspect, date further back than the memorable ‘“‘ Black Thursday,”
when tremendous fires raged over this tract of country. . It may also be inferred, from
the constant discoveries during the process of clearing of blackfellows’ stone
tomahawks, that much of this country, now covered by a dense scrub of gum
saplings, Pomaderis apetala, Aster argophylla, and other arborescent shrubs, that the
country was at that time mainly an open forest.
I might go on giving many more instances of this growth of the Eucalyptus
forests within the last quarter of a century, but those I have given will serve to show
how widespread this re-foresting of the country has been since the time when the white
man appeared in Gippsland, and dispossessed the aboriginal occupiers, to whom we owe
more than is generally surmised for having unintentionally prepared it, by their
annual burnings, for our occupation.
The age of the new forests does not, however, depend merely on the general
observation that they have sprung up since the settlement of the country in 1840.
I have been enabled to make some direct observations, which show the size of
certain trees of known age, and which will serve as comparison for the general growth
of the forests.
In 1864 the discovery of auriferous quartz reefs in the Crooked River district,
caused a township, which is now called Grant, to be formed on the summit of the
mountains, near the sources of Good LuckCreek. In part of the Government reserve,
upon which the warden’s quarters and police camp stood, and which was cleared of
timber, a few young E. amygdalina trees grew, and were permitted to remain. One
of these was lately kindly measured for me by Mr. W. H. Morgan, M.M.B., who
found it to be 56ft. high and 10ft. in girth 3ft. above the ground. ‘This tree is an
example of very many others of the same species now growing on the surrounding
ranges. At Omeo, in the Government reserve, a number of young E. viminalis are
112 THE EUCALYPTS OF GIPPSLAND.
now 60ft. high, which in 1863 were only small saplings under 5ft. in height. On the
road from Sale to Port Albert, which was formed somewhere about 1858-59, there
are numerous places where E. viminalis and E. muelleriana and other species are now
growing, upon the ditches formed at the sides of the road. Those, for instance, at
Lillies Leaf are on an average about 30ft. high.
These instances show how the occupation of Gippsland by the white man has
absolutely caused an increased growth of the Eucalyptus forests in places. I venture,
indeed, to say with a feeling of certainty, produced by long observation, that, taking
Gippsland as a whole, from the Great Dividing Range to the sea, and from the
boundary of Westernport to that of New South Wales, that, in spite of the clearings~
which have been made by selectors and others, and in spite of the destruction of
Eucalypts by other means (to which I am about to refer), the forests are now more
widely extended and more dense than they were when Angus M‘Millan first descended
from the Omeo plateau into the low country.
I have spoken just now of the destruction of Eucalypts by other means than the
hand of man, for clearing his holdings, and the following are the facts I have gathered
concerning the subject :—
About the year 1868-4 I observed that a belt of Red-gums which extended across
the plains between Sale, Maffra, and Stratford were beginning to die. Gradually all
the trees of this forest, as well as in other localities, perished. At that time my
attention was not drawn to the investigation of the cause. Later, however, probably
about 1878, I observed the Red-gum forests of the Mitchell River Valley to be dying,
just as those at Nuntin and elsewhere had died years before. I then investigated the
subject, and found the trees were infested with myriads of the larve of some
one of the nocturnal Lepidoptera. These devoured the upper and under epidermis
of the leaves, thus asphyxiating the tree. Some 75 per cent. of that forest died that
year, and subsequently almost all the surviving trees died also. Since then I have
observed the same larvee at work, some of which, when kept until they had passed through
their several metamorphoses to the perfected insect, were pronounced by Professor
M‘Coy to be examples of Urubra Iugens. Whether this insect has in all cases been
the agent in destroying the red-gums I cannot affirm. Probably not wholly, but Iam
satisfied that the greater part of the Red-gum trees which have died in Gippsland
from obscure causes have been killed by its agency.
The inference may be drawn from the above observations of forests having been
killed by infesting insects, that each species of Eucalypt, or at any rate each group
of allied species, will have attached to it some particular insect which preys upon it
rather than upon any other Kucalypt.
THE EUCALYPTS OF GIPPSLAND. 113
If this is so, we ought to find some one tree selected for destruction out of a
number of species, and it is the case with the Red-gum, for it falls a victim to Urubra
lugens, whilst its neighbours the White-gum (EK. viminalis), the Swamp-gum (EK.
gsunni), and the Yellow-box (E. melliodora) are untouched and in vigorous
health.*
I feel little doubt that this will explain why it is that in many parts of the
country, at all elevations above sea level, certain tracts of dead forest are to be found.
Twenty-five years ago I noticed that during the course of three years all the White-
oums (HE. viminalis) in part of the Omeo district died, whilst E. pauciflora and E.
stellulata remained alive.
I have said that in my opinion the increased growth of the Eucalyptus forests
since the first settlement of Gippsland has been due to the checking of bush fires
year by year, and to the increase thereby of the chance of survival of the seedling
Kucalypts, and to the same cause we may assign the increase of the leaf-eating insects
which seem in places to threaten the very existence of the Red-gum.
Bush fires, which swept the country more or less annually, kept down the
enormous multiplication of insect life, destroymg myriads of grasshoppers and
caterpillars, which now devastate parts of the Gippsland district, spoiling the oat
crops, and eating the grass down to the ground.
The ravages of the larvae of Lepidoptera are at present greatly aided by the
sickly state in which many of the Red-eum forests in Gippsland now are. The
long-continued use of the country for pasturage, and the trampling of the surface of
the ground by stock, has greatly hardened the soil, so that rain which formerly, in
what I may call the ‘‘ normal state’? as regards Eucalypts, soaked in, now runs off.
In the course of successive droughty seasons the soil of such places becomes
thoroughly dry and hard, so that the Red-gum is deprived of much moisture which
it otherwise would have in reserve. The trees are wanting in vigour, and thus
unable to withstand the attacks of insect pests.
The effects produced by man’s interference with the balance of nature, by settling
new countries, is not only of great scientific interest, but is also of importance in
showing us how and why it is that the labours of the graziers and farmers are being
carried on year by year under the increasing attacks of insect pests.
The subject is a tempting one, but to pursue it further would be foreign to the
subject of these Notes, which is the ‘‘ Eucalypts of Gippsland.”
* I have observed, however, in some localities E. melliodora and E. piperita have been slightly attacked by
Urubra lugens.
OF GIPPSLAND.
EUCALYPTS
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THE EUCALYPTS OF GIPPSLAND.
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EXPLANATION OF PLATES.
Puate 8.
Eucalyptus amygdalina (b).
Lower leaf of sapling, Mount Livingstone.
Lower leaf of sapling, Dargo.
Upper leaf of sapling, Wentworth River.
Leat of aged tree, Wentworth River.
Seedling, Mount Livingstone.
Buds, natural size.
Section of fruit.
Anthers x 24 linear.
Anthers x 6 linear.
Fruit, natural size, Mount Livingstone.
PLATE 9.
E. amygdalina (d).
Lower leaf of sapling, Stradbroke.
Lower leaf of sapling, Wild Horse Creek.
Leat of young tree, Dargo,
Leaf of aged tree, Tambo River.
Leaf of aged tree, Carrajung.
Seedling, Bruthen Creek.
Anthers x 24 linear.
Filaments and anthers x 6 linear, Stradbroke.
Cluster of buds, natural size, Stradbroke.
Fruit, natural size, Dargo.
Puate 10.
E. amygdalina (e).
iB
2.
3.
4.
Terminal shoot of young sapling.
Cluster of fruit.
Buds.
Seedling.
THE EUCALYPTS OF GIPPSLAND. 117
5. Leaf of aged tree, Tambo River.
6,7. E. amyedalina (c), Leaves of aged tree.
8. Section of fruit.
9. Cluster of fruit.
10. Buds.
11. Anthers x 6 linear.
12. Anthers x 24 linear.
Puate 11.
E. muelleriana.
1. Seedling, Insolvent Track.
2. Seedling, Toongabbie.
Seedling, Shady Creek, South Gippsland.
4. Spray, with buds and flowers, Woodside.
5, 6. Anthers x 24 linear, Woodside.
7,8. Anthers x 24 linear, Toongabbie.
9,10. Anthers x 24 linear of E. eugenioides, Agnes River.
11,12. Anthers x 24 linear, E. piperita, Toongabbie.
18, 14. Anthers x 24 linear, Wild Horse Creek.
15. Section of fruit of E. muelleriana, Woodside.
16. Cluster of fruit of EK. muelleriana, Woodside.
ise)
Pratt 12.
FE. nuelleriana.
1. Leaf of sapling (8ft.), Daraman.
2. Leaf of sapling, Insolvent Track.
3. Leaf of sapling, Woodside.
4. Leaf of aged tree, Four Mile Creek.
5. Leaf of aged tree, Woodside.
6. Leaf from top of sapling, Tambo River.
7. Leaf of aged tree, Toongabbie.
Prate 13.
1. Leaf of sapling of E. piperita, Toongabbie.
2. Leaf of sapling of E. capitellata, Ostler’s Creek.
3. Leat of sapling of EK. obliqua, Wallialla.
4. Leaf of sapling of E. macrorhyncha, Dargo.
5. Leaf of sapling of E. eugenioides, Drouin West.
118 THE EUCALYPTS OF GIPPSLAND.
Fruit of E. piperita.
6. Bruthen.
7. Fainting Range.
8. Tambo River
9. Wild Horse Creek.
10. Stockyard.
11, 12. Monkey Creek.
13. Groggingee.
14. Bruthen.
15. Bulgaback.
16. Insolvent Track.
17. Bruthen.
18. Toongabbie.
19. Insolvent Track.
E. eugenioides.
20. Agnes River.
21. Drouin.
E. muelleriana.
22. Tambo River.
23. Insolvent Track.
24. Toongabbie.
25. Woodside.
E. capitellata.
26, 27. Lilies’ Leaf.
28. Mirboo North.
29. Ostler’s Creek.
E. macrorhyncha.
30. Drouin West.
31. Fanwick.
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32. Dargo.
Prats 14.
1. Seedling, E. piperita, Toongabbie.
Seedling, E. capitellata, Ostler’s Creek.
Seedling, E. macrorhyncha, Bulgaback.
4. Seedling of E. eugenioides, Agnes River.
5. Seedling of E. obliqua, Agnes River.
co be
THE EUCALYPTS OF GIPPSLAND. 119
Puate 15.
1, 2,3, 4. Anthers x 24 linear, K. pulverulenta, Boolarra.
5, 6, 7,8. Anthers x 24 linear, EK. stuartiana, Lilies’ Leaf.
9. Pair of leaves of aged tree, E. pulverulenta, Monkey Creek.
10. ‘Terminal leaf (alternate), E. pulverulenta, Monkey Creek.
11. Leaf of aged tree, E. stuartiana, Toongabbie.
12. Seedling, EK. stuartiana, Toongabbie.
13. Seedling of E. pulverulenta, Monkey Creek.
Buds and Fruit of E. viminalis (6).
14. Squirrel Forest.
15. Omeo.
16. Morwell.
17. St. Pancras Peak, Woolgulmerang.
18. Highteen Mile Creek, Dargo High Plains.
19. Fraser’s Plain.
20. Dargo High Plain.
21. Grant.
22. Woolgulmerang.
E. viminalis (a).
23. Monkey Creek, Port Albert Road.
24. Alberton.
25. Coal Creek.
26. Wentworth River.
27. Tucker Creek.
28. Wild Horse Creek.
29. Sources of ‘Tucker Creek.
30. Woolgulmerang.
31. Upper Wentworth River.
E. pulverulenta.
32. Monkey Creek, Port Albert Road.
33. Boolarra.
34. Ostler’s Creek.
E. stuartiana.
35. Lilies’ Leaf.
36. Stradbroke.
37. Cuidmundie, Dargo.
38. Insolvent Track.
All natural size,
120 THE EUCALYPTS OF GIPPSLAND.
PuatE 16.
E. viminalis (6).
1. Lower leaf of sapling, Dargo.
2. Pair of upper leaves, sapling, Dargo.
3. Leaf of young tree, Dargo Plains.
4. Leaf of aged tree, Dargo Plains.
5. Leaf of aged tree, Omeo.
6. Young plant, Omeo.
7,8, 9. Anthers x 24 linear of E. viminalis (a), Alberton.
10, 11, 12. Anthers of K. viminalis (6), Cuidmundie, Dargo.
13, 14, 15. Anthers x 24 linear, E. viminalis (0), Squirrel Forest.
16,17, 18. Anthers x 24 linear, E. viminalis (6), Omeo.
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ARTICLE IV.—A New Famity or HyprompEa, TOGETHER WiTH A Description
OF THE STRUCTURE OF A New Species or Prumutaria, By W. Batpwin Spencer,
M.A., Proressor or Bronocy in THe University oF MELBOURNE.
(Read Noy. 12th, 1890.)
The following deals with two hydroid forms obtained by Mr. J. Bracebridge
Wilson, M.A., from the neighbourhood of Port Phillip. One of these is so distinct
from any hitherto described that it must be taken as the type of a new Family, to
which the name of Hydroceratinide is given, the other is a new and somewhat
curious species of the genus Plumularia. The paper is, therefore, divided into two
parts—(1) On the Hydroceratinide, a new family of the order Hydroidea, and (2) On
Plumularia procumbens, a new species of the genus Plumularia.
(1) On tHe Hyproceratinip®, 4 New Faminy oF THE OrpER Hyprorpea.
The form in question undoubtedly calls to mind, in the general appearance of
the dried specimens, one of the two genera which were placed in a distinct family—
the Ceratellade—by Dr. Gray,* and, when this paper was read before the Royal
Society of Victoria, I provisionally referred it to this family.f Dr. Gray had doubtfully
* Norz.—Up to the time of reading this paper I had only the necessarily
somewhat imperfect descriptions (since taken from dried specimens only) of Dr. Gray
and Mr. Carter to be guided by together with the figures of the former. Those of
Dehitella bore a general resemblance to the specimen in question which was
placed provisionally in that genus as a new species. Through the courtesy of Dr.
EK. P. Ramsay I have since been able to examine specimens of Ceratella and Dehitella
from the Australian Museum, Sydney, and have received from there specimens
obtained from the New South Wales coast, and from Lord Howe Islands by Mr.
Whitelegge, to whom I am much indebted for kind assistance and information. The
examination of these specimens has shown that the form with which this paper deals
differs in such important points not only from the Hydractiniide but also from
the Ceratelladz that it must be placed in a separate family and I desire to here record
my indebtedness to the authorities of the Australian Museum, Sydney, for the
courteous assistance received from them.
* Proc. Zool. Soc., Nov., 1868.
122 A NEW FAMILY OF HYDROIDEA.
placed the family in the group Porifera. Mr. Carter subsequently had the opportunity
of examining both Dr. Gray’s specimens and others of allied forms from New
Zealand and the Cape, and rightly referred them all to the Hydroidea, placing them
in the family Hydractiniide, This classification was adopted by Dr. v. Lendenfeld, and
also, at first, by Mr. Bale in his valuable contribution to the literature of Australian
zoology—the ‘Catalogue of Australian Hydroid Zoophytes.”* Subsequently, Mr.
Bale had the opportunity of examining spirit-preserved specimens with the hydranths
and soft parts present, and, finding that these differed very considerably from those
of the Hydractiniide in the presence of irregularly distributed capitate tentacles,
&e., rightly separated them once more from the Hydractiniide, and adopted the
name of Ceratelladz for the family.| I was unaware of this short paper of Mr.
Bale’s until my attention was kindly drawn to it by Mr. Whitelegge, of the
Australian Museum.
Whilst Dehitella in its general form (in the dried specimen) is somewhat like
the specimen obtained by Mr. Wilson, there can be little doubt that the two are
markedly distinct, even so far as the skeleton goes. Dehitella has most clearly what
have well been called hydrophores', and agrees closely in this respect and in the
general formation of its skeleton with Ceratella ; in their soft parts we may
reasonably suppose that a corresponding agreement exists. Now the soft parts of
the Ceratella are known,§ and they differ very strongly from those of the form with
which this paper deals, so that we are probably correct in assuming that a similar
difference exists between the latter and Dehitella. There is, on the other hand (as
will be shown subsequently), quite as marked a distinction between the new form
and any member of the family Hydractiniide as exists between the former and the
Ceratelladx, so that it is necessary to create a new family for its reception.
Ihave not so far, though numerous sections have been cut, been able to
detect any reproductive elements, and therefore the description is incomplete, but will
quite sufficiently serve to distinguish it from other forms.
Family Hydroceratinide.
Hydrophyton, consisting of a mass of entwined hydrorhiza, with a skeleton in
the form of anastomosing chitinous tubes; the surface is studded with tubular
hydrotheecx, into which the hydranths can be completely retracted. Hydranths
sessile and connected with more than one hydrorhizal tube, claviform with a single
verticil of filiform tentacles. Defensive zooids present with a solid endodermal axis
and nematocysts borne at the distal end.
* Catalogue of Australian Hydroid Zoophytes. W. M. Bale, 1884.
+ Bale, Proc. Linn. Soe., N.S.W., Vol. III., pt. 2, p. 748; also Brazier, Proc. Linn. Soc., N.S.W., new series, Vol. I.
page 575; Whitelegge, loc. cit., p. 578.
+ Bale, Proc. Linn. Soc., N.S.W., 1888, p. 749.
§ Bale, loc. cit.
A NEW FAMILY OF HYDROIDEA. 123
Clathrozoon wilsont, n. gen. et. n. sp.
Hydrophyton irregularly sub-dichotomously branched, expanded in one plane fan-
shaped. Main stem somewhat flattened and ridged, arising from an expanded base, dark
brown in colour, with the surface showing a series of tortuous grooves. Smaller
branches cylindrical. The whole composed of a number of branching and
anastomosing tubes, with chitinous walls complete, except those of the most external
ones. Hydrothece, which have the form of tubular spaces with chitinous walls,
project slightly from the surface of all the branches, irregularly placed on
the main stems, spirally on the branches. The whole, except the external
openings of the hydrothec, completely enclosed in a thin transparent chitinous
layer, from which arise very numerous cylindrical tubes enclosing the defensive
zooids, and which forms also a funnel-shaped collar, projecting beyond the lips of the
hydrothece.
Locality—Near Port Phillip Heads, Victoria. Mr. J. Bracebridge Wilson.
The description of the species is the same as that of the genus, and I have much
pleasure in dedicating it to Mr. Wilson.
The specimens were obtained from two spots, each within a distance of five
miles of Port Phillip Heads, that is, in Bass Straits, close to the Victorian shore.
They were dredged in water of from twenty to twenty-two fathoms, and were placed
directly into strong alcohol so that, like all received from Mr. Wilson, they are in
excellent histological preservation. The polypes are, of course, in a state of retraction,
and though very many sections from various parts of the hydrophyton have been cut,
nowhere as yet have I been able to detect the presence of reproductive organs, or of
individuals modified in connection with these, and though, therefore, unable to give
any account of the reproductive organs, I have thought it advisable to describe and
figure the animal carefully so far as can yet be done.*
The largest colony secured by Mr. Wilson measures 10in. in height, by 4in. in
greatest width, and at first glance recalls to mind, to a certain extent, one of the
dark coloured fan-shaped gorgonid forms. A cursory examination, however, at
once shows that it does not belong to this group of animals.
If the whole colony be secured, it is found to be attached by a broad flattened-out
base, attached to some solid structure; from this arises a single large stem, perhaps 2in.
in thickness, slightly flattened out in one plane, and distinctly ridged (Fig. 1). From
the edge at either side of the plane lateral branches are given off, varying much in size;
* That the animal is not at all common is shown by the fact that, though during this summer (1890-91), Mr. Wilson
has been kind enough to spend a considerable amount of time in dredging, with the special object of procuring more
specimens, not a trace of this hydroid has been brought up in the dredge, though the same ground which previously
yielded it has been visited.
124 A NBW FAMILY OF HYDROIDEA.
from these other branches are given off on either side, which again divide in a roughly
but not constantly dichotomous manner, the smaller branches being cylindrical in
shape. All the branches in the living and spirit-preserved specimens usually le in the
one plane, but when dried they often become slightly twisted, and thus thrown out of
the plane. There is thus a considerable resemblance in general appearance between
the form Clathrozoon on the one hand, and Dehitella and Ceratella on the other,
especially between the two first mentioned, owing to the rounded branches of
Dehitella. If, however, we come to examine the structure of the two more minutely,
we find that the differences are very marked; whilst both have the skeleton in the
form of a horny network, that of Clathrozoon has more the appearance of being
composed of a series of anastomosing tubes, and rather less of the appearance of an
open meshwork than is the case in the Ceratellade ; but, what is of much greater
importance, it presents on its surface a large number of circular openings at the
extremities of slightly projecting chitinous tubes, which form true hydrothece.
These are very distinct from what Mr. Carter and Dr. Gray aptly described as scoop-
like projections of the chitinous network of the Ceratellade, though they
unfortunately gave to them the name of hydrothece, that of hydrophore being
more correct and suggestive of their real function, as at most they can but afford a
support for the proximal part of the hydranth.
Skeleton (Figs. 4, 5, 6, 7, 8, 9, 11, 12, 18, 16).
When the branches of spirit-preserved specimens, with the soft parts present,
are viewed under the lens, the surface is seen to be covered with a great number of
tortuous grooves, filled with light yellowish coloured material—the ccenosarcal
tubes. The edges of the grooves are formed by the chitinous perisare of a dark-brown
colour (Fig. 6), and arranged in a spiral manner are circular projections, the external
opening of chitinous cylinders forming the hydrothec and containing the retracted
polypes: These hydrothece are encircled by the tortuous grooves, and supported by
extensions of the ordinary perisarc. The whole surface is covered over by an
extremely thin and delicate colourless layer of perisarc common to the whole branch.
This layer is not usually recognised until sections are cut, but the whole surface is
seen to be studded with small cylindrical tubes, which are really formed from this
thin layer and the spaces in which are continuous with the tortuous grooves below
the latter (Figs. 7, 8, 9).
Sections show also that this thin layer rises up somewhat from the general
surface, and is attached to the lips of the hydrothece ; beyond the margin of the
latter it is continued on so as to form a very thin collar-like extension, acting as an
operculum. The arrangement is represented in the figures, especially in Fig. 13,
which is meant to be a diagram showing the relationship of a hydrotheca to the
A NEW FAMILY OF HYDROIDEA. 125
perisarc, as seen in a very thick section cut transversely to the length of a branch.
This operculum-like structure is very thin and liable to be torn away, but is present
in all well-preserved and cut specimens. Sometimes it projects as a stiff collar,
sometimes (Figs. 4, 16) it is thrown into folds, and at others is withdrawn into the
hydrotheca. With only specimens in which the polypes are completely retracted to
examine, I cannot say how much of the body of the polype is covered by this thin
collar when the animal is fully expanded.
When the soft parts are dissolved away by potash the whole branch appears to
consist of a meshwork of tubes with chitinous walls (Fig. 11), which anastomose so
freely that the skeleton appears, in surface view, to consist of an irregular chitinous
network, which has a considerable superficial resemblance to the skeleton of a horny
sponge, though this resemblance is even more strongly marked in the case of the
Ceratellade. On cutting sections, however (Figs. 7, 8, 9), the tubular arrangement can
be recognised. The structure is essentially identical in the branches and branchlets,
whatever be the size, the only difference consisting in the greater number of tubes
entering into the composition of the larger branches as compared with the smaller
ones. The tubes (Fig. 7) run roughly parallel to the length of the branch, continually
branching and anastomosing. The spaces which they contain vary much in calibre.
Towards the exterior the chitinous walls are much stronger and thicker than in the
interior, and the most external series are incomplete, forming grooves rather than
tubes. The thin external layer of the branch, previously alluded to, touches the
chitinous lips of these grooves (Figs. 8, 9), and thus a complete inclosure for the most
external-lying ccenosarcal tubes is formed.
Sections also show that the larger circular openings, visible on the outside, lead
down into tubular spaces, the walls of which are formed of chitinous material similar
to that of the tube walls with which they are connected. These form the hydrothece,
and the spaces within them are continuous with one or more of those within the
tubes surrounding them (Figs. 4, 7,12). The thin external chitinous (?) layer always
passes up to be attached to the rim of the hydrothece. The cylindrical structures
(P’) on this thin layer are very simple in form, open at both ends, and serve during
life to contain the defensive polypes which are directly connected with the external
layer of ccenosarcal tubes. (Fig. 5).
Measurements of the Skeleton.
Diameter of branch taken, ‘7 mm.
Number of tubes, as seen in longitudinal section, entering into the
composition of a branch of this size, 7—9; as seen in transverse
section, 40—50.
Thickness of the tube, varying, but averaging about ‘07 mm.
126 A NEW FAMILY OF HYDROIDEA.
Thickness of wall of tube, 0525 mm. at the exterior, some of those in the
centre being not more than ‘00525 mm.
Hydrotheca.— Transverse section, -175 mm. — °1225 mm.
Hydrotheca.—Longitudinal section, *525 mm. (average).
Soft Parts. (Figs. 8, 4, 5, 10, 12, 14, 15).
The chitinous tubes are filled by the ccenosarcal tubes which branch and
anastomose freely. No one of the latter tubes is more prominent than any
other. ‘he polypes are distributed over the whole surface, having apparently no
definite arrangement on the larger stems or branches, but, in the smaller ones,
a very distinct spiral arrangement (Figs. 6,11). One corresponds to each of the
hydrothees, seen in the skeleton, and is capable of complete retraction within the
latter. When thus retracted the opening is protected by the thin collar-like projection
formed of the outer layer of perisarc mentioned above.
Each polype has the typical hydroid form. It is somewhat tubular, with a
conical hypostome, from the base of which arises a single circlet of solid tentacles,
which vary in number from six to ten, and are provided with minute nematocysts,
the threads of which are short, stiff, and unbarbed. When the tentacles are retracted
they curl over towards the mouth, much in the same manner as do those of
Pedicellina, amongst the Polyzoa.
The body of the polype is formed of the typical layers—an ectoderm of somewhat
columnar cells, an endoderm of larger cells less defined in shape, and between the
two a very strongly marked layer of mesogloea (.00875 mm. in thickest part). From
the bases of the ectoderm cells pass off long processes (Fig. 14), much more
strongly marked than is usual in hydroid forms. In its greatest breadth one of
these fibres measures (0058 mm. ‘They give rise to a circular band of what are
evidently in function, ‘muscle fibres.” In figure 14 these are represented as seen
when an oblique longitudinal section of a polype is cut.* Possibly some of them
may have lost their connection with the ectoderm cells from which they have been
derived. Hach has a very definite outline and contents, which, when stained, have a
somewhat granular appearance. ‘They all lie external to the mesoglea, and form a
very definite band round the body of the polype, the position of which is represented
in the diagram (Fig. 12).
Sections, transverse and longitudinal, show that the basal ends of the polypes
are continuous with more than one (sometimes as many as four or five) of the
* These bear a considerable resemblance to those described by Weismann as occurring in the ceenosare of Plumularia
echinulata. See Entstehung der Sexualzellen bei den Hydromedusen, Pl. VIII., Fig. 9b.
A NEW FAMILY OF HYDROIDEA. 127
ceenosarcal tubes. In this they resemble the polypes in the group Hydrocoralline.
The cavities of the polypes and of the meshwork of tubes are hence directly
continuous. The tubes are formed of the two primary layers with, if present
at all, an extremely thin layer of mesoglea between them. The endoderm is always
unilaminar and distinct ; the ectoderm is composed of smaller cells, the nuclei, but
not the outlines, of which are clearly marked. It appears to be frequently more than
one cell in thickness, and to be irregular in outline, spirit specimens showing
projections (Fig. 10) which come in contact with the chitinous walls. Probably
during life there is very little space between the hard and soft parts. Within the
ccenosarcal tube are frequently seen large globular structures containing darkly
staining portions (Fig. 10. x.)
The whole of the external layer of ccenosarcal tubes is studded over
with an enormous number of small defensive polypes of very simple structure
(Figs.8, 5,15). One corresponds to each of the cylindrical structures which rise up
from the outer thin perisare layer, and consists of a stalk and ahead. The former
is directly continuous with the ccenosare tube, and consists of an internal axis, the
nature of which is difficult to determine. It is solid, and contains no prolongation of
the tubular canal of the canosarc, but appears to be continuous with the endoderm,
‘though I cannot as yet make out the cellular structure. It is very much shrunken
in spirit specimens. The outer layer is thin and continuous with the ectoderm of
the ceenosare (Fig, 15). In spirit-preserved specimens it has the appearance of a
thin layer of granular protoplasm with nuclei, the diameter of which is greater than
the thickness of the protoplasmic layer, in which the outline of cells cannot be
distinguished. No trace of muscle elements can be distinguished.
The head consists of a little mass of nematocysts, with the remnants of the cells,
which have given rise to them, and the nuclei of which can be clearly seen. Each
nematocyst is fusiform in shape, and the whole form a little group lying close to the
open end of the cylindrical tube, through which the threads can doubtless be ejected.
None of the nematocysts have the threads put out, and their exact form cannot be
therefore described.
In relative size and structure these defensive polypes resemble more than
anything else the machopolypes of the Plumulariidz, and have no resemblance to the
defensive polypes of Hydractinia or Podocoryne, whilst none have as yet been
described in the Ceratellade.
In figure 3 is represented a diagrammatic restoration of a branch of the colony
only the soft parts being drawn. The upper surface is supposed to have been cut
away, and the network of ccenosarcal tubes is shown branching and anastomosing.
The connection of the alimentary polypes with the tubes is drawn, and the small
128 A NEW FAMILY OF HYDROIDEA.
defensive polypes are shown arising from the outermost layer of ccenosarc. The hard
skeleton parts represented in figure 11 occupy the spaces between the tubes in figure 3,
the polypes corresponding in position to the large circular openings leading down
into the hydrothece. The general relationship of the polypes to the skeleton and the
ccenosarcal tubes is represented diagrammatically in figure 12.
A ffimties of the Hydroceratinde.
Clathrozoon differs in important points from any hydroid hitherto described,
and it is somewhat difficult to determine its affinities, more especially in the
absence, as yet, of any knowledge of its reproductive elements.
At first glance it would appear to be related to the Ceratellade, but this is simply
in consequence of the superficial resemblance in the skeleton of the two groups ;
each consists of a branching mass of entwined perisarcal tubes, and here the
resemblance ends, for, whilst the branching network of tubes may be compared with that
constituting the skeleton of the Hydractiniide and the Ceratellade, there are present two
structures entirely wanting in the members of these two families, (1) the hydrothecz, and
(2) the external layer of thin perisarc, with its projecting cylindrical tubes. The first
of these structures are quite unlike those of any other hydroid in their simple
shape, the thickness of their walls, the relation of these to the surrounding
perisarec tubes, and the number of openings leading through the wall into the
enclosed space. At the same time, if the cavity in the scoop-like projection, formed
of a small network of perisarcal tubes in the Ceratellade, were to become deeper
and to penetrate the substance of the branch, and if the branches of the network
bounding the cavities thus produced were to ‘‘run together’ and thus give rise to a
tubular structure, in which apertures were left, we should have produced a structure
similar to the hydrothece of Clathrozoon. ‘The second of these structures is,
apparently, not present in any other hydroid, and the manner of its origin is difficult
to conceive. There is no such external layer of ccenosare investing the colony as is
present in the Hydractinude, and which might be supposed to secrete such a layer.
On the contrary, the surface of the colony shows a series of much branching tortuous
perisare tubes, each incomplete externally, and the whole covered in by this thin
continuous layer, which does not give the idea of having been formed as a separate
outer covering for each most external tube, though it must apparently have been pro-
duced in this way. The little cylinders arising from it enclose each, it is important to
notice, a defensive zooid, which arises directly from the ccenosarcal tube immediately
below. In one or two of the sections cut transversely near to the end of a growing
branch, the perisare tubes appear to have a thin imnermost lining distinguishable
from the rest of the perisarc; possibly this may be comparable to this outer layer,
but at all events the latter, as shown. in sections, passes continuously from lip to lip
A NEW FAMILY OF HYDROIDEA. 129
of the outer grooves, and has nothing to do with any layer lining the latter internally.
That it is connected in development with the soft structures in these there can be
little doubt if only on account of its relationship to the protective zooids to which
they give rise.
The extension of this layer beyond the mouth of the hydrothece so as to form, as it
were, an operculum for these, is a curious feature, and one not met with, so far as I am
aware, in any other hydroid. This particular part is very flexible, being capable of being
thrown into folds (Fig. 16), or even of being pulled back within the outer rim of the
hydrotheca by the retreating zooid. Taken altogether the skeleton, though in certain
respects showing a resemblance to the Hydractiniide and Ceratellade, differs essentially
from that of these forms in the two important points dealt with above.
In dealing with the soft structures we find a most curious combination of
characters, each one characteristic of hydroid forms, belonging to groups perfectly
distinct from one another.
The network of ccenosarcal tubes resembles, to a certain extent, that of the
Hydractinide, Ceratellade and Hydrocoralline, but even here we have to note the
entire absence of an external continuous layer, characteristic certainly of the first
and third, and, probably, also of the second group.
The alimentary polypes are sessile, and distinctly “ claviform,” that is, have
tubular bodies with a conical hypostome and a single circlet of simple solid tentacles,
in which points they resemble the genus Clava. Those of the Ceratellade*, on the
other hand, have scattered capitate tentacles, and in the Oa Eat they are
provided with a strongly developed hydrocope.
The gastrozoids again resemble those of the Hydrocoralline, and differ from all
others amongst the Hydrozoa in being connected with several of the ccenosarcal
tubes.
The defensive zooids, or dactylozooids, resemble more than anything else, certain
individuals characteristic of the Plumulariide, to which the name of ‘« machopolype ”
has been applied... They consist of a solid stalk bearing a number of nematocysts at
its free end, each zooid being enclosed in a distinct protective case or nematophore.
The structure of the gastrozooids thus calls to mind the genus Clava, that of the
dactylozooids the family Plumulariide, and the relationship of the gastrozooids to
the coenosareal tubes, the sub-order Hydrocoralline. This combination of characters,
together with the nature of the skeleton, serves to render the Hydroceratinide distinct
from any family of Hydroidea yet known,
* Bale, loc. cit.
130 A NEW FAMILY OF HYDROIDEA.
2. A New Species oF PLumuuaria.
This form was dredged by Mr. Wilson in Port Phillip, and differs, as far
as I can ascertain, from any yet described. In (1) that the nematophores are
perfectly independent of the hydrothec, (2) that no intrathecal ridge is present,
(3) that the hydrothece are cup-shaped, with smooth untoothed margin, and are set
at some distance apart from one another, the form in question shows the features
characteristic of the group to which Allman gave the name of Eleutheroplea.
The arrangement of the hydrothece would prevent it from being placed in any
of the three divisions—Isocola, Anisocola, and Monopyxis—into which Kirchenpauer
proposed to divide the genus.
The species may be described as follows :—
Plumularia procumbens, n. sp.
Hydrocaulus upwards of 6in. in length. The whole colony procumbent with
large polysiphonic branches running in a roughly horizontal direction. From
one side of these principally arise smaller polysiphonic branches, all lying in one plane.
From the sides of the branches arise numerous pinne (at largest }in. in length)
irregularly arranged; in addition to these, hydrocladia (pinnules) may arise direct
from the hydrocaulus. Two pinnules, alternate, arise from each joint of the primary
pimne. Pinnules composed of alternate small and large joints, the latter only
bearing hydrothece and nematophores. Nematophores bithalamic with simple
terminal aperture, one beneath each hydrotheca, two at the level of the mouth of the
hydrotheca, all independent of the latter. Two nematophores in the angle between
the pinnule and main stem of the pinna; nematophores scattered irregularly in great
numbers over the surface of the polysiphonic stem and branches.
Colour.—Light-brown stems.
Hab.—Port Phillip, Mr. J. Bracebridge Wilson.
The colony reaches a considerable size, and from the fact that the main branches
run in one direction (Fig. 20) with the smaller ones arising principally from one side,
it is inferred that in all probability it is procumbent in habit. The branches of the
hydrocaulus are all polysiphonic and strong, the pinne and hydrocladia, which arise
from them, being very small indeed in comparison. The two latter are given off all
round the branches, and reach at most the length of jin., whilst the main branch
may be 6in. in length.
Skeleton.
(a) Structure of large branches (Figs. 24, 25, 26, 27, 28).
A NEW FAMILY OF HYDROIDEA. 131
Under the lens the tubes making up the branches can be clearly seen, the
surface being marked by darkish brown lines, due to the edges of the perisarcal
tubes seen in optical section (Fig. 25). The tubes on the exterior run parallel
to one another along the length of the stem, branching very rarely. From the
surface arise irregularly, and on all sides—(1) large pinnate shoots, the proximal
parts of which are covered with one or more layers of dark-brown perisare, within
which only one tubular cavity is contained; (2) smaller branches (hydrocladia),
corresponding to the pinnules in structure ; (8) a great number of nematophores,
or protective cases, for the minute defensive zooids which are characteristic of the
group.
In transverse section (Figs. 27, 28) the branch is seen to be made up of a great
number of tubes—as many as 40-50 in a branch of average size—the number varying
with the size of the branch.
Each tube has a definite and thick perisarcal wall, which shows clear indications
of arrangement in layers. In places the cavities within the tubes are seen to be (Figs.
24, 27, 28) in communication with one another. A striking feature of both trans-
verse and longitudinal sections is the presence of a distinct central tube, always clearly
recognisable, both on account of its size and the slightly yellow colour of its walls
when compared with the surrounding ones (Fig. 24). This central tube, which, from its
relations to the other parts, is probably to be regarded as homologous with the main
stem of a monosiphonic form, passes along all the branches of the colony, and from
it arise every one of the pinne and hydrocladia, though, as will be shown shortly,
these may be also connected with the other tubes which go to form the polysiphonic
stem, Professor Allman has figured* in Aglaophenia coarctata a connection existing
between the various tubes of a polysiphonic stem, and this is most clearly seen in the
species under consideration. The connection is, however, somewhat different from
that obtaining in 4. coarctata, which is thus described by Allman :—‘‘ Communication
is effected by very short processes, which are given off from the component tubes,
those of two juxtaposed tubes meeting one another and inosculating in such a way
as to suggest the conjugation of a zygnema.” In longitudinal section (Fig. 24) the
various tubes are seen to be arranged so that they run parallel to the central one,
and at intervals their walls are pierced by apertures. Where the pinne and
hydrocladia pass through, the walls of the latter are connected with those of the
stem-tubes, and apertures are formed opening into the cavities of the latter,
(6) Structure of the pinne, etc. (Figs. 21, 22, 23, 25, 26).
The origin of these from the stem is shown in figure 25, their structure in
figures 21, 22, and 26.
*« Challenger” Reports, Hydroidea, Part I., Plumulariide.
132 A NEW FAMILY OF HYDROIDEA.
The basal portion of each pinna is strengthened by the development of a very
strong layer of dark brown coloured perisarc, continuous with that of the tube walls.
This thick external layer ends abruptly just before reaching the distal end of either
the first or second joint (Fig. 25). Each jot beyond the first one or two carries
two pinnules. There appears to be a slight variation in the most proximal joints ;
sometimes the first, sometimes the first and second, differ from the rest, in bearing
each only one pinnule.
The pinnules are alternate, and consist of a varying number of joints, which are
alternately shorter and longer, the latter only bearing the hydrothece and
nematophores (Figs. 21, 26). This arrangement is constant in all specimens which
IT have examined, In other Plumularie the pinnules of which are composed of
alternately longer and shorter joints, such, for example, as P. setaceoides, goldsteint,
delicatula,* the shorter ones always bear a nematophore. This is absent in
P. procumbens. The hydrothece are cup-shaped, with a smooth margin, and are
placed on the side facing the central stem of the pinna. The most distal pinnules
carry one hydrotheca each, the proximal as many as six. The cavity is separated
from that of the jomt by a septum pierced by a circular opening, which lies
near to the external wall. There is no trace of an intrathecal ridge.
The nematophores are three in number on each of the larger joints; one is
placed below the hydrotheca, two at the level of its upper margin. Tach is
bithalamic with the terminal chamber cup-shaped, and the proximal one somewhat
canaliculate. The walls are thin (Fig. 22), except where the division into the two
parts is formed, at which spot they thicken considerably, and give rise to a circular
ridge projecting upwards into the distal chamber. ‘The opening is single and terminal,
and each nematophore is only attached by its proximal end, where the walls are thin,
and may be thrown into slight folds.
The walls of the pinnule joints show internal ridges, which are always more
prominent on the side facing the central stem of the pinna, and thin away towards
the opposite surface. They are always arranged thus—(1) In the larger joints
there is a ridge close to each extremity, with a third one corresponding in position
to the septum of the hydrotheca; this varies much in development, being sometimes
searcely noticeable. (2) In the smaller joints there are uniformly two ridges. (8)
In the projection from the joint of the pinna bearing the pinnule there is always one
ridge. Taken altogether the result is that each line of division in the pinnule has
one ridge immediately on either side of it (Fig. 21).
In the axil of the pimnules there are present—(1) Two nematophores correspond-
ing in structure exactly to those described above, (2) between these a curious
* Bale. Catalogue of Australian Hydroids, Pl. XI.
A NEW FAMILY OF HYDROIDEA. 133
structure formed of the perisarc, having the shape of a cone with the apex cut off
(Fig. 21). The space within the latter communicates by the narrow end with
the exterior, and by the broader with the cavity of the pinna joint. Into it
cells of the ectoderm may enter to a slight degree, but more usually it appears
to be unoccupied (in spirit-preserved specimens), and I am quite unable to
attach any meaning to it, though it is a perfectly constant structure. It has
nothing to do with the reproductive structures. Possibly it may serve as a means
of allowing of the ingress and egress of water to and from the perisarcal tubes.
Any space between the ectoderm and the perisarc in the very numerous tubes which
compose the colony must presumably be filled by liquid. The openings leading into
the hydrothece and nematophores from the stem are small and narrow, and quite
filled up by the soft parts. When sudden contraction takes place part of the soft
portions must be withdrawn through these openings and occupy space within
this perisarcal tube previously, presumably, occupied by fluid. If there be
some means of expelling this fluid then the sudden contraction of the polypes and
machopolypes is rendered more easy. It may be that these openings serve this
purpose. The openings are guarded, as it were, by two machopolypes.
Soft Parts (Figs. 17, 18, 19, 24).
(a) Larger branches.
These are all polysiphonic, The ccenosareal tubes of which each is composed
may be divided into two divisions—(1) a central one, (2) others varying in number
and surrounding this. The tubes very rarely branch in such a way that two
running longitudinally arise from a common one, though (Figs. 24, 27, 28) they
frequently are united with one another by short transverse branches passing through
openings in the perisarcal walls. Hach consists of ectoderm and endoderm
containing a cavity. From the outermost series arise a great number of minute
machopolypes or defensive zooids. The central one gives origin to all the branches
passing out into the pinne and hydrocladia, with which the branches are irregularly
studded. As they pass from the centre to the external surface they are connected
with a varying number of the surrounding ccenosarcal tubes, a feature which is
especially marked in the case of the pinne (Figs. 17, 24). ‘This arrangement is
clearly seen when sections are cut, and has not, so far as 1 am aware, been noted before.
Bale,* in speaking of the structure of the stem and branches in the Plumulariude,
says that in most polysiphonic species ‘‘ the primary jointed stem is slender (the
requisite strength being given by the compound stem, which is only developed as the
zoophyte increases in size), and the branches spring, not from the jointed stem, but
from the supplementary tubes which grow up in contact withit. For example, in
* Genera of the Plumulariide, with observations on various Australian Hydroids. Proc. R. 8. Victoria, 1886.
134 A NEW FAMILY OF HYDROIDEA.
Aglaophenia longicornis we find at the back of the original slender-jointed stem a
stouter secondary tube, and from this spring, at regular intervals, the alternate
pinnately arranged branches.* . . . fa 28 2) Keeper mr amind the
hydrorhizal origin of the polysiphonic atin we see thatin A glaophenia longicornis, for
example, every one of the main pinne is equivalent to a separate shoot of such
species as A. parvula, a fact which is further illustrated by the presence, near the
base of the stem in the latter species (and, indeed, in many others), of a long
oblique joint similar to that which exists near the base of each pinna in A. longicornis.
: I have not hitherto met es any species with branches springing
ly ai the jointed stem and the added tubes.”
It will be seen at once that an important difference exists in this respect between
the species now described, and those examined by Mr. Bale, and, so far as I am
aware, any other investigator. Not only is there a very distinct connection between
the soft and hard parts of all the tubes of the polysiphonic stem, but the central
one, though distinguishable from the rest, has the same fundamental structure as
the latter so far as its walls are concerned, and shows no traces of joints.+ The
question naturally arises What elements are we to consider as entering into the
structure of the polysiphonic stem in this form ?
We may regard it as composed of—
(1.) A central tube equivalent to the hydrocaulus of a monosiphonic form,
which is surrounded by a series of added but modified hydrorhizal
elements, or
(2.) A number of hydrorhizal elements forming branch-like structures, and
giving off pinne, or
(8.) A number of hydrocauli in close apposition to one another.
The last is the most improbable, since, if it were the case, we might expect pinne
to be given off from all or any of the tubes composing the stem, whilst they all arise
primarily from the central one. The same objection applies to the second, and the
distinction which undoubtedly obtains between the central and all the other tubes
seems to point to the fact that the former is the fundamental portion, the latter being
secondary structures. At the same time a hard and fast line of distinction between
hydrocaulus and hydrorhiza cannot possibly be drawn. In some forms of
Plumulariide we find the pinne arising directly from the hydrorhiza, in others they
arise from the sides of a hydrocauline structure which grows upwards from the
* The italics are mine.
+ At the free extremity of each branch it is continued directly on into a jointed stem forming the centre of a pinna,
and may probably be regarded as having lost its jointed nature subsequent to its being completely enclosed by the
surrounding tubes of the polysiphonic stem.
A NEW FAMILY OF HYDROIDEA. 135
hydrorliza. Sometimes the structure bearing the pinne, or the central stem of the
pinne itself, may be strengthened by the addition of tubes clearly growing up from
the hydrorhiza. These tubes, in some monosiphonic forms, may be feebly developed,
whilst in typical polysiphonic ones apparently homologous structures may be constant
and greatly developed. In the former case the distinction between hydrocaulus and
hydrorhiza is clear enough, but then if the true polysiphonic form be investigated, we
find that the supplementary structures grow around the primary jointed stem, and (1)
finally (as in A glaophenia longicornis) give rise to the pinnate branches, just as does the
hydrocauline tube in such a form as P. falcata, the jointed stem bearing no branches;
or (2) they form (as in P. procumbens) an enclosing mass for the original tube which
alone gives off the pinne, though these may be connected with the hard and soft parts
of the enclosing branches. Thus what must be regarded as homologous structures,
may either (1) be feebly developed, and retain their original hydrorhizal nature ; or (2)
be strongly developed and (a) give rise to pinne, and, as it were, usurp the function of
the original hydrocauline tube which they support, or (6) assume an intermediate form,
the original tube which they inclose alone giving rise to pinne, with which, however,
as well as with the former they are in organic connection.
(b) Pinna, etc.
There is little to say with regard to the structure of the soft parts contained in
the pinne ; the hydranths have the form typical of the genus Plumularia, with the
body divided into two parts by a central constriction, the distal part bearing a single
circlet of solid tentacles at the base of a broad hypostome, the proximal half being
somewhat globular and presenting no special feature.
The machopolypes of the main branches and the pinne are of precisely similar
structure, corresponding to those of other forms which Lendenfeld has distinguished
as ‘“‘guard animals with corticating capsules.”* There is no trace of any with
adhesive cells such as are found in the genus Aglaophenia.
Each machopolype consists of a proximal tubular part lying in the proximal
half of the nematophore and a distal swollen part, which contains rounded
nematocysts. I have been unable to study the living form, and have only seen
spirit specimens, in which the soft parts are of course much contracted ; in these,
lines running from the head down the stalk probably indicate muscle fibres, the
machopolype being capable of great extension.
In figure 17 is represented a restoration of the soft parts of a small portion of
one of the polysiphonic stem, together with one of the hydrocladia and pinne which
arise from it.
* Zeitsch. f. Wissen. Zool., Vol. XXXVIIL., p. 338.
136 A NEW FAMILY OF HYDROIDEA.
(c) Reproductive structures.
The only specimen bearing reproductive structures was a male colony. The
gonothece (Figs. 18, 19, 23) are simple and pear-shaped, with a large terminal
opening and short stalk springing from the axil of a pinnule. Each contains one
gonophore (Fig. 18), which in longitudinal section is seen to arise from the
blastostyle and to fill nearly the whole cavity. The blastostyle in the spirit-
preserved specimen examined was much shrunken with a terminal swelling (D) below
the opening of the gonotheca. ‘The cell-layers could not be distinguished. The two
prominent parts of the gonophore are the mass of sperm cells (sp), which stain deeply,
and the spadix (Fig. 19, end.) Outside the sperm cells a very thin layer of ectoderm
can be distinguished. I failed after long searching to recognise any trace of
reproductive elements in the ccenosarcal tubes or blastostyle, or any appearance of
the formation of more than one gonophore in each gonothece, though in
P. echinulata Weismann states that very often a second gonophore may be found
before the contents of the first have passed out.* In P. halecioides also it seems
that two may be present at the same time.
* Entstehung der Sexualzellen bei den Hydromedusen, p. 180.
+ Loc. cit., p. 184.
DESCRIPTION OF PLATES.
Plates 17, 18, 19, and 20 refer to Clathrozoon wilsoni.
Prats 17.
Figure 1.—The skeleton of a dried specimen of Clathrozoon wilsont, life size,
from a photograph.
Figure 2.—Skeleton of part of a colony of Clathrozoon wilsoni, drawn from a
spirit specimen.
Puate 18.
Figure 3.—Restoration of the soft parts. The hard skeleton parts are entirely
omitted. Only the end of one of the branches of a colony is represented, but this
is typical of the structure of the whole. The upper surface of the branch is
supposed to be cut away to show the connection of the gastrozooids with the
ccenosarcal tubes. The outermost of the latter are studded with the defensive zooids.
The whole is much magnified, the actual size of the branches being represented in
figures 1 and 2.
Figure 4.—Represents a longitudinal section of a small portion of a branch to
show the connection of a polype (retracted) with the ccenosarcal tubes. In the
figure the polype is connected with three of these. (A) The perisare walls are shown,
and the thin external layer forming at the mouth of the hydrotheca an operculum
P. Outline drawn with the camera under Zeiss A. oc. 2.
Figure 5.—Transverse section of a small portion of the outermost part of a
branch to show a defensive polype, connected with one of the outer ccenosarcal tubes,
which is connected again with a deeper lying one. The defensive poly pe is enclosed
by the nematophore. LE. is one of the most external tubes, the outer wall of which is
formed of the thin layer (P.) of perisare, on which lies an accumulation of foreign
substances, spicules, &c. Outline drawn with camera under Zeiss A. oc 2.
Puate 19.
Figure 11.—A small branch much enlarged showing the skeleton only, the soft
parts having been dissolved in potash. The hydrothece are shown arranged spirally
and projecting with circular margins slightly from the surface. The network of
chitinous tubes is seen to represent somewhat in appearance the clathrate horny
skeleton of certain sponges. The somewhat prominent lines on the surface corres-
138 A NEW FAMILY OF HYDROIDEA.
pond to the dark curving ones in figure 6. The thin external layer with the
nematophores is only shown at the edges, in reality it covers the whole surface.
Letters as before. x 50.
Figure 12.—A diagrammatic drawing to show the relationship of the hard and
soft parts as seen at the external part of a longitudinal section of a branch. A
gastrozooid is shown partially expanded. Hy. the walls of the hydrotheca. C.
ccenosarcal tube. Ma. protective zooid arising from the outermost ccenosarcal tube.
P. the thin external layer of perisarc. P’. the continuation of the latter to form a
collar beyond the opening of the hydrotheca. P”. the nematophore. T. tentacle.
Figure 13.—A semi-diagrammatic drawing of a hydrotheca as seen in a thick
transverse section of a branch very much enlarged. The walls of the hydrotheca are
supported by numerous extensions of the perisarcal network, and the openings into
the internal-lying end of the hydrotheca are shown. ‘The perisare tubes have
branched and anastamosed to such an extent that their walls form a network, and
the tubular structure is not so evident as it is in the more deeply lying parts.
Letters as before.
Figure 14.—The basal part of a gastrozooid cut obliquely to show the strong
band of muscular fibres derived from the ectoderm together with the thick layer of
mesoglea. Ect. ectoderm. End. large granular endoderm cells. M. thick mesog-
lea layer. Mus. muscle fibres, some in connection with the ectoderm cells, from the
basis of which they arise, others cut at a deeper level, at which their connection with
ectoderm cells is not seen. Possibly some have lost their connection with ectoderm
cells. Drawn under Zeiss F. oc. 2.
Figure 15.—A defensive zooid highly magnified. It consists of a stalk and head.
The former is in connection with a ccenosarcal tube one wall of which, as seen in
longitudinal section is shown. The centre of the stalk is solid and continuous with
the endoderm, though no cellular structure can be determined. Outside this is a
thin layer of ectoderm in which nuclei are scattered. The head consists of nemato-
cysts at the bases of which remnants of the cells in which they have been formed
can be seen with their nuclei. ct. ectoderm. End. endoderm. M. mesoglea.
N. nematocyst. Drawn under Zeiss apo. 4.0 mm. apert. 0.95 oc. 12.
Figure 16.—The collar-like operculum surrounding the margin of a hydrotheca
seen from above. The collar is thrown into folds. Drawn under Zeiss F. oc. 2.
Pruate 20.
Figure 6.—A terminal branch, much enlarged to show the circular openings of
the hydrothecx, within which the polypes are withdrawn, and also the general spiral
arrangement of the hydrothece. The surface is covered with tortuous grooves,
A NEW FAMILY OF HYDROIDEA. 139
bounded by dark lines, indicating the edges of the perisarc, and is studded with
nematophores seen at the edges of the branch. Hy. Hydrothece. P. nematophores.
Figure 7.—A longitudinal section of a small branch, showing the skeleton. The
tubes of which the branch is composed are seen to be very irregular, but to run,
especially towards the central part, in a direction generally parallel to the length of
the branch. E. spaces or grooves, immediately beneath the thin external layer.
Hy. hydrothece. P. thin external layer of perisarc. PP’. the continuation of P.
to form an operculum at the mouth of the hydrothece. PP’. the nematophores.
x 36.
Fioure 8.—A transverse section of a small branch, to show the tubes of which it
is composed. The tubes vary in size, and open into one another. The outer ones
are incomplete externally, forming grooves, the lips of which are usually touched by
the thin external Jayer of perisare. ‘Two hydrothece are cut through. The perisare
is thicker towards the outer than in the inner part of the branch. Letters as in
Fig. 7, x 70.
Figure 9.—A small portion of the external part of a branch very much enlarged
to show the roughly concentric layers of which the perisare is formed. Letters
as in Fig. 7.
Figure 10.—Small portion of a ccenosarcal tube with the perisare walls from
the interior of a branch. Ect. the ectoderm in which the outline of the cells cannot
be clearly distinguished, and which varies in thickness in various parts coming in
contact, in certain places, with the perisarc walls. End. the unilaminar endoderm,
the cells of which are larger than those of the ectoderm. x. globular structures of
unknown significance containing darkly staining parts which lie apparently within
the cavity of the tube. Drawn under Zeiss F. oc. 2.
Plates 21, 22, 23 refer to Plumularia procumbens.
Puate 21.
Figure 17.—Restoration of the soft parts only of Plumularia procumbens. The
polysiphonic stem is shown in section with the transverse connections between the
various tubes which compose it. Down the centre runs the main tube from which
arise all the lateral branches—pinne and hydrocladia—which are in connection with
a varying number of the surrounding tubes. The most external ones give off
numerous machopolypes, and on the pinne are shown the groups of polypes, each
consisting of one gastrozooid and three machopolypes. One blastostyle is shown
with a gonophore, arising from the angle between a pinnule and the main stem of the
pimna. x 30.
Figure 18.—Longitudinal section of a male gonangium with its contained
blastostyle and gonophore. In the centre of the gonophore lies the manubrium. Bi’.
140 A NEW FAMILY OF HYDROIDEA,
blastostyle. D. swollen distal, end of blastostyle. Sf. sperm. G. walls of
gonotheca.
Figure 19.—Transverse section of a male gonangium and gonophore. On the
left side between the gonophore and the wall are seen traces of the much compressed
blastostyle. End. endoderm of manubrium. Ect. ectoderm outside sperm cells (sp.)
Pruate 22.
Figure 20.—Portion of a colony of Plumularia procumbens showing the poly-
siphonic stem and the pinne arising irregularly from this, x 1}.
Figure 21.—Much enlarged portion of a pinnule or hydrocladium to show the
alternate longer and shorter joints. The latter bears no hydrotheca or
nematophores ; the former bears a hydrotheca with the nematophores at
the level of the mouth of the former, and one in the median line
below. The thickenings in the walls of the pinnule are shown in their positions
and the two nematophores in the axil between the pinnule and the main stem of the
pinna, and at A, the conical structure opening to the exterior. N'. upper pair of
nematophores. NN. lower median nematophores. N*. nematophores in the axil of
the pinnule and pinna stem. A. conical process in the axil leading from the interior
of the pinna to the exterior.
Figure 22.—Much enlarged view of a nematophore seen in optical section, and
showing the two chambers.
Figure 23.—Much enlarged view of a male gonangium.
Figure 24.—Semi-diagrammatic drawing of a longitudinal section of the poly-
siphonic stem of the same to show the relationship of the hard and soft parts and the
distinctness of the central tube with its walls slightly yellower than those of the
surrounding tubes. C. central tube. H. central stem of pinna. H/’. hydrocladia.
L. lateral tubes. N.nematophores. T. transverse connections uniting the various
tubes. x 80.
Prate 23.
Figure 25.—Much enlarged view of the termination of a polysiphonic stem,
showing the numerous nematophores on the stem and the pinne &c., arising
irregularly. The basal parts of the pimne are strengthened by the formation of a
thick perisare wall continuous with that of the tubes forming the stem. x 20.
Figure 26.—A highly magnified portion of a pinna.
Figures 27 and 28.—Transverse sections of the polysiphonic stem to show the
component tubes—skeleton only—with the large central one, from which in figure 28
a pinna is arising. The external tubes are studded with nematophores.
Trans, PS Victoria. Plate 17
CLATHROZOON WILSONI.
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TRANSACTIONS
h
OF THE
ROYAL SOCIETY OF VICTORIA.
VOL: PART II.
1891.
CONTENTS.
ON THE OCCURRENCE OF THE GENUS BELONOSTOMUS IN THE
ROLLING DOWNS FORMATION (CRETACEOUS) OF CENTRAL
QUEENSLAND, sy R. Erueriper, Junr., PALEONTOLOGIST TO THE
GroLoaicAL Survey or New Sourn Wates, SypNEY, AND ARTHUR
Smith Woopwarp, F.Z.S., &c., oF THE DrparTMENT oF GEOLOGY,
British Muszum (Nat. Hist.), Lonpon. (With Plate 1) - = -
ON THE STRUCTURE OF CERATELLA FUSCA (GRAY), sy W. Batpwin
Spencer, M.A., Proressor or Brotoay in THE UNIVERSITY OF
Metsourne. (With Plates 2, 3, and 3a.) - : : - -
ADDITIONAL OBSERVATIONS ON THE VICTORIAN LAND
PLANARIANS, sy Artuur Denpy, D.Sc. (With Plate 4) : :
LAND PLANARIANS FROM LORD HOWE ISLAND, sy W. Batowin
Srencer, M.A., Proressorn or Brotogy IN THE UNIVERSITY OF
Metsourne. Parr I.—Description or Species. (With Plates 5 and 6.)
EDITED BY PROFESSOR W. BALDWIN SPENCER, M.A., HON. SEC.
J" MELBOURNE.
PUBLISHED FOR THE ROYAL SOCIETY
PAGE
25
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BY THE SPECTATOR PUBLISHING COMPANY LIMITED, 270 POST OFFICE PLACE
FesBruary, 1892.
PRICE, 85s.
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ReAWaOUth Mid On
merece SOOTY” Ol” VICTORLA.
Watron:
HIS EXCELLENCY THE EARL OF HOPETOUN, G.C.M.G.
President :
PROFESSOR W. C. KERNOT, M.A., C.E.
Vice=Presidents :
E. J. WHITE, F.R.A.S. | H. K. RUSDEN, F.R.G.S.
‘bon. Treasurer
C. R. BLACKETT, F.C.S.
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PROFESSOR W. BALDWIN SPENCER, M.A.
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Council :
R. L. J. ELLERY, F.R.S. | PROF. ORME MASSON, M.A.,, D.Sc.
SS DENDY, D.Sc. | H. MOORS.
G. §. GRIFFITHS, F.R.G.S8. | PROF. B. T. LYLE, M.A.
JAMES JAMIESON, M.D. ALEXANDER SUTHERLAND, M.A.
Ape S-) FUCKS, MUA. /B.Se: Cay, MORE ins alu:
PROF. H. LAURIE, LL.D. Asi. SWAY, ONDA
TRANSACTIONS
OF THE
RON SOCIETY OF “ViCTORiLA
VOR: PART II.
1891.
CONTENTS.
ON THE OCCURRENCE OF THE GENUS BELONOSTOMUS IN THE
ROLLING DOWNS FORMATION (CRETACEOUS) OF CENTRAL
QUEENSLAND, sy R. Erneriper, Junr., PALEONTOLOGIST TO THE
GroLogicaL Survey oF New South Watses, SypDNEY, aND ARTHUR
Smith Woopwarp, F.Z.S., &c., or THE DEPARTMENT oF GEOLOGY,
British Museum (Nar. Hist.), Lonpon. (With Plate 1) -
ON THE STRUCTURE OF CERATELLA FUSCA (GRAY), sy W. Batpwin
Spencer, M.A., Proressor oF Biology IN THE UNIVERSITY OF
Mexpourne. (With Plates 2, 3, and 38a.) 2 2 z 2 -
ADDITIONAL OBSERVATIONS ON THE VICTORIAN LAND
PLANARIANS, sy Artuur Denny, D.Sc. (With Plate 4) s 2
LAND PLANARIANS FROM LORD HOWE ISLAND, sy W. Batpwin
Spencer, M.A., Proressor or BrioLogy IN THE UNIVERSITY OF
Mesourne. Part I.—Description or Species. (With Plates 5 and 6.)
EDITED BY PROFESSOR W. BALDWIN SPENCER, M.A., HON. SEC.
MELBOURNE.
PUBLISHED FOR THE ROYAL SOCIETY
PAGE
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BY THE SPECTATOR PUBLISHING COMPANY LIMITED, 270 POST OFFICE PLACE
FEBRUARY, 1892.
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ARTICLE I.—On tox OccurRENCE oF THE GENUS BELONOSTOMUS IN THE ROLLING
Downs Formation (CRETACEOUS) OF CENTRAL QUEENSLAND, BY R. ETHERIDGE,
JunR., PALEONTOLOGIST TO THE GEOLOGICAL SuRvEY oF New Sours Watgs,
Sypney, anp ArTtHUR SmitH Woopwarp, F.Z.S., &c., of THE DEPARTMENT OF
Grotocy, British Museum (Nat. Hist.), Lonpon. (With Plate I.)
I. Iyrropvuction.
The fine specimen about to be described was entrusted to our care by Mr.
George Sweet, of Brunswick, Melbourne, by whom it was obtained when on a
collecting tour in Central Queensland, during the year before last (1889); together
with many other very interesting fossils, which one of the present writers hopes some
day to have the pleasure of working out.
The principal fossil exhibits a long, slender fish, with deep narrow ganoid scales
and feeble fins, bent upon itself at about the middle point, and wanting the greater
part of the head. The remains are preserved on counterpart slabs, one side being
shown, reduced one-half in Plate I., Fig. 1, while various details are illustrated of
the natural size in the accompanying Figs. 2-7.
The information afforded by this specimen is supplemented by four other
fragments in a similar matrix, numbered M. 31, M. 26 A, M. 28, and M. 80 respectively.
The first exhibits an imperfect cranium, much fractured and imbedded in hard rock.
In the second the remains of the gill arches are preserved. No. M. 28 shows part
of the operculum, supraclavicle, supratemporals, and adjoining scales in counterpart ;
and No. M. 80 isaconnected series of six large vertebral centra, bearing parts of their
arches. Taken together, the specimens make known nearly all the more important
features in the skeleton of the fish; and it is at once obvious that we are concerned
with an unusually large species of the well-known Upper Mesozoic ‘“ Ganoid ”
Belonostomus*, and with which we have much pleasure in associating the name of
Mr. Sweet as B. sweets.
Species that are apparently allied have already been recorded from the Upper
Cretaceous of Western Europe, India, and Brazil; and the present discovery is of
great interest as extending still further the ascertained geographical range of the
genus during Cretaceous times.
* L. Agassiz, Poiss. Foss., 1884, II., Pt. 2, p. 140.
2 ON THE OCCURRENCE OF THE GENUS BELONOSTOMUS IN THE
2. Previous Notices oF FIsH-REMAINS IN THE Roxiiinc Downs Formation.
The first record of the existence of fish-remains in the immense expanse of
rocks, now termed by Mr. R. L. Jack, Government Geologist of Queensland, the
‘* Rolling Downs Formation,” is due to the late Mr. Charles Moore, of Bath, England,
who found numerous fragments of teeth and scales in the Wollumbilla blocks
submitted to him by the late Rev. W. B. Clarke, M.A. He regarded the teeth and
certain of the scales as those of Hybodus, and other scales as representing the genus
Lepiodotus.* Subsequent researches have so far failed to reveal further traces of these
genera.
The next reference to Australian Cretaceous fish with which we are acquainted
is the brief record by Mr. R. Etheridge, F.R.§., of the caudal portion of the vertebral
column and several scales of Asfidorhynchus in the Hughenden beds,+ Central
Queensland, associated with Ammonites, and other fossils of Cretaceous age.
After a lapse of some years, Mr. R. L. Jack, whilst engaged with General
Fielding in the exploratory traverses for the Queensland Trans-Continental
Railway, discovered at Kamilaroy, Leichhardt River, a bed of magnesian limestone,
containing, with other fossils, sharks’ teeth and small vertebre.{ The associated
fossils clearly proved this bed to belong to that portion of the great Rolling Downs
Formation extending from Cloncurry to Hughenden. These remains were
determined by one of us to be the teeth of Ofodus appendiculatus, Ag., and the
vertebree those of a small Teleostean fish. Subsequent to this these teeth were
described and figured,|| together with a portion of the vertebral centrums of a large
Lamna from the Walsh River beds of the Rollimg Downs Formation. This interesting
specimen forms a portion of the collection of the Queensland Museum, Brisbane, and
was kindly communicated by Mr. C. W. de Vis, M.A., the Curator of that Institution.
For the Selachian of which these bones formed a portion, the name of Lamna
daviesii was proposed. ‘The old genera, Otodus and Lamna have now been united, {
and in consequence the teeth before mentioned must in future be known as Lamna
appendiculata. What relation they may bear to the vertebrie of L. daviesii remains
for the future to prove.
No other remains of Fish have been described from the Rolling Downs
formation.
* Quart. Journ. Geol. Soc., 1870, XXVI., p. 238.
+ Quart. Journ. Geol. Soc., 1872, XXVIII., p. 346.
t Reports on the Geological Features of Parts of the District to be traversed by the proposed Trans-Continental
Railway. Queensland Parl. Papers, 1885 (pp. 22, feap., Brisbane, by Authority, 1885), p. 8.
§ R. Etheridge, Jun. Appendix to R. L. Jack’s Report, loc. cit.
|| Proc. Linn. Soc. N.S.W., 1888, III (2), Pt. I., p. 156.
4; A.S. Woodward, Cat. Foss. Fishes Brit. Mus., 1889, p. 392.
joy)
ROLLING DOWNS FORMATION (CRETACEOUS) OF CENTRAL QUEENSLAND.
3. Belonostomus sweett, sp. Nov.
The state of preservation of the large fossil does not permit any precise estimate
of the original proportions of the fish. Judging, however, by the analogy of the
allied Brazilian species, the trunk cannot have been less than 0°9 in length, with a
maximum depth of 0-14 in the abdominal region ; while the head, with the opercular
apparatus, may have added about 0-4, making the total length approximately 1:3.
The individual under consideration is thus and by far the largest member of the
genus hitherto discovered.
(a.) Head and Opercular A pparatus.—As shown by the fragment M.31, the cranium
is well ossified in the occipital, otic, and prefrontalregions. The exoccipital extends
considerably into the lateral wall of the brain-case, and is bounded in front by a large
pro-otic element, separated only by suture, extending downwards to the base of the
cranium, and exhibiting near its hinder border a large oval foramen, evidently for the
exit of the glosso-pharyngeal nerve. The membrane bones of the cranial roof are
thick; and there is a robust parasphenoid bone, expanding in front of the orbit. The
jaws are not preserved, and the only element of the suspensorium exhibited is the
hyomandibular, which lies beneath the opercular apparatus of the left side in the
large fossil. As usual in Belonostomus, this bone is lamelliform, with a much
expanded, squamous, inferior moiety, and a narrow, more robust, superior portion,
from which at the hinder border would arise a short, stout process for the articulation
of the operculum. Externally, remains of large suborbital membrane bones
(Fig. 1, so.) are conspicuous between the orbit (orb.), and the preoperculum (p.0f.) ;
these being ornamented with coarse tuberculations rarely fused into irregular ruge.
The preoperculum (f.0f.) is a deep triangular bone, tapering, though notably
robust, above, and terminating in a truncated expansion below. Its upper extremity
reaches a point not far below that of the operculum, and the length of its inferior
border exceeds half the height of the bone. The operculum (of.) is approximately
quadrate in form, with a somewhat truncated and rounded postero-superior
angle. Its maximum depth and breadth are nearly equal; the upper border
is somewhat turned inwards, and the articular pit (a) for the reception of
the process from the hyomandibular is especially prominent near the antero-
superior angle. The sub-operculum (s.of.) is comparatively small, deepest and
truncated in front, gradually tapering to a posterior apex at the postero-inferior angle
of the operculum. All these bones are superficially ornamented with prominent
tuberculations of ganoine, which are partly fused into irregular radiating series, but
still more conspicuously arranged in concentric lines.
(b.) Branchial Apparatus.—As shown by the small fossil M. 26 A, the gill-arches
are slender, and shaped like those of modern Teleosteans, being a series of narrow,
4 ON THE OCCURRENCE OF THE GENUS BELONOSTOMUS IN THE
elongated lamin, —-shaped in transverse section. To each arch is affixed a close
series of long, slender, compressed appendages, only superficially calcified, and
apparently to be interpreted as supports of the gill-filaments.
(c.) Axial Skeleton of the Trunk.—The vertebral centre (Figs. 2 and 3) are well
ossified throughout, and the slender neural and hemal arches, with their spines seem
to be fused to them. They are deeper than long, and so far as can be determined
from impressions of the terminal coneavities (Fig. 3), the notochord was completely
constricted, no central perforation of the bone being distinguishable. A secondary
development of longitudinally-fibrous bone completely surrounds the primitive
double-cone of each centrum externally, and imparts to the vertebra the robust
appearance well shown in Fig. 2.
(d.) Appendicular Skeleton.—Of the fins, only fragmentary remains of the
pectoral and caudal are preserved in the large fossil (Fig. 1). The rays are all robust,
laterally compressed, undivided for some distance proximally, but soon branching
and marked by numerous articulations. Of the supporting elements, nothing can be
discerned, and even of the membrane bones accompanying the pectoral arch there
remains little evidence. It is clear from the small fossil, M. 28, that there was a
pair of large supra-temporal bones externally ornamented with tubercles fused into
radiating and somewhat reticulating ruge. The same specimen also exhibits a small,
triangular, supraclavicle, adjoining the truncated postero-superior angle of the
operculum, and ornamented with tubercles of irregular size, partly fused into
concentric lines. Of the clavicle and pectoral arch itself, however, there are no
fragments sufficiently worthy of note.
(e.) Squamation.—As usual in Belonostomus one series of flank scales is
excessively deepened, these being abruptly truncated below, but slightly rounded
and reflexed forwards at the upper extremity. A restoration of one example
from the anterior abdominal region is given, of the natural size, in Fig. 4. This
scale is about five times as deep as broad, crossed at its flexure by the lateral line ;
and the overlapped portion is relatively narrow. Immediately posterior to the
hinder margin of the overlapping scale is an area of nearly smooth ganoine, bounded
by a nodose longitudinal ridge. A parallel and more prominent longitudinal ridge,
separated from this, and from the hinder area by a furrow, divides the scale into two
nearly equal halves, and the posterior half is much ornamented. This ornament
consists first of large irregular tubercles, passing behind into small, nearly parallel,
though sometimes bifurcating transvere ridges, at right angles to the posterior
border. ‘The scale immediately below the principal flank series is much deeper than
broad, and then follow about four small vential scales, equally broad, but their depth
not exceeding half their breadth. One of the latter is shown, somewhat restored, in ~
Fig. 5; its overlapped superior border, with the large articular spine, is conspicuous,
ROLLING DOWNS FORMATION (CRETACEOUS) OF CENTRAL QUEENSLAND. 5)
and there are some superficial ruge in the lower half of the scale parallel with its
inferior border. The squamation above the principal flank series is partly shown in
the hinder half of the trunk, and some of the scales are well preserved (Fig. 6).
There are at least two series between the upper extremity of the principal scale and
the dorsal azygous series, both comprismg imbricated rhomboidal scales, well
ornamented with ruge of ganoine, and each with a prominent oblique ridge in the
middle of its exposed portion, almost diagonal; the ruge are partly concentric
with the hinder and inferior borders, partly at right angles to the former. Towards
the extremity of the tail the flank scales gradually become less deep in proportion
to their breadth, and at the origin of the caudal fin, all the scales of each ventral
row are of nearly equal size (Fig. 7), rhomboidal, and closely resembling the
ordinary dorso-lateral scales, except that the diagonal has become more prominent,
and few of the rugz are concentric with the borders.
4,—Sprciric DETERMINATION.
In the absence of definite information as to the proportions of the species now
described, it is necessary to refer exclusively to the scales for diagnostic characters.
It seems, however, almost certain that we are concerned with a much less slender
fish than the typical Jurassic species, Belonostomus tenutrostris, B. sphyrenoides,* &c.;
while neither of these, so far as we can discover, exhibit so conspicuous and elaborate
a superficial ornamentation. It thus suffices to refer only to the known
Cretaceous species, which may be enumerated as follows :—
Belonostomus attenuatus, Dixon, Geol. and Foss., Sussex, 1850, p. 368, Pl. XXXV.,
f. 4 [Portion of mandible], Chalk, Sussex.
Belonostomus cinctus, Lu. Agassiz, Poiss. Foss., 1844, Vol. IL., Pt. IL., p. 142,
Pl. LXVIa, Figs. 10—13; F. Dixon, op. cit., p. 367, Pl. XXXYV., Fig. 3;
A. 8. Woodward, Quart. Journ. Geol. Soc., Vol. XLIV., p. 145, Pl. VIL,
Figs. 7—13, and Proce. Geol. Assoc., Vol. X., p. 805.—Chalk; S.E. England.
(Mandible and scales. ]
Belonostomus comptoni, Agassiz, A. 8. Woodward, Proc. Zool. Soc., Nov. 18th,
1890. Aspidorhynchus comptoni, Li. Agassiz, Edin. New Phil. Journ.,
1841, Vol. XXX., p. 83, and Comptes Rendus, 1844, Vol. XVIIL.,
p. 1009.—Upper Cretaceous; Serra de Araripe, North Brazil. {Imperfect
fishes. |
* L. Agassiz, Poiss. Foss., II., Part 2, p. 140; A. Wagner, Abth. Math. Phys. Cl. K. Akad. Wien, 1863, IX.,
p- 689.
6 ON THE OCCURRENCE OF THE GENUS BELONOSTOMUS IN THE
Belonostomus crassirostris, O. G. Costa, Pal. Regno Napoli, Pt. II., p. 33,
Pl. IL., Figs. 1, 2. Belonostomus gracilis, O. G. Costa, ibid., p. 35,
Pl. Il., Fig. 8.—Upper Cretaceous; Pietraroja, near Naples. [Imperfect
fish. ]
Belonostomus (?) indicus, A. S. Woodward, Rec. Geol. Surv. India, 1890, Vol.
XXIII., p. 28. Upper Cretaceous (Lameta beds) ; Dongargdon, Nagpur,
India. [Imperfect skull and mandible. |
Belonostomus lesinensis, F. Bassani, Denkschr. math.-naturw. Cl. k. Akad.
Wiss. “Wien, “1882; Vol OULV qt) 198.0 Ply a. Pao MOx? Tipper
Cretaceous; Isle of Lesina, Dalmatia. [Imperfect fish.]
Of these species B. attenuatus, and B. ? indicus, are not strictly comparable,
being known only by parts of the head; but the absence of ornamentation on these
fossils renders most improbable any intimate relationship with the Queensland fish.
The feeble character of the ornament of B. cinctus, which is known by scales, as
well as jaws, also excludes this species from comparison. 8B. crassirostris, and B.
sleinensis are small slender fishes, apparently as much elongated as any of the
Jurassic species. Indeed, the only form which seems to approximate at all closely
to the species now under consideration is the large B. comptoni from Brazil. The
few proportions that can be compared are very similar, and the superficial
ornamentation in adult fishes is equally conspicuous. The principal flank scales of
B. comptoni, however, never appear to exhibit the prominent, fine, transverse
striations, so characteristic of the hinder margin of all the flank scales of B. sweeiz,
and we thus venture to regard the latter as specifically distinguished by its superficial
ornament.
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ROLLING DOWNS FORMATION (CRETACEOUS) OF CENTRAL QUEENSLAND.
Pa Ae ae
DeEscRIPTION oF FiGurReEs.
Fig. 1.—Portion of fish, with remains of opercular apparatus.
op. operculum.
p.op. pre-operculum.,
s.op. sub-operculum:
orb. orbit, half nat. size.
Fig. 2.—Portion of a vertebra, side view, with parts of the neural and
hemal arches.
Fig. 3.—The same, end view.
Fig. 4.—Restoration of a scale from the anterior abdominal region.
Fig. 5.—A ventral scale, somewhat restored.
Fig. 6.—Scales from above the hinder principal flank series.
Fig. 7.—Scales of one of the ventral rows at the origin of the caudal fin.
ARTICLE II.—On tHe Srructure or CERATELLA Fusca (Gray), By W. BaLpwin
Spencer, M.A., Proressor or Bionogy In THE University or MELBOURNE.
(With Plates 2, 3, and 3a.)
(Read Thursday, June 11th, 1891.)
IT have to thank Dr. Ramsay and the Trustees of the Australian Museum,
Sydney, for the opportunity of examining the structure of this interesting hydroid
form. The specimens examined came from Bondi on the New South Wales coast
and from Lord Howe Island, where they were collected by Mr. Whitelegge of the
Australian Museum, whom I have to thank for kind assistance. The Lord Howe
Island specimen had the zooids beautifully expanded.
Dr. Gray* was the first to describe and figure Ceratella and the closely allied
genus Dehitella from specimens in the British Museum. He had dried specimens
only at his disposal and his description necessarily refers only to the hard parts
and these having the form of a horny network somewhat resembling in general
appearance the skeleton of a horny sponge, led him to place the two forms
provisionally amongst the sponges. For their reception he constituted the family
Ceratellade. Five years later Mr. H. J. Carter+ published a paper entitled
“Transformation of an Entire Shell into Chitinous Structure by the Polype
Hydractinia, with short Descriptions of the Polypodoms of five other Species.” In
this he refers to Ceratella fusca and Dehitella atrorubeus and describes two new forms
belonging to the former genus under the names of C. procumbens and C. spinosa.
Both of these came from South Africa the former from the Cape of Good Hope and Natal
the latter from Natal. In the same paper he describes a new genus Chitina with a
single species C. ericopsis which came from New Zealand. Reference to these will
again be made after the anatomy of Ceratella fusca has been described, meanwhile
it is sufficient to note that Mr. Carter’s investigations of a species of Hydractinia
(H. levispina) with a skeleton composed of a horny network which incrusts and eats
its way into univalve shells led him to re-examine the two forms placed by Dr. Gray
in the family Ceratellade and in consequence of his finding undoubted traces of
thread-cells in the dried specimens both of these and of the two new species
of Ceratella mentioned above, he rightly recognised them as belonging to the
* Proc. Zool. Soc., Nov., 1868.
+ An. and Mag. Nat. Hist., Jan., 1873.
ON 'THE STRUCTURE OF CERATELLA FUSCA (GRAY). 9
Hydroidea and not to the sponges in which group Dr. Gray had provisionally placed
them. At the same time it may perhaps be well to note that the finding of
thread-cells in dried specimens with a skeleton in the form of a horny network
is not in itself absolute proof of their belonging to the Hydroidea inasmuch as at the
present time sponges are known to exist the substance of which is pierced by hydroid
erowths so that it would be possible to find thread-cells in the dried skeletons of
sponges. At the same time Mr. Carter was perfectly right, as events have proved, in
removing the Ceratellade from the Porifera though the subsequent discovery by
Mr. Bale of the nature of their soft parts has shown that they cannot be placed,
where Mr. Carter put them, in the family Hydractinude.
The next notice of these forms occurs in the Proceedings of the Linnean Society
of New South Wales for 1886 (p. 575) when Mr. Brazier recorded the occurrence of
Ceratella fusca from various localities near Sydney such as Bondi Bay and Coogee.
In the Proceedings of the same Society for the year 1888 (p. 745) Mr. Bale for the
first time gave some description of the soft parts and showed that the zooids were
formed on a very different type from those of the Hydractiniide. The latter have a
single circlet of filiform tentacles surrounding the hypostome whilst those of Ceratella
are irregularly distributed over the body and are capitate. Mr. Bale accordingly
removed Ceratella from its position amongst the Hydractiniide assigned to it by
Mr. Carter and placed it in a distinct -family to which he gave the name Ceratellide.
He apparently overlooked the fact that Dr. Gray had already adopted the name of
Ceratellade for the family including his two genera Ceratella and Dehitella, so that
this name given in 1868 must now be retained. With a more abundant supply of
material I have been enabled to work out the structure in greater detail than was
possible to Mr. Bale to whom we owe the first description of the soft parts and
the determination of the fact that Ceratella belongs to a family distinct from the
Hydractiniide.
DeEscRIPTION OF THE STRUCTURE OF CERATELLA FUSCA.
I have endeavoured so far as possible to give a complete account of the anatomy
of both soft and hard parts the only figures yet published being those of the external
form given by Dr. Gray* both of which suffice to clearly identify the genera.
Skeleton.
The colonies of Ceratella procured on the New South Wales coast measure from
14—5 inches in height and are of a rich brown colour. The largest specimen which
Ihave seen is the one procured by Mr. Gabriel which came from Flinders Island
in Bass Straits where it had been washed ashore. ‘The figure given by Dr. Gray
Maca Clits
10 ON THE STRUCTURE OF CERATELLA FUSCA (GRAY).
represents fairly well the macroscopic characters of the skeleton but conveys the idea
of one which has been much water-worn and has lost many of its smaller branches.
The branching is much richer and closer than is represented by him and the
projecting hydrophores are more distinct and regular. At the same time there is a
certain amount of difference in the growth of various colonies some of which are
more bushy in appearance than others. That secured by Mr. Gabriel has the
branches more distinctly arranged in one place than is the case with the others.
The whole colony may be described as follows:—The colony arises from a much-
branching root-like base encrusting foreign objects. The root-branches unite to
form a strong stem common to the colony which is flattened in the same plane as
that in which the branches of the colony, generally, expand. The common stem
may have the appearance of being formed of intertwined branches and from it arise
irregularly larger or smaller branches the former being more or less flattened in the
same plane with itself.
From the larger arise irregularly on either side smaller branches which again
branch until a somewhat bush-like or fan-shaped colony is formed the whole being
more or less flattened in the plane of the maim stem. Except the main stem close
to the root branches and the latter all the branches may bear bracket-like
projections—the hydrophores—which are arranged in a roughly spiral manner and
are especially abundant on the smaller branches except the growing ends which are
somewhat swollen and flattened in a plane at right angles to that in which the
branching of the colony takes place.
In addition to considerable variations in the general form of the colony some
appear to have the hydrophores so arranged that they le along the two opposite sides
only of the branches whilst in others they are arranged all around. The more
bush-like the growth is, to the greater extent does the latter obtain and vice versa.
Usually there is only one main stem arising from the root-branches but at times one
or more smaller ones may arise independently. (Fig. 1.)
The skeleton when examined by the lens has much the same appearance as that
of Hydractinia and differs considerably from that of Clathrozoon and more still from
that of the polysiphonic stem characteristic of certain species of Plumularia. It forms
in the larger branches a meshwork of chitinous tissues so similar at first sight to
that of certain horny sponges that as Mr. Bale says ‘‘a portion broken off and
examined separately might well be mistaken for sponge tissues.” ‘This however is
only true with certain limitations for, if the soft parts are present, the distinction
between the two is readily seen as is also the case when the hydrophores are present.
Apart from this also nothing comparable to the layer of cells secreting the sponge
skeleton is present nor are the chitin fibres of the two similar in their minute
_ structure.
ON THE STRUCTURE OF CERATELLA FUSCA (GRAY). el
The meshwork is extremely irregular in the larger branches as is shown in the
sections represented in figs. 9 and 11. There seem to be two not clearly
distinguishable portions present (1) large and strong fibres (2) smaller ones which
form connecting bars and often have the appearance of thin web-like plates which
are especially well developed on the hydrophores. Fig. 5 represents a medium-sized
branch, from which the soft parts have been removed by potash, seen by reflected
light. At times the fibres seem to run for some distance parallel to the long axis of
the branch though this is much more strongly marked in some branches than in the
one figured. There are no definite hydrothece present within which the zooids can
be completely retracted but the branches are studded sometimes only at the sides
sometimes all over with little bracket-like projections which give a characteristic
‘serrate appearance to the branches, very different however from that which is
produced by the hydrothecx of the Sertularians. Mr. Bale has aptly suggested that
the term ‘‘ hydrophores” (originally applied by Allman to the calyces of Halecium)
should be used to describe these structures which form merely supports for the
hydroid zooids. Each one may perhaps be best likened to a very concave scallop
shell with ribs formed by the strong fibres continuous with those of the branch whilst
the spaces between them are filled up by a thin fenestrated web of chitin. In some
hydrophores the ribs are more strongly marked than in others and project as small
points around the margin. (Figs. 5 and 6.) The growing ends of the smaller
branches usually contain two or three longitudinal fibres connected by transverse
bars, often somewhat web-like, and each of these growing ends is clearly
distinguishable (1) by its being flattened in a plane at right angles to that in which
general growth takes place and (2) by the entire absence of hydrophores and zooids.
It may be added that the longitudinal arrangement of the fibres in the branches is
more clearly noticeable in specimens with the soft parts present than in those in
which the latter are absent since the soft parts conceal from view largely the
connecting and the deeper-lying fibres and thus prevent to a large extent the network
structure from being seen. The whole skeleton also in living specimens is
completely enclosed by the soft parts though only a thin layer of tissue is present
over the external surface. (Figs. 9, 10and11H.) There is no protective covering
for the reproductive structures.
Soft Parts.
The structure of these has, as yet, been only briefly described once and that by
Mr. Bale, whose short account refers only to the external form of the hydroid zooids.
He pointed out, as stated before, that these possessed irregularly scattered capitate
tentacles and that hence they differed considerably from those of Hydractinia.
Amongst the specimens from the Australian Museum is one collected by
Mr. Whitelegge on Lord Howe Island with the zooids fully expanded and others
with the soft parts well preserved.
12 ON THE STRUCTURE OF CERATELLA FUSCA (GRAY).
(1). Hydroid zoords (Figs. 2, 4, 6, 7, 9, 13).—These may be found on all the
branches, large and small except at the growing ends. Often also they are absent on
the larger main branches and are always fewer in number on these than on the
smaller ones (Fig. 4). Each zooid, in a well preserved spirit specimen, reaches a
length of about 1-4 m.m. and in the general form of the body resembles, as Mr. Bale
says, those of Coryne. The body (Figs. 4 and 6) is somewhat elongate with a
terminal almost conical portion, at the apex of which lies the mouth opening. The
basal portion which is seated upon the hydrophore is broad, this is followed
by a shghtly contracted portion, then comes a slightly swollen part which gradually
diminishes in size towards the mouth end. Over the surface are scattered irregularly
the capitate tentacles from 10-14 or perhaps even more in number (Figs. 4 and 6),
and one or two of these are frequently placed close to or upon the basal region.
The minute structure is generally that which is typical of hydroid forms and is
represented in Figs. 2 and 7. The ectoderm (£ct.) is unilaminar over the general
surface the cells being in close apposition each with a large nucleus and except in
the capitate ends of the tentacles there are no thread cells present. At the ends of
the tentacles the ectoderm is swollen out and apparently forms a mass of cells in
which large nematocysts are present with barbed threads (Fig. 7). The nematocysts
here and elsewhere seem to be all of the one size. The ectoderm lies on a thin layer
of mesogloea scarcely visible in extended zooids but more prominent in retracted
ones. There cannot be detected any fibrous muscular elements such as form so
distinct a feature around the basal region of the zooids in, for example, Clathrozoon.
The endoderm consists of large vacuolate cells each one subtending the base of, as a
general rule, at least three ectoderm cells. The protoplasm appears to be always
concentrated at the inner ends where the nuclei are placed and where in preserved
specimens the cell outlines are completely lost. In zooids which are feeding
(Fig. 7) this inner end of the cell is filled with minute food particles the digestion
being evidently, in part at least, intracellular.
The ectoderm is continuous at the base with the layer which covers externally
the whole colony whilst the endoderm is continuous with that of two or more of the
coenosare tubes.
(2.) Gonophores.—I have only been able so far to find the male gonophores
which are present on three colonies secured at Coogee on the New South Wales
coast. Hach of these colonies carries numbers of minute somewhat pear-shaped
structures which are only from one-third to one-quarter of the length of the hydroid
zooids and which are seen when rendered transparent or cut into sections to be
medusoid in nature.
ON THE STRUCTURE OF CERATELLA FUSCA (GRAY). 13
These gonophores arise directly from the cenosarc and are not carried by modified
zooids or blastostyles.
They may be very numerous indeed especially on the medium-sized branches
where their number in one specimen far exceeds that of the hydroid zooids. Two
are represented as seen by reflected light in figure 6 and in all specimens examined
the gonophores are at the same stage of development. I have been unable to detect
any indication of the formation of reproductive elements in the hydrophyton.
The gonophores much resemble in structure those figured by Weismann* in
Pennaria cavolina or Cladocoryne floccosa. In essential structure the transverse section
of the gonophore of the former is identical with that of a similar section of a
Ceratella gonophore as shown in figure 8. The longitudinal section again of the
gonophore of Cladocoryne or of Pennaria as figured by Weismann agrees almost
precisely with that of Ceratella represented in figure 12. In the latter the
manubrium is well developed and surrounding this lie the reproductive elements
which in longitudinal section (Fig. 12) form a horseshoe-shaped mass and in transverse
section aring. External to the latter isa thin layer of ectoderm (ect.') which comes
in contact with the external layer of ectoderm (ect.) at the part corresponding to the
mouth of the medusa bell where the former layer dips inwards. (Fig. 12, M.) This layer
must correspond to the sub-umbrella ectoderm of the medusa and the special point
mentioned indicates also the position at which, in development, the “ glockenkern ”’
of Weismann grew in by proliferation of the ectoderm cells. I have been unable to
find any gonophore younger than the stage figured though many have been cut in
section, the manubrium being in every case well developed.
Between the two layers of ectoderm lies the endoderm with four radial canals
seen clearly in transverse sections (Fig. 8) whilst in longitudinal sections (Fig. 12)
the indication of a ring-shaped space around the distal end can always be detected.
Hach gonophore may be connected with more than one of the ccnosare tubes
and its ectoderm is continuous with that which covers the colony externally.
(3.) The Hydrophyton—This may be divided into two parts (1) the external
layer common to the whole colony and (2) the branching network of tubes.
The external layer is formed entirely of ectoderm cells (Figs. 3 and 12) and is
only one cell thick though it may come into contact with the ectoderm of the tubes
lying immediately beneath it. It is especially well marked in the younger branches
and may be worn away to a greater or less extent in older ones though typically it
forms a covering for the whole of the colony (Figs. 9 and 13). It is directly
* Die Entstehung der sexualzellen bei den Hydromedusen. Pl. XVII., Figs. 1-5 and Fig. 7; Pl. XVIII., Fig. 1.
14 ON THE STRUCTURE OF CERATELLA FUSCA (GRAY).
continuous with the ectoderm of both the hydroid zooids and the gonophores, and
has in many respects a close resemblance to the superficial layer of ectoderm as
described and figured by Professor Moseley in Millepora.* In the latter the exact
relationship of the superficial ectoderm to the zooids could not be ascertained but
in Ceratella where the latter are not retracted imto spaces within the skeleton its
direct connection with the ectoderm of the zooid can easily be seen in sections.
Figure 8 represents a small portion of the layer as seen under a high power. The
outlines of cells cannot be definitely distinguished in the specimen in question
though a somewhat light space with a fairly distinct outline surrounds the thread
cells. The inner ends of the cells are in contact with the ectoderm of the tube
beneath and the layer thins out just where it passes over the projecting point of a
portion of the skeleton. In younger branches (Fig. 14) the cells of the layer are much
more definite in form and outline being each cubical with a distinct nucleus whilst
comparatively very few thread-cells are present.
This superficial layer is only known to exist m the Hydrocoralline, the
Hydractiniide and, now, in the Ceratellade. In the last mentioned the soft parts of
only Ceratella fusca ave known as yet but the skeleton of Dehitella is so closely
similar to that of the former that we may with much probability infer that a close
agreement exists in the nature of their soft parts. The ccenosare tubes form a
richly branching network of tubes occupying all the spaces in the chitinous
meshwork which forms the skeleton. In Figures 9 and 10 this is represented
diagrammatically by the grey colour the former being a longitudinal and the latter
a transverse section of a branch. Throughout the whole system the endoderm is
never more than one cell thick whilst the ectoderm is very irregular. Very often the
endoderm cannot be recognised or else it forms an indistinct layer which stains more
darkly than the ectoderm and contains no space, a result probably due to the action
of reagents. At other times (Figs. 8, 11, 12, and 14) a distinct tubular space
can be distinguished. The number of tubes varies naturally according to the size of
the branch. Figure 13 represents a longitudinal section through a portion of a small
branch with a smaller offshoot which formed part of a specimen brought by
Mr. Whitelegge from Lord Howe Island. The general appearance of a part of the
same specimen viewed as a solid object is represented in Figure 4. This particular
specimen has the branches much finer than those of the others and the skeleton and
ccenosarc tubes somewhat more regular in arrangement, whilst the hydrophores
are not very strongly developed. Up the small offshoot pass the main skeletal ribs
united by cross bars which are thin and almost web-hke and up the centre runs a
single ccenosare tube (B) which is connected with at least three of those in the larger
branch. This tube consists of an internal layer of endoderm (Fig. 18 a, end.) and
* On the Structure of a Species of Millepora occurring at Tahiti, Society Islands. Phil. Trans. R.S. London,
1876, Vol. CLXVII., p. 117.
ON THE STRUCTURE OF CERATELLA FUSCA (GRAY). 15
an irregular external layer of ectoderm (ect.) the former beg continuous with that of
the zooids. The whole is covered by aunilaminar ectoderm (£). <A few thread cells
are present. Up the layer branch the tubes run more regularly than usual and three
or four may be traced for a considerable distance running parallel to its length but
eiving off lateral branches. In sections both transverse and longitudinal of Ceratella
these connecting bars or webs crossed by ccenosare tubes form a very characteristic
feature (Figs. 11 and 138. C).
As stated above there is a strongly-marked difference between the endoderm and
ectoderm. The former (Fig. 3) is regular and takes stain somewhat more readily
than the latter, which is often very irregular and several cells thick, though most
often the outlimes of cells cannot be recognised, and a structure resembling a
syncytium is formed. In the latter are found (1) nuclei, (2) thread cells, (3) bodies
surrounded by a clear space and staining evenly and deeply (Fig. 3A). The
thread cells are apparently confined to the ectoderm, though of this I cannot feel
absolutely sure, and are found in great abundance in the inner parts of the branch
whence they must migrate to the surface if they are to be of service to the colony.
It is a somewhat curious fact that they are as a rule present in far greater numbers
in the ectoderm of the coenosarc tubes than in the most external layer. Figure 3
represents a small portion of the latter on a part of a colony where the
gonophores were numerous and here thread cells were present in greater numbers
than elsewhere. Of the nature of the third-named structures it is difficult
to be certain but it is probable that they are ectoderm cells in which thread
cells are being formed. ‘There is at all events a curious agreement in appearance
between them and the structures which Professor Moseley has described as developing
thread cells in Millepora.* He says “the thread cell appears to be developed out of
the nucleus of the ectodermal cell, the ectodermal cell becoming much enlarged and
forming a wide chamber, in which the process of development takes place. The
ovoid nucleus becomes enlarged together with the cell, but not at all in the same
proportion the cell always appearing as a wide cavity around it. The nucleus as it
enlarges has a rounded nucleolus developed at one end of it.’ The nucleolus has
large granules developed within it, whilst the nucleus becomes finely granular. In
the next stage one large coil of the thread appears in the nucleus.”
The earlier stage seen in Millepora when the nucleus with nucleolus at one
end of it lies in the cell which forms a clear cavity around it, corresponds exactly to
that represented in figure 3A.c. in the case of Ceratella. Though a complete series of
stages could not be obtained still those drawn in figure 3A. will serve to show that in
all probability Ceratella resembles Millepora closely in the formation of thread cells.
In figure 3A.a. the cell is small and the nucleus but little larger than that of an
* Toc. cit., p. 129.
16 ON THE STRUCTURE OF CERATELLA FUSCA (GRAY).
ordinary ectoderm cell though stained very deeply and having a homogeneous
appearance ; in b. the cell has increased in size, the nucleus is much larger, and has
a clear space all around it between it and the cell wall; this is clearly marked in c.
and d. where, in the former, a nucleolus is present and in the latter two darker thread-
like lines possibly indications of the commencing formation of the thread; in ec. what
is evidently a very young thread cell is seen—it is somewhat darkly stained without
a definite thick wall such as is seen clearly in later stages, and down the centre is a
lighter line corresponding to the thicker attached part of the thread. It has also the
shape of the thread cell but there is no trace of the clear space present in earlier
stages, a certain amount of stained protoplasm being attached to it. Inf. and g. two
later stages are shown in which the thick cell wall is present and the coiled thread
can be clearly seen. ‘These thread cells evidently resemble closely in structure the
three-barbed ones described by Professor Moseley in Millepora.
The only other point to notice in regard to the ccenosare is the structure of the finer
erowing branches which are somewhat flattened out. A longitudinal section of one
of these is represented in figure 14. Up the centre runs a ccenosare tube with a large
cavity and clearly-marked endoderm the ectoderm being as usual irregular. From
this central tube short branches are given off (D) which run outwards towards the
external layer with which, as at the point x, they may come into direct contact. At
this point the cells of the two layers are well marked, and in all probability this
shows us the earliest stage in the formation of a zooid. It has already been noted
that the ectoderm of the latter is in direct connection not with that of the ccenosare
tubes but with the common external layer and this method of formation would
explain this otherwise somewhat curious fact. ‘The endodermal process grows out
into a bud—the early stage of a zooid—carrying with it the external ectoderm layer
which thus, as further growth takes place, naturally gives rise to that of the zooid
itself. At the same time the ccenosarc tube branches as the stem increases in size
and thus the zooid, if the branching be near the base of the latter, will become
connected with two or more tubes.
AFFINITIES OF THE CERATELLAD®.
When Dr. Gray first described these forms there were only two specimens
available neither of which possessed the soft parts. The hard parts whilst agreeing
in important points differed from each other sufficiently to be regarded by him as
species of two distinct genera Ceratella and Dehitella.
Mr. Carter, with more material at his disposal, recognised the fact that they
were hydroids and owing to similarities in their skeleton and that of Hydractinia
placed them in the family Hydractiniide, thus abandoning Dr. Gray’s family
Ceratelladee which had been created under the assumption that the two forms were
allied to the sponges.
ON THE STRUCTURE OF CERATELLA FUSCA (GRAY). 17
Undoubtedly in many respects Ceratella and Dehitella call to mind the
Hydractiniide, but it is doubtful if even our knowledge were confined to that of the
structure of the hard parts whether Mr. Carter’s classification could be upheld. ‘The
one point of resemblance—and at first sight it is the most striking feature—is that
the skeleton of both consists of a very irregular branching chitinous network. In the
Ilydractiniide however this has the form of an encrusting network with at most very
feebly developed branches arising from it; these may more correctly be described as
spines and they do not appear to carry any zooids. In the Ceratelladx the whole
colony consists of a much-branching structure arising from a comparatively small
encrusting root-portion which may itself be made up of branches more or less
entwined. In addition to this all the branches bear hydrophores or special
developments of the network to support the hydroid zooids. These are never
present in the Hydractiniidze but always in the Ceratelladee. Now that the soft parts
are known there can be no doubt about separating the two families. The hydroid
zooids are quite different those of Ceratella being provided as are those of Coryne
with scattered capitate tentacles whilst there is no trace of protective zooids such as
are present in Hydractinia and Podocoryne. In addition to this the gonophores
arise directly from the ccenosare and not from modified zooids. ‘The most important
points of agreement lie in (1) the existence in Hydractinia and Ceratella of a
common external layer of ccenosare which covers over the whole skeleton mass
whether this be encrusting or branching in nature; (2) the presence in both of a
network of ccenosare tubes forming the hydrophyton. It may however be noted
that in both these points we find a similar agreement to exist between Ceratella and,
for example, Millepora amongst the Hydrocorallinz as between the first named and
Hydractinia.
The presence of this external layer which, in the Hydrocoralline and Ceratella
at all events, consists simply of a layer of ectosarc is very difficult to explain.
Professor Moseley* has represented it in Millepora as if it formed the outer layer of
the surface ccenosare tubes though even in this case it passes over all the
parts (occupied by the calcareous skeleton) which on the surface he between the
tubes, and is very different in appearance and in the relative size of its cells from
the ectoderm which elsewhere forms the onter wall of the tubes. In Ceratella it is
perfectly independent of the tubes all of which have their own ectoderm covering
though at the surface this comes in direct contact with the outer layer. I am not
aware of any determination in Hydractinia of the exact relationship of this outer
layer though very probably it will be found to agree with that of Ceratella.
It is not apparently connected in any special way with the formation of the
chitinous network as this ies deep within the structure of the branch, and the only
* Loe. cit., Pl. 3., Figs. 10 and 16.
18 ON THE STRUCTURE OF CERATELLA FUSCA (GRAY).
purpose which it can apparently serve is that of a covering layer which prevents
foreign objects from passing in between the meshes of the network and interfering
with the general welfare of the colony. It is strange however to note, if this be its
function, that most usually the internal ccenosare contain a far greater number of
thread cells than this external layer does in Ceratella.
In Millepora Professor Moseley was unable to determine its exact relationship to
the zooids but in Ceratella by its means all the ectodermal structures lying on the
external surface are brought into direct continuation with one another.
It may be noted in passing that though in the genus Clathrozoon* the branches
of the colony are formed of a somewhat similar network of soft parts there is nothing
present resembling this external layer the whole branch being in this instance covered
with a thin protective perisare,
Taking both the hard and soft parts we find the following points of agreement
to exist between the Hydractiniide on the one hand and the Ceratellade on the
other though it must be borne in mind that we only know as yet the structure of the
soft parts in one member of the latter family.
(1.) The skeleton has the form of a branching chitinous network.
(2.) The hydrophyton consists of a network of freely branching and
anastomosing ccenosarcal tubes.
(3.) The zooids arise directly from this network and no true hydrothece
or gonothece are formed.
(4.) A common external layer is present enclosing the whole colony.
The two differ from one another in the following points :—
(1.) Hydractiniide form encrusting masses with at most spmulose branches
arising from the surface which do not bear zooids. The Ceratellade
always form freely branching masses either erect or procumbent: the
basal part which serves to attach the colony being alone of an
encrusting nature whilst even this has the form of intertwined
branches.
(2.) The Ceratellade always possess hydrophores or special developments of
the skeleton which serve asa support for the basis of the hydroid-
zooids and nothing similar to which is found in the Hydractiniide.
(3.) The hydroid zooids Ceratellade possess scattered capitate tentacles
those of the Hydractiniide being filiform and arranged in a single
circle beneath the mouth.
* Trans. R.S., Victoria, 1890, p. 121. Pl. 18, Fig. 3; Pl. 19, Fig. 12.
ON THE STRUCTURE OF CERATELLA FUSCA (GRAY). 19
(4.) The gonophores of the Ceratellade arise directly from the ccenosare
and are not developed on special zooids as in the case of the
Hydractinide.
Whilst the poimts of agreement detailed above serve to show a general
resemblance between the members of the two groups those of difference are of
sufficient importance to justify their separation into two distinct families.
As stated previously Dr. Gray’s name Ceratelladz will be retained and the
following gives the characters of the family (modified from Dr. Gray’s and Messrs.
Carter and Bale’s descriptions) and the list and characters of the genera and species
yet known.
Family Ceratellad@ (Gray, Proc. Zool. Soc., 1868, p. 575).
Forming branching colonies. Skeleton in the form of a chitinous network with
slight bracket-like or tubular projections (hydrophores) serving as a support for the
bases of the gastrozooids.
Hydrophyton a network of branching anastomosing tubes the whole enclosed by
a common ectoderm layer.
Gastrozooids naked.
Gonophores medusoid : fixed and arising directing from the hydrophyton.
Genus. Dehitella. (Gray.)
Colony dichotomously branched, expanded growing in a large tuft from a broad
creeping base. Stem cylindrical, smooth; branches tapering and cylindrical.
Hydrophores shghtly tubular and on the smaller branches divergent nearly at right
angles from the stem.
(1.) Dehitella atrorubens. (Gray.)
The description of the species is the same as that of the genus. It is known at
present simply from that given by Dr. Gray* who states that the genus “is
distinguishable from Ceratella by the greater thickness and cylindrical form of the stem,
by the more tufted and irregular manner of growth and by the tufts of spicules (oscules
or cells) being more abundant and equally dispersed on all sides of the branches and
branchlets.” The ‘“oscules or cells” of Dr. Gray must be the structures which,
following Mr. Bale, have been described above as ‘ hydrophores.”
Locality. —Delagoa Bay, Africa.
* Proc. Zool. Soc. 1868, p. 579. Fig. 1.
20 ON THE STRUCTURE OF CERATELLA FUSCA (GRAY).
Genus. Ceratella. (Gray.)
Colony irregularly branching; more or less expanded in one plane; growing
from a creeping base. Main stem flattened, branches rounded and beset with
bracket-lke hydrophores.
(2.) Cevatella fusca. (Gyay.)
Colony branching and fan-shaped ; expanded in the one plane; erect. Skeleton
consisting of alight or dark-brown chitinous network; the main stem broad and
flattened; branches numerous with the bracket-shaped hydrophores arranged on
them in a roughly spiral manner and formed of ribs continuous with the fibres of the
stem and united by thin perforated laminz the ribs projecting at the outer margin.
All the spaces within the chitinous network filled by a much branching hydrophyton
and the whole enclosed by an external layer of ectoderm. Gastrozooids seated on
the hydrophores, erect, with capitate tentacles tregularly scattered (10-14).
Gonophores medusoid, fixed.
Localities.—Coogee, Bondi (N.S.W.), Broughton Island, Flinders Island, Lord
Howe Island.
(3.) Ceratella procumbens. (Carter*).
Colony procumbent, thickly branched on the same plane; the larger stems
chiefly on one (the lower) side, hard, flexible, of an ochre-brown colour, tinged here
and there with purple. Trunk short, solid, compact, compressed vertically, soon
dividing irregularly or subdichotomously into round branches which are confined to
the lower surface, ending in branchlets with sub-clavate ends, that appear on the
upper or opposite side, not reuniting or anastomosing. Hydrophores consisting of
a little semitubular plate, extending outwards and forwards from the side of the stem
on the proximal border of an aperture in the latter; scattered thickly over all the
branches, but most prominent on the branchlets ; frequently represented by the little
hole alone in the stem where the projecting portion has been worn off; scanty on the
lower side of the main stems. Minute structure ; composed of clathrate chitinous
fibre throughout, whose meshes are subrectangular; hydrophore formed of the
semitubular scoop-lke plate mentioned supported on its proximal side by an extension
of the clathrate structure of the stem and bordering the little hole also above
mentioned, which extends into the centre of the stem; surface of the larger stems
bluntly microspined. Size of largest specimens 11 inches long by 5 inches broad,
and about 1 ineh thick or vertically.
Locality.—Cape of Good Hope, Natal.
* Ann. and Mag. Nat. Hist., 1873. Transformation of an entire shell into chitinous structure by the Polype
Hydractinia, with short descriptions of the Polypidoms of five other species (Pl. 1). The descriptions of C. procumbens,
C. spinosa and Chitina ericopsis are taken with only slight alterations from this paper.
ON THE STRUCTURE OF CERATELLA FUSCA (GRAY). 21
(4) Ceratella spinosa. (Carter.)
Colony procumbent; thickly branched hard flexible of a dark rich red-purple colour.
Main branches round, brownish, covered with small, smooth, often subspatulate erect
spines. Stem dividing subdichotomously into purple branchlets, which terminate in
abruptly pointed extremities. Hydrophores the same as in the foregoing species;
most prominent in the round branchlets to which they give, en profil, a serrate
somewhat sertularian appearance, the teeth of which are inclined forward. Minute
structure: Main stems composed of clathrate chitinous fibre, of which the meshes are
more or less oblong, passing into prominent longitudinal lines on the branchlets
where they terminate on the backs of the semitubular plates which respectively form
the floors of the hydrophores, to which they thus give support. Size of specimen,
which is merely a branch 43 inches long by 2 broad.
Locality.—Port Natal.
Mr. Carter adds that ‘‘the spines on the surface distinguish this from the
foregoing species, add to which its longer and more pointed branches, longitudinally
ridged clathrate fibre and rich red-purple colour.”
Genus. Chitina. (Carter.)
Colony erect, bushy, fragili flexible, fawn coloured. Trunk long, hard,
irregularly round, composed of many stems united clathrately and obliquely into a
cord-like bundle, which divides and subdivides irregularly into branches which again
unite with each other in substance (anastémose) when in contact and finally form a
straggling bushy head. Hydrophores long clathrate tubular, terminating the ends
of the branchlets, or prolonged from some of the proliferous tubercles which beset
the surface of the trunk and larger stems. Minute structure : Composed of clathrate
chitinous fibre throughout, whose network is subrectangular and massive in the stems,
where there is no difference between the centre and circumference, with the exception
that the fibre is stouter in the former or oldest part; hydrophores composed of several
longitudinal fibres or ridges lattice-worked together transversely into a tubular form,
somewhat contracted at the extremity, in the centre of which is an aperture of the
meshwork a little larger than the rest. Height of specimens about 14 inches, trunk
about 1 inch in diameter; hydrophores averaging 1-3rd of an inch long by
1-60th of an inch in its broadest part and an aperture 1-90th of an inch in
diameter.
(5) Chitina ericopsis. (Carter.)
The description of the species is the same as that of the genus.
Locality.—New Zealand.
22 ON THE STRUCTURE OF CERATELLA FUSCA (GRAY).
DeEscrRIPTION OF PLATES.
PLATE II.
Fig. 1—Ceratella fusca. Life size. The specimen of which this is a drawing
was washed up on Flinders Island, Bass Straits. The main stem of the colony
springs from a root-like structure made up of intertwined branches. On the left side
arises from the roots a very small independent stem.
Fig. 2.—Transverse section through a portion of the wall of an expanded
gastrozooid. Externally is the layer of ectoderm consisting of cubical cells. The
mesogleea is an extremely thin layer and the endoderm consists of large vacuolate
cells with granular protoplasm aggregated at their inner ends which face into the
gastral cavity. The nuclei are conspicuous and placed at the same end. Camera.
Zeiss H, oc. 2.
Ect. ectoderm. End. endoderm. WM. mesoglea.
Fig. 8.—A small portion of the external surface of a branch in the region in
which gonophores are numerous to show the external ectoderm containing thread
cells and a portion of one of the ccenosare tubes. In the latter the endoderm forms
a definite layer of darkly-stained, small cubical cells. Surrounding this is the
ectoderm in contact with the external layer and containing developing thread cells.
E. external layer of ectoderm. Ect. ectoderm of ccenosare tube. End.
endoderm. A. developing thread cells. Sk. skeleton. Camera. Zeiss H, oc. 2.
Fig. 8a.—Developing thread cells found in the ectoderm within the branches.
a. an ectoderm cell in which the nucleus is slightly larger than usual and appears
homogeneous. b. the nucleus has increased in size, stains very darkly, lies at one end
and is surrounded by a clear space. c. the nucleus begins to show a darker spot
within it. d. two darkly-staining thread-like structures are present. e. the nucleus (?)
does not stain so deeply, and is not surrounded by a clear space but has a small
amount of protoplasm clinging to it; within it at the somewhat pointed extremity can
be seen a light line indicating the larger terminal part of thethread. f. the wall
of the thread-cell and the thread itself are clear. 9. The fully developed thread-cell.
PLATE III.
Fig. 4.—Portion of a small branch with the zooids expanded. The skeleton is
covered by the soft parts but the dark lines indicate the external parts of the chitinous
network which show through the soft structures.
ON THE STRUCTURE OF CERATELLA FUSCA (GRAY). 238
Fig. 5.—Portion of the skeleton of a somewhat larger branch showing the
chitinous network of which it is composed and the little bracket-like hydrophores.
(Ayyy x12:
Fig. 6.—Highly magnified small portion of a branch seen partly by direct and
partly by transmitted light. Two gastrozooids are shown each of which is placed upon
a hydrophore. The latter shows prominent ribs which project beyond the margin
and are connected by a thin fenestrated web of chitin. Two male gonophores arise
from the ccenosare and they and the gastrozooids are quite naked. The whole branch
is covered by a thin external layer of ectoderm. x 20.
G. gastrozooids. Gon. gonophore. £. external layer of ectoderm. Hy. hydro-
phore. Hy.' web connecting the ribs of the latter.
Fig. 7.—Transverse section across the body of a gastrozooid. The tentacles
are solid with swollen ectoderm ends filled with thread cells. The zooid has been
feeding and the endoderm cells are full of little food particles. /.—Remnant of
small crustacean on which the zooid is feeding. It lies close against the endoderm
on one side and the food particles are passing into the interior of the cells. Outline
drawn with Camera, Zeiss H, oc. 2.
Fig. 8.—Transverse section across a gonophore. Mn. endoderm of the
manubrium. Sf. sperm cells in the ectoderm of the latter. Ect. ectoderm of the
medusa. . radial canals. Ect’. ectoderm of the sub-umbrella surface. Outline
drawn with Camera, Zeiss E, oc. 2.
PLATE IIIA.
Fig. 9.—Semi-diagrammatic drawing of a longitudinal section through a
small branch. The skeleton is coloured brown the soft parts grey. E. external
layer of ectoderm. Gon. gonophore. G. gastrozooid. Hy. hydrophore. Sf.
general network of skeleton cut across in various directions. Outline drawn with
Camera, Zeiss A*, oc. 2.
Fig. 10.—Semi-diagrammatic drawing of a transverse section across a good-sized
branch. Letters as in figure 9. Ect. ectoderm of endosare tubes. End. endoderm.
Outline drawn with Camera, Zeiss A*, oc. 2.
Fig. 11—More highly magnified portion of a branch cut in transverse section.
The ectoderm of the gastrozooid is directly continuous with the external layer of the
branch and the base of the zooid is continuous with two of the ccenosare tubes.
At C. is represented the characteristic feature of the latter crossing over five
connecting strands of the chitinous network. C. points at which the soft parts
24 ON THE STRUCTURE OF CERATELLA FUSCA (GRAY).
cross thin connecting strands of the skeleton. &. external layer of ectoderm.
Iict. general ectoderm. G. gastrozooid. Hy. hydrophore. Sk. general skeleton.
Outline drawn with Camera, Zeiss A*, oc. 2.
Kig. 12.—Portion of a longitudinal section of a branch from which arise two
gonophores. The latter are cut in longitudinal section. &. external layer
continuous with the ectoderm of the gonophore. Ld. endoderm of the manuubrium.
Ect. ectoderm. Ect'. ectoderm of sub-umbrella layer of the medusa. G.
eastrozooid. M. point at which the ectoderm dips in corresponding to the mouth
of the medusa. RR. radial canals. Outlme drawn with Camera, Zeiss C, oc. 2.
Fig. 18.—Longitudinal section of a branch from which another small one arises.
In this specimen the skeleton has more than usual a more or less definite
arrangement into ribs which run parallel to the length of the branch and are
connected by transverse bands. Outline drawn with Camera, Zeiss A*, oc. 2.
Fig. 18a.—A small portion of the small branch in figure 13 more highly
magnified to show the single caenosare tube passing along the centre. The ectoderm
is irregular. Letters as before. Drawn under Zeiss F, oc. 2, *
Fig. 14.—Portion of a terminal branch devoid of zooids. Up the centre runs
a single tube with an internal unicellular layer of endoderm and an irregular ectoderm.
‘he endoderm gives off hollow processes which at certain parts (x) come into direct
contact with the external ectoderm (Z). Further growth of these will probably give
rise to the gastrozooids the external layer of the colony thus forming their ectoderm.
Drawn under Zeiss H, oc. 2.
>
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ARTICLE III.—Apprrionan OpsErvAtTIONS ON THE VicTorRIAN Lanp PLANARIANS,
By ArtHuR Denpy, D.Sc. (With Plate IV.)
(Read June 12th, 1891.)
Since my last paper ‘‘On the Victorian Land Planarians”’ was read before this
Society I have been able to collect a considerable amount of further information
concerning this interesting group of worms, and the present observations are intended
to bring our knowledge of the group up to date by describing species or varieties
hitherto unknown and especially by giving full details as to the variation in colour
and markings and as to the distribution within the colony of those species which
have already been described.
In my previous memoir I recorded fifteen Victorian species ; this number has
now been brought up to twenty-two, the additional seven species being Geoplana
dendyi, Spencer, G. frosti, Spencer, G. howitti, Dendy, G. ventropunctata, Dendy,
all of which are new to science since I last wrote; with G. swlphurea, Fletcher and
Hamilton, and G. sanguinea, Moseley, hitherto recorded only from New South
Wales, and Bipalium kewense, Moseley, an introduced species recorded by Fletcher*
from near Hltham. The genus Rhynchodemus has not again been met with.
G. howitti and G. ventropunctata are now fully described and figured for the first
time, and I have also taken the opportunity of giving figures of several other species
or varieties which I had not been able to figure previously.
The question of the distinctness of the different ‘species’? remains where it
was, but I hope that the observations concerning the variation in pattern and colour
recorded in the present contribution will be of assistance in ultimately settling it.
These observations are given systematically under the headings of the different
species, and I have thrown out hints as tothe possible relationship of the different
forms, which may or may not be followed by future workers.
I have also been in communication with Professor L. von Graff, of Gratz, who is
engaged in the preparation of an elaborate monograph of the whole group and wrote
to me for specimens of our Australian Planarians. Iam glad to have beeu able to send
Professor von Graff a large number of our species, which he intends to examine
* Linnean Society of New South Wales, February 25th, 1891. (Zoologischer Anzeiger, April 20th, 1891, p. 139.)
26 ADDITIONAL OBSERVATIONS ON
anatomically, and when the results of this anatomical mvestigation are combined with
the observations of Australian zoologists upon the living animals we may hope to
arrive at a fair idea of the relationship of the various forms.
In the systematic part of this paper the species are arranged in what appears to
me to be the most natural order, commencing with the forms without stripes.
Fifteen of the species thus appear to fall into five groups, within each of which the
species are all closely related. The remaining seven species do not to my mind fall
into any of these groups and are also very distinct from each other. The following
is the complete list up to date, the related species being bracketed together in
groups :—
1. Geoplana alba, Dendy.
2. 3 sanguinea, Moseley.
3. “i spenceri, Dendy.
4, FA cerulea, Moseley.
Lis, $5 dendy1, Spencer.
6. 3 walhalle, Dendy.
he , frosti, Spencer.
S i sugdent, Dendy.
9. mediolineata, Dendy.
| 10. rr quinquelineata, Fletcher and Hamilton.
ue me m'mahont, Dendy.
12. , sulphurea, Fletcher and Hamilton.
(rey - hoggit, Dendy.
(ela: - quadrangulata, Dendy.
ena mi, ventropunctata, Dendy.
16. “ munda, Fletcher and Hamilton.
ie ae ade, Dendy.
18. 4 fietcheri, Dendy.
19. Ms lucast, Dendy.
20. 53 howitti, Dendy.
21. Fhynchodemus victoria, Dendy.
22. Bipalium kewense, Moseley.
With regard to the distribution of the species within the colony, I have been
able to collect a considerable amount of information; and, for the benefit of
collectors, to whom it may be of service to know what localities have been explored
and what species found in each, I give the following table of distribution. I at
first thought of giving also a list of the months of the year in which each species
has been found, but this appears to me to be superfluous. Land Planarians may,
I believe, be found at all times of the year by diligent searching, but they seem to
THE VICTORIAN LAND PLANARIANS. 27
be much more abundant in spring (or early summer) and autumn than in the great
heat and drought of the summer or the cold and excessive moisture of the winter
months.
Toorax.—Geoplana cerulea.
HEIDELBERG.
G. quinquelineata; G. munda.
Eiruam.—Bipalium kewense.
Croypon.—G. alba; G. mediolineata; G. sulphurea; G. munda; G. ade.
Fern Tree Guiiy.—G. alba; G. mediolineata; G. sulphurea; G. munda; G.
ade; G. m‘mahoni; G. ventropuncttata.
Warracui.—G. alba; G. sugdeni.
Between LinypaLe and Sevitte.—G. m‘mahoni.
Warsurton.—G. sfencerz; G. mediolineata; G. ada.
M‘Manon’s Creek (Upper Yarra).—G. alba; G. m‘mahoni ; G. spencert.
Between Marysvitte and the Source or tHe Yarra.—G. alba; G. spencers;
G. dendyi; G. walhalle; G. frosti; G. mediolineata; G. m‘mahont ;
G. sulphurea; G. munda.
WaLHALLA.—G. spenceri ; G. mediolineata; G. walhallea ; G. sulphurea.
Upper Wetuineton River.—G. alba; G. sulphurea; G. quadrangulata, vay.
wellingtoni ; G. lucasi; G. howttti.
Croasincotone.—G. caerulea; G. dendyi; G. alba; G. lucasi; G. sulphurea;
Rhynchodemus victoria.
CASTLEMAINE.—G. sanguinea.
SaNDHURST.—G. quinquelineata.
Batuarat.—G. alba; G. quinquelineata ; G. munda.
Creswick.—G. sugdeni; G. mediolineata; G. quinquelineata; G. hoggi; G.
quadrangulata; G. munda.
Macepon.—G. alba; G. sugdent; G. mediolineata; G. hoggiu; G. ade; G.
quadrangulata; G. fletchert.
Orway Forest.—G. sulphurea; G. quinquelineata; G. munda.
From this it will be seen that, so far as Planarians are concerned, a very large
proportion of the colony is unexplored, and we may expect considerable additions to
our Planarian fauna in the near future.
Bipalium kewense is undoubtedly an introduced species, whose natural habitat is
unknown. It was obtained from the Kew Botanical Gardens near London, and
28 ADDITIONAL OBSERVATIONS ON
described by Professor Moseley. Since then it has been found in various parts of
England and also in Sydney,* and still more recently, as I have already noticed, Mr.
Fletcher has recorded its occurrence at Eltham, and likewise in Samoa. Good figures
of this interesting Planarian are given by Bell in his ‘‘ Note on Bipalium kewense,
and the Generic Characters of Land-Planarians.”’t
Concerning Rhynchodemus and Bipalium I have nothing further to add, and we
may pass on to the systematic observations on the 20 species of Geoplana. As the
present memoir is merely a continuation of my earlier paper ‘On the Victorian
Land Planarians,” published in the Transactions of this Society for 1890, I have not
thought it necessary to give constant references to that paper, to which I would
refer the reader for further information and references.
1. Geoplana alba, Dendy.
This is one of the most widely distributed and abundant of our Victorian
Land Planarians. Since the publication of my description and figures Professor
Spencer has recorded the species}—on the occasion of the Field Naturalists’
Expedition to the Yarra Falls—and has figured two specimens which are of a rather
darker colour than usual and thus approach G. sanguinea. As already noted in my
original description the species varies a good deal as to the depth of colouration, from
almost pure white to peach-coloured or yellow flesh. I may add that the eyes are
very small and difficult to make out. In addition to the localities already mentioned
I have now obtained the species from Croydon, Fern-Tree Gully, Ballarat (abundant
under stones close to the edge of the lake, Mr. Avery), and the Upper Wellington
River.
2. Geoplana sanguinea, Moseley.§
This species has not hitherto been recorded from Victoria, but I have now
received seven specimens collected by Mr. T. S. Hall, M.A., at Castlemaine, under
fallen bark and stones. The specimens were received by me in alcohol, with the
following note as to the colours of the living worm :—* Rich Indian red above,
paler below.’ The specimens (in spirit) are about 25 mm. long, and 4 mm.
broad, much flattened on the ventral surface but pretty strongly arched on the dorsal.
The anterior end of the body tapers off gradually, the posterior end terminates much
* Pletcher. ‘Remarks on an introduced Species of Land-Planarian, &c.,” Proc. Linn. Soc., N.S.W., May 25, 1887.
+ Proc. Zool. Soc. Lond., 1886, p. 166, Plate XVIII.
t Proceedings of the Royal Society of Victoria, 1891, p. 88.
§ Quarterly Journal of Microscopical Science, Vol. XVII., N.S., p. 285.
THE VICTORIAN LAND PLANARIANS. 29
more abruptly. The peripharyngeal aperture is situate at about the junction of the
middle and posterior thirds of the body and the genital aperture nearer to it than to
the posterior end.
The eyes are, as in G. alba, very small and difficult to make out.
Geoplana (Cenoplana) sanguinea was very imperfectly described by Moseley as
follows :—‘‘ Closely resembles C. Coerulea, with the exception that it is coloured of
a uniform light red, which is lighter upon the under surface of the body. Actual
length when living 7 cm.; breadth 4 mm., Parramatta, near Sydney. Amongst
earth at the roots of a Eucalyptus stump.” (Loc. cit.)
It seems probable that Fletcher and Hamilton’s G. rubicunda* is identical with
G. sanguinea, a conclusion at which I understand Mr. Fletcher has himself now
arrived. My G. alba is also evidently closely related to G. sanguinea and may have
to be considered merely as a variety of that species.
G. sanguinea (as represented by the Castlemaine specimens), G. rubicunda and
G. alba all agree in the general shape of the body, in the position of the apertures
and in the curious indistinctness of the eyes.
The Castlemaine specimens of G. sanguinea are all markedly smaller than the
average size of G. alba, but Moseley’s examples seem to have been of about the same
size as the latter.
3. Geoplana spenceri, Dendy.
This species has again been met with by Professor Spencer} on the occasion of
the Field Naturalists’ expedition to the Yarra Falls, in a locality intermediate between
the Upper Yarra district and Walhalla, whence it had been previously recorded.
Though extremely abundant in these parts it does not seem to occur in other localities,
so that its distribution appears to be curiously limited in extent.
4. Geoplana cerulea, Moseley.
@isy.,, Wie. %)
In my previous memoir on the Victorian Land Planarians I recorded this species
from Croajingolong, where it was collected by Professor Spencer. Since then
I have only met with a single specimen and that in a somewhat remarkable
* Proceedings of the Linnean Society of New South Wales.”’ Series II., Vol. IL, p. 370.
+ Loc. cit.
30 ADDITIONAL OBSERVATIONS ON
situation, viz., in the earth around the roots of a chrysanthemum which
was given to me by Mr. G. W.: Officer and which was taken out of the ground in
Toorak, near Melbourne, in June, 1890. This specimen when alive was of a very
dark Prussian blue colour on the dorsal surface and paler blue on the ventral.
There was a narrow median dorsal stripe of yellow, and an indistinct median
ventral very pale blue stripe traceable back nearly to the genital aperture. There
was no different colour at the anterior extremity. In spirit the specimen measures
about 35 by 2°5 mm.; the peripharyngeal aperture is very slightly behind the
middle of the body and the genital aperture at rather more than one-third of the
distance from the peripharyngeal opening to the hinder end of the body. The eyes
are easily visible with a lens. ‘This specimen, although now preserved in spirit for a
year, has preserved its blue colour remarkably well. It is impossible, of course,
to determine how it was imtroduced into Mr. Officer's garden at Toorak, and
it is interesting to note in this connection a remark made by Messrs. Fletcher
and Hamilton* concerning the same species to the effect that the specimens usually
found by them have an orange-red tip (as appears to have also been the case in the
Croajingolong specimens) but that ‘‘ on three different occasions we have found on
the pavement in Hyde Park alongside the enclosure at Captain Cook’s statue a
number of blue Planarians (about fourteen altogether), which are without the red tip,
and in which the median stripe varies from a dirty white to a distinct yellow,
changing to white in spirit. . . . . Theenclosure referred to has probably been
stocked with these Planarians from the Botanic Gardens, but we do not know from
what locality. The differences in living specimens in the two cases seem to be
constant, and are sufficiently marked to make one a variety of the other, if not to
separate them as distinct species.’ ‘They consider that the red-tipped variety is
probably the typical form, although Moseley does not mention the red colour of
the anterior extremity. It is curious that the ‘ introduced” Toorak specimen, if
I may use the term, should exhibit exactly the same peculiarity as the ‘‘ introduced”’
Sydney specimens, and this is an argument for supposing them to belong to a
distinctly marked variety whose natural habitat is not known, but which is
characterised by the absence of any red colour at the anterior extremity. }
5. Geoplana dendyi, Spencer. |
This species is evidently very closely related to Geoplana cerulea, being
distinguished from it perhaps only by the presence of the median dorsal line of blue
* Loe. cit.
+ On the evening on which this paper was read another specimen of this variety was found on the floor in the house
at Toorak from the garden of which the first specimen was obtained almost exactly twelve months ago. Mr. Officer kindly
brought this second specimen to me alive, so that owing to this remarkable coincidence I am able to give a figure of
the worm in its living condition. The second specimen agrees so closely with the first that no special description is
necessary.
t) oc: cit.
THE VICTORIAN LAND PLANARIANS. 31
in the middle of the yellow stripe. Some of the specimens brought by Professor
Spencer from Croajingolong (in spirit), which otherwise resemble G. c@rulea, have
the dorsal light band thus divided into two by a dark line, and I find that I had
accordingly placed them in separate bottles away from those which | definitely
identified with cwrulea. Probably, therefore, these Croajingolong specimens are also
referable to G. dendyz, so that we have the two closely allied species, cwrulea and
dendy1, inhabiting the Croajingolong district. The Croajingolong specimens appear
to be more flattened than those described by Professor Spencer.
6. Geoplana walhallea, Dendy.
Only a single specimen of this species has been obtained since it was first
described, and that by Professor Spencer, who found it in the country lying between
Marysville and the source of the Yarra along the Wood’s Point Road, and has
described and figured it in the Proceedings of the Royal Society of Victoria.* The
genital opening, as figured by Professor Spencer, is much nearer to the peripharyngeal
opening than in the type specimens.
7. Geoplana frosti, Spencer.t
This species, as pointed out in the original description, is obviously very closely
related to G. walhalle, from which it is distinguished only by the presence of two
light yellow stripes one on each side of the mid-dorsal line and by the absence of the
brown speckles along the mid-ventral line. I do not feel disposed to lay any stress
on the latter character. Professor Spencer has figured an interesting connective
form in which the dorsal yellow stripes are absent throughout the greater part of the
length of the body.
T have identified} a small specimen which I found near Mount Wellington,
Gippsland, with this species. In life the ground colour of the dorsal surface was
very dark indigo blue, so dark that the exact colour was difficult to determine, with
two narrow stripes of whitish separated by a very fine, dark, mid-dorsal line. The
ventral surface was pale brown flecked with darker brown. Even in the spirit-
preserved specimen a light mid-ventral line, devoid of specks, is distinctly visible,
and the opening into the peripharyngeal chamber is situate in about the centre of
the body and the genital aperture (?) only a short way behind it. The specimen
(in spirit) measures 12mm. in length by a little under 3mm. in greatest breadth.
* Loc. cit.
{ Loc. cit.
t Victorian Naturalist, June-July, 1891, p. 44.
32 ADDITIONAL OBSERVATIONS ON
8. Geoplana sugdent, Dendy.
T have received specimens of this species from Creswick, collected by Mr. Fiddian,
and have also obtained more specimens from Macedon, whence it was originally
recorded. At Macedon I have again observed it crawling about in broad daylight.
Professor Spencer informs me that he has found this species at Warragul.
9. Geoplana mediolineata, Dendy.
I identify with this species specimens obtained from Fern Tree Gully, Croydon
and Creswick, and Professor Spencer has figured* a variety of the species from the
country lying between Marysville and the source of the Yarra along the Wood’s Point
Road. As Professor Spencer has pointed out, specimens in which the lateral stripes
are well developed make an approach to Geoplana quinquelineata ; on the other
hand, the species seems also to merge into G. sugdeni. Hence although typical
specimens of G. sugdeni and G. quinquelineata appear very different indeed it seems
not impossible that they may have to be united with G. mediolineata as varieties of
one species. G. mediolineata occupies an intermediate position between the other
two, not only as regards the stripes but also in the shape of the body, being not
usually so flattened as G. guinquelineata and less cylindrical than G. sugdenti.
In G. mediolineata the median stripe is always much darker in colour than the
others (if they be present). In this it agrees with G. quinquelineata and differs from
G. m‘mahoni.
10. Geoplana quinquelineata, Fletcher and Hamilton.
This species seems to be characteristic of the Sandhurst district and in addition
to the localities mentioned in my previous memoir I have now to record it from
Ballarat, Creswick and Heidelberg. In some of the Creswick and Ballarat specimens
the lateral stripes show a strong tendency to disappear and it is difficult to say
whether the specimens should be called quinquelineata or mediolineata. The genital
aperture is very difficult to observe but appears to me to be rather nearer to the
peripharyngeal aperture than to the hinder end of the body, while the peripharyngeal
aperture is more easily visible and situate about the middle of the ventral surface.
Hence G. sugdem, G. mediolineata and G. quinquelineata agree pretty closely in the
position of the apertures.
= Loc; ictt:
THE VICTORIAN LAND PLANARIANS. 33
11. Geoplana m‘mahoni, Dendy.
(BIVLV., Haga)
Since the publication of my previous memoir I have met with several interesting
specimens of this species at Fern Tree Gully. The ground colour in the living worm
was bright canary yellow on the dorsal surface, paler on the ventral. Some
specimens had (like the original type) only the two stripes on the dorsal surface, both
broad and black, or nearly so. Others had, in addition, a much narrower median
dorsal stripe of the same colour, and yet others were intermediate between these two,
the median band being very narrow and discontinuous. The paired stripes are,
in this species, the characteristic and constant ones, and not, as in G. mediolineata
the least persistent; they are always much more prominent than the median
stripe. The position of the genital and peripharyngeal orifices in the Fern Tree
Gully specimens agree in one specimen with that described for the type, though
in another the peripharyngeal aperture is almost central. Ido not set much value on
slight differences in this respect, as the position of the apertures must depend to a
certain extent upon the state of contraction of the specimen.
Professor Spencer* records the species from the country lying between
Marysville and the source of the Yarra along the Wood’s Point Road.
Thave also a slight variety of this species from between Lilydale and Seville.
The colour when alive was bright yellow all over except for the brown anterior tip
and two strong dorsolateral stripes (one on each side) of brown starting from the
anterior tip and dying out about one-third of the way down the body, and also a very
short, thin, median dorsal stripe of brown starting from the anterior tip
but soon dying out. When alive the specimen measured about 60 mm.
in length. In spirit it is rather broad and flat and the peripharyngeal aperture is
situate shghtly in front of the middle of the body and the genital aperture not very
far behind it, much nearer to it than to the posterior end of the body; obviously,
however, the anterior half of the body is much more contracted than the posterior,
thus bringing the apertures forward.
The occurrence of the Fern Tree Gully variety with the median stripe is
interesting as showing how an even-striped form may be derived from an odd-striped
form by disappearance of the median stripe, and as pointing out a possible relationship
between G. mediolineata (and its close allies), and G. m‘mahoni.
* Loc. cit.
34 ADDITIONAL OBSERVATIONS ON
12. Geoplana sulphurea, Fletcher and Hamilton.
(Geoplana sulphureus, Fletcher and Hamilton, Joc. cit.).
This species was found by Professor Spencer in the country between Marysville
and the source of the Yarra, alongthe Wood’s Point Road, and is identified and
figured by him in the Proceedings of this Society.* I have in my collection a
number of specimens from other parts of Victoria, viz., the Otway Forest (two very
large specimens), Croydon, Walhalla, Croajingolong, Fern Tree Gully, and the
Upper Wellington (a number of very small specimens). Some of these specimens
have been in my possession for a long time but the original description is so short
that I had not ventured upon an identification. Neither Messrs. Fletcher and
Hamilton nor Professor Spencer mention the position of the external openings. In
spirit specimens I find the peripharyngeal aperture situate in about the centre of the
body and the genital aperture at about one-third of the distance between the
peripharyngeal aperture and the posterior end.
In G. sulphurea the outer stripes are broader than the inner, in G. hoggii, which
é
comes very near it, the reverse is the case.
The ground colour in G. su/phurea is yellow. The inner stripes are dark brown,
or sometimes greenish grey, the outer ones are darker, usually black or nearly so.
In one of my Wellington specimens the two inner stripes are so close together that
in the spirit-preserved specimen they appear as one, so that the specimen appears to
have three stripes of equal width. In an interesting specimen from Fern Tree
Gully all four stripes are extremely thin and discontinuous and almost entirely absent
from the posterior part of the body; the outer stripes, however, are still distinctly
stronger than the inner ones.
13. Geoplana hoggit, Dendy.
Thave little to add to my original description of this worm. I have again
collected it at Macedon, where it is very abundant, and have also received typical
specimens from Creswick, collected by Mr. Fiddian. The form certainly comes
very near to G. sulphurea, and had I realised what G. sulphurea was really like at
the time when I wrote my description I should possibly have considered it as a
variety of the latter. G. hoggii, however, differs from G. sulphurea, as pointed out
above, in having the inner stripes broader than the outer ones, the green colour being
at the same time more pronounced; hence it may perhaps stand as a fairly
well-marked variety, though probably not a distinct species.
* Loc. cit.
THE VICTORIAN LAND PLANARIANS. 35
14. Geoplana quadrangulata, Dendy.
I have again collected this very beautiful and distinct little Planarian at
Macedon, where it is by no means uncommon, and I have also received a specimen
from Creswick, collected by Mr. Fiddian. These specimens all exhibit in a more or
less marked degree the pale spots along the junction of the dorsal and lateral
surfaces ; these spots are merely the foreshadowings of the characteristic mottled
appearance of the Mount Wellington variety.
14a. Geoplana quadrangulata, var. wellingtont, nov.
This pretty little variety was obtained in abundance under logs near the foot of
Mount Wellington and its occurrence has already been recorded by me in the “ Victorian
Naturalist.” * The worm is arather difficult one to collect as it crawls into the crevices
of the rotten wood in a very provoking way and readily breaks to pieces.
When alive the ground colour of the dorsal surface was pale brown, with a
median dark brown line and on-each side of it numerous thickly scattered speckles
of dark brown.
The lateral surfaces were very pale brown thickly flecked with dark brown, and
the ventral surface was white, devoid of speckles. All the specimens obtained were
a good deal smaller than the typical form of the species from Macedon usually is.
More than a dozen specimens were collected.
15. Geoplana ventropunctata, n. sp.
(Pl. IV., Figs. 2, 2a.)
I obtained this very beautiful little species in abundance under rotten logs at
Fern Tree Gully, on the occasion of the Field Naturalists Club’s excursion to that
locality in March last.
When at rest the body is very broad and flat (Pl. IV., Fig. 2a), but when
crawling, which is done very actively, the body becomes greatly elongated and very
narrow, measuring about 20 mm. in length anda little under 2 mm. in greatest
breadth.
The eyes are arranged as usual on the horse-shoe shaped anterior extremity and
sides of the body.
* Loe. cit.
36 ADDITIONAL OBSERVATIONS ON
The opening of the peripharyngeal chamber (in spirit) is somewhat behind the
middle of the ventral surface and the genital aperture about one-third of the way
between it and the posterior end of the body. The protruded pharynx (in spirit) is
cylindrical.
The ground colour of the dorsal surface is really nearly white, but almost entirely
obscured by the abundant markings, which are arranged as follows:—(1) A rather
narrow, nearly black median stripe with irregular margins. (2) On each side of the
black stripe and extending to the lateral margin of the body are a great number of
small, closely set, irregular longitudinal streaks of various shades of brown, in some
specimens a good deal lighter than in others. Each side of the body, between the
dorsal and ventral surfaces, is occupied by an irregular stripe of very dark brown,
made up of a series of splotches of pigment more or less run together. The anterior
extremity is dark brown.
The ventral surface is white, very beautifully spotted all over with small, distinct
and isolated specks of dark brown.
When the animal is alive the lateral stripe is visible from the ventral surface
but not from the dorsal ; in spirit it is just visible from the ventral surface.
In general appearance and size this species closely resembles Geoplana
quadrangulata var. wellingtoni, but we may note the following points of distinction :—
(1) The body is not so distinctly quadrangular in section and the sides do not slope
inwards nearly so much, so that the ventral surface is much wider. (2) There is a
distinct, though irregular lateral stripe of a very dark colour, instead of the sides of
the body being lighter in appearance than the dorsal surface as in G. quadrangulata
and its variety. (8) The ventral surface is, in all specimens which I have seen (a
considerable number), very distinctly spotted with brown; while in all specimens of
G. quadrangulata which I have seen it has no spots at all.
It a curious fact that while this species was found abundantly at Fern Tree
Gully in March, in company with large numbers of Geonemertes australiensis,
yet we could not find a single specimen of either of these worms when we visited the
spot a few weeks later in May.
16. Geoplana munda, Fletcher and Hamilton.
This species has proved to be one of the commonest and most easily recognisable
of our land Planarians. Since writing my last memoir on the subject I have had
specimens from Croydon, Fern Tree Gully, Heidelberg, Creswick and Ballarat. At
THE VICTORIAN LAND PLANARIANS. 37
Croydon, on December 17th, 1890, I found more than fifty specimens of this species
under a single log, all agreeing closely in colour and markings.
I was at first much puzzled at not being able to find the genital aperture in this
species. I could not find it in the Otway Forest specimens recorded in my previous
memoir, nor could I find it in the specimens collected at Croydon in December, 1890,
though I carefully examined a considerable number. When, however, I came to examine
specimens collected at Creswick in March, 1891, I found it at once in ten out of eleven
specimens, the eleventh being a very small one. The Croydon specimens were fully
of the average size and the obvious conclusion seems to be that the genital aperture
is visible only at certain times of the year (autumn). With this we may associate the
fact, recorded in my last memoir, that Geoplana mediolineata copulates in the beginning
of April. Possibly all our land Planarians breed in the autumn.
The peripharyngeal aperture in G. munda is nearly central and the genital
aperture only a short way behind it, as originally described by Fletcher and Hamilton.
Professor Spencer* has also recorded the species from the country lying between
Marysville and the source of the Yarra along the Wood’s Point Road, and has
given an excellent coloured figure of the living worm.
I have now to mention a slight variety of the species which I found at
Heidelberg and which was again obtained by Mr. Fiddian at Creswick. The
Heidelberg specimens, three in number, are thus described in my notes on the living
animals :—‘“ In size and shape closely resembles G. munda, the ground colour is also
yery similar, on the dorsal surface a slightly mottled olive brown; ventral surface
pale yellow. In the mid-dorsal line a fine, very pale brown band, edged on either
side by a fine dark brown line which forms a margin to the ground colour. No other
stripes.” The Creswick specimens presented exactly the same peculiarity, viz., the
absence of the dorso-lateral dark stripes; they were collected in May, and show the
genital aperture distinctly.
17. Geoplana ade, Dendy.
(Pl. IV., Figs. 3, 3a.)
I have again collected this very handsome Planarian at Macedon and have also
found it at Croydon and Fern Tree Guliy. ‘The genital aperture, which I had not
been able to distinguish when I first described the species, is situate only a short way
behind the peripharyngeal and a long way from the posterior end of the body ; it
appears to vary in distinctness with the time of the year, as in G. munda.
¥ Loc. cuts
38 ADDITIONAL OBSERVATIONS ON
My original description and figure did scanty justice to the beauty of this
species, partly because the specimens were not so fine as those which I have since
obtained.
The two specimens from Croydon were about 65 mm. in length and 4 mm. in
width when crawling. The ground colour of the dorsal surface was light, mottled
brown, and the paired stripes were very broad, of a dark greeny brown colour, with
a fine, well-defined, very dark outline on each side. One of the specimens shows a
fine, ight-brown mottling on the ventral surface, except in the mid-ventral line, the
other only shows a trace of this mottling at the anterior end.
At Fern Tree Gully, on March 14th, 1891, a number of fine specimens were secured
which again showed some interesting variations in colour. The ground colour of the
dorsal surface was very pale, yellow brown, generally, at any rate, more or less
mottled, but only very slightly. The ventral surface was very pale yellow brown,
often shehtly mottled and sometimes with two broad bands of distinct, slightly
mottled brown one on each side of a narrower median band of almost pure
white. In some specimens the three stripes on the dorsal surface were coloured
as in the specimens originally described and figured. The most striking variety,
however, is that represented in Figs. 3, 3a., Pl. IV. In this specimen the ground
colour of the dorsal and ventral surfaces was very pale brown very slightly
mottled, with slght imdications of a lighter median stripe on the ventral
surface. The two broad stripes on the dorsal surface, mstead of being simply
brown, or purple brown as in some specimens, were of a dark brown colour with very
minute blue specks thickly scattered all over. Viewed as a whole these stripes
appeared of different tints according to the light thrown on them, but the general
impression was dark olive green.
When at rest G. ade is very broad and flat,and the green variety closely
approaches G. frosti in appearance. The broad, dark bands are, however, edged on
the outside with ground colour, and do not extend to the lateral margin asin G. frosti.
In other words, what appears as ground colour in G. frost: appears as a definite stripe
in G. ade.
18. Geoplana fletchert, Dendy.
(ELLY jelio.iO5)
I have again found this species at Macedon but have not seen it from any other
locality. I take this opportunity of emphasising the fact that this Planarian, which
at first sight might easily be mistaken for one of the common yellow forms, is really
very distinct indeed. The much flattened ventral surface, sometimes becoming
THE VICTORIAN LAND PLANARIANS. 39
strongly concave in spirit, and the markedly posterior position of the apertures, the
genital being close to the hinder extremity of the body, readily distinguish the
species.
In one of my specimens the copulatory organs are partially protruded, and
exhibit the very remarkable peculiarity that while the female copulatory organ is
single there are two distinct male organs. The appearance and arrangement of these
parts 1s shown in Figure 6. Whether or not this condition is constant I cannot say,
but I am inclined to think that the species possesses anatomical peculiarities which
would well repay investigation, and which may even necessitate the erection of a new
genus for its reception. The eyes are arranged as is usual in Geoplana.
19. Geoplana howitti, Dendy.*
(PRIN. Big. 5.)
Body (in spirit) much flattened on the ventral surface, strongly convex on the
dorsal, much broader behind, where it terminates rather abruptly and bluntly, than
in front, where it tapers gradually. Peripharyngeal aperture (in spirit) well behind
the middle of the ventral surface, only 8mm. from the hinder extremity of the body.
Genital aperture about 4mm. behind the peripharyngeal. Length of body (in spirit)
about 25mm., greatest breadth 4mm. ‘The eyes are arranged as usual in the genus
but are difficult to make out owing to the dark pigment around them, so that I had
to cut a slice off the side of the head end before I could find them.
When alive the ground colour of the dorsal surface was yellowish white ; in the
mid-dorsal line a fairly broad band of ground colour, on each side of this a stripe of
about equal width of dark purplish brown, outside this a rather broader band of
ground colour thickly flecked with dark purplish brown and edged on the outside by
a fine line of dark purplish brown, outside this fine line a very narrow edge of ground
colour.
It is noteworthy, as showing a tendency towards variation in pattern, that in the
mid-dorsal band of ground colour, just in one place near the posterior end, there are
a few small flecks of dark purplish brown, as shown in the figure. The horse-shoe-
shaped anterior extremity is, during life, dark purplish brown, and all the dark bands
unite at each end of the body.
The ventral surface in life is pale yellowish white or grey, with no markings.
I found a single specimen of this Planarian beneath a log near the bank of the
Upper Wellington River, at the foot of Mount Wellington, Gippsland, on the occasion
* A brief preliminary diagnosis of this species was given in the Victorian Naturalist for June-July, 1891, p. 43.
40 ADDITIONAL OBSERVATIONS ON
of an expedition made to that locality in company with Mr. A. W. Howitt and
Mr. A. H. 8. Lucas in December, 1890. I have much pleasure in naming the species
after the leader of the expedition.
In the shape of the body and the markedly posterior position of the apertures
Geoplana howitt: resembles G. fletcher:.
20. Geoplana lucasi, Dendy.
(PL: -TY,;, Hig. 94")
I found a single specimen of this interesting species under a log near Mount
Wellington, Gippsland, on the same occasion and in the same locality as G. howitti.
As I had not before seen this Planarian alive I was very glad of the opportunity of
making careful notes of the appearance of the living animal, andI am also able to
give a figure of the natural colour.
In life the dorsal surface is white, with dark grey or black markings arranged as
follows :—There is a rather narrow median stripe of black and the remainder of the
dorsal surface is thickly flecked with small specks and longitudinal dashes of dark
grey. On either side of the median black line these specks and dashes are absent,
or nearly so, leaving a very narrow band of almost clear ground colour, then they
appear comparatively large and very close together and more in irregular longitudinal
dashes which seem to be formed each of a number of smali dots. Towards the
margin of the body the dots and dashes get smaller and scarcer, ‘The horse-shoe-
shaped anterior extremity is reddish brown. The ventral surface is greyish white,
with no markings.
When crawling the worm measured about 60 mm. in length.
The specimen after preservation in spirit closely resembles the spirit specimens
from Croajingolong originally described. The eyes appear to extend (in abundance)
for an unusual distance down the sides of the body, if not along the entire length of
the animal, but it is difficult to be certain on this point owing to the resemblance
between the eyes and the smaller specks of pigment.
In concluding these observations I desire again to express my indebtedness to
those gentlemen who have kindly assisted me in collecting specimens of Land
Planarians, viz., the Rev. W. Fielder, Mr. Officer, Mr. Avery, Mr. Fiddian, and Mr.
W. Mann, of the Melbourne University ; Mr. H. R. Hogg, Mr. Hennell, and other
members of the Field Naturalists’ Club of Victoria.
32>
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THE VICTORIAN LAND PLANARIANS. 41
Description oF Prats LY.
Fig. 1—Geoplana m‘mahoni, crawling, dorsal surface. From Fern Tree Gully.
Fig. 2.—Geoplana ventropunctata, crawling, dorsal surface. From Fern Tree
Gully.
Fig. 2a.—Geoplana ventropunctata, at rest, dorsal surface. From Fern Tree
Gully.
Fig. 3.—Geoplana ade, variety with green stripes; crawling; dorsal surface.
From Fern Tree Gully.
Fig. 3a.—Geoplana ade, same specimen as represented in Fig. 3; at rest ; dorsal
surface.
Fig. 4.—Geoplana lucasi, crawling, dorsal surface. From Upper Wellington
River.
Fig. 5.—Geoplana howitti, crawling, dorsal surface. From Upper Wellington
River.
Fig. 6.—Geoplana fletcherz. Ventral surface of posterior end, showing copulatory
organs protruded. From Macedon.
Fig. 7.—Geoplana cerulea. Variety with blue tip. Dorsal surface. From
Mr. Officer, Toorak.
Norr.—All the figures are enlarged. The actual measurements are given in the
text. ;
ARTICLE IV.—Lanp Puianartans From Lorp Hower Isnanp, sy W. Batpwin
Spencer, M.A., Proressox or BroLoay in THE University oF MELBOURNE.
Part [.—Description oF Species. (With Plates 5 and 6.)
(Read Thursday, June 11th, 1891.)
The following paper deals with an interesting collection of land Planarians
which were obtained by Mr. Thomas Whitelegge for the Australian Museum during
a visit paid by a collecting party, despatched by order of the Trustees of the Museum
to Lord Howe Island in 1887.
The material was first placed for examination in the hands of Mr. J. J. Fletcher,
to whom we are indebted for the description of the various species of land planarians
found in New South Wales. Mr. Fletcher being unable to devote the necessary
time to their elucidation kindly offered, with the consent of the trustees of the
Museum, to place the material at my disposal.
The specimens are excellently preserved, and I have had the advantage of the
short notes made by Mr. Whitelegge at the time of collecting them giving some
little information with regard to their colour and length. The specimens all came
from under the leaves of palm trees.
On the Australian continent two genera of land planarians are well-known, viz.,
Geoplana and Rhynchodemus. Of the former 35 species have been described,
and of the latter six species, five from New South Wales, and only one as yet from
Victoria. The latter is known only from one specimen which I collected in
Croajingolong, near to the New South Wales border, and to which Dr. Dendy has
given the name of R. victorie. In addition to these two genera, the introduced form
Bipalium kewense has been recorded by Mr. Fletcher from both New South Wales
and Victoria.
Thus in Australia the genus Geoplana is the predominant one. New Zealand
has as yet been scarcely worked for land planarians and from the Polynesian region so
far as | am aware, the only land planarians recorded come from the Samoan Islands,
LAND PLANARIANS FROM LORD HOWE ISLAND. 43
from which place Grube has described two species of Rhynchodemus, viz.,
R. bistriatus and R, quadristriatus.
Lord Howe Island lies far away from the nearest mainland, in the north of what
it is now proposed to call the Tasman Sea at about one-third of the distance between
the Australian eastern coast and the most northerly point of New Zealand. It lies in
about Lat. 32° S. and Long. 158° EH. and in the line of the comparatively shallow
soundings which indicate probably the existence of an ancient land connection
between New Zealand and the north of the Australian continent. It is interesting
hence to note that as far as our, at present, limited knowledge extends the genus
Rhynchodemus would appear to be more common as we go northwards on the
mainland, whilst there is only one species recorded from Victoria, five are known in
New South Wales, and it will be of interest to see whether a knowledge of Queensland
forms confirms this relative distribution of the two genera Geoplana and
Rhynchodemus. Had a species of Geoplana been at any rate at all plentiful on Lord
Howe Island it would have hardly escaped the notice of so careful a collector as Mr.
Whitelegge though at the same time it must be remembered, judging by experience
in Victoria, that forms commonly met with at one special time may at another be
few and far between. Still there is not in Mr. Whitelegge’s collection a single
example of Geoplana.
The collection proves to contain eight species all, so far as can be ascertained,
new to science. Jam however unable to refer to the description of R. bistriatus and
R. quadristriatus described by Grube from the Samoan Islands. It is just possible
that these may be identical with two of the species now to be described but taking
into account the distance which separates the two islands and the fact that even
Victoria and New South Wales, though so close together, have very few species
indeed of land planarians in common it seems most likely that the species of
Rhynchodemus will be distinct on the Samoan and Lord Howe Islands.
The most interesting part of the collection consists of two species which
must be referred to a new genus for which the name of Cotyloplana is proposed in
consequence of the presence of a distinct sucker-like structure on the ventral surface
anteriorly.
In Part IT. of this paper I propose to describe the anatomy of this genus. The
figures which accompany the paper are drawn from the spirit specimens and to
preserve the relative dimensions proportional compasses have been used: at the
same time it must be remembered that the form of these soft bodied creatures often
changes very much when they are placed in spirit; still the markings and relative
position of the pharyngeal and genital apertures may, together with colour markings,
be taken as discriminative of the species.
44 LAND PLANARIANS FROM LORD HOWE ISLAND.
The following is a list of the forms described :-—
Genus. Cotyloplana, gen. nov.
1. C. whitelegger, sp.n. (Figs. 1, 2, 3, 4.)
2. C. punctata, sp.n. (Figs. 5, 6, 7, 8.)
Genus. Rhynchodemus. (Leidy.)
3. R. fasciatus, sp.n. (Figs. 9, 10.)
R. laterolineatus, sp.n. (Figs. 11, 12, 13.)
R. grandis, sp.n. (Figs. 14, 15, 16, 17, 18, 19.)
R. mediolineatus, sp.n. (Figs. 20, 21, 22, 23.)
R. dubius, sp.n. (Figs. 24, 25.)
t. fletcheri, sp.n. (Figs. 26, 27.)
We ED) Ub
Cotyloplana, gen. nov.
Body flattened. A sucker is present on the ventral surface close to the anterior
extremity. Hyes two.
Cotyloplana whiteleggei, sp.n. (Pl. V., Figs. 1, 2, 3, 4.)
Anterior end somewhat spathulate with the antero-lateral margins of the body
crenate and a median semicircular projection with smooth margin. yes two ;
dorsal ; one at each side of the base of the semicircular projection. A sucker is
placed on the ventral surface at the anterior extremity with its longest diameter at
right angles to the long axis of the body.
The dorsal surface is of a brown colour and the posterior part is covered with
dark mottligs which become restricted anteriorly to form a broad median band
which stops short of the anterior extremity. A dark band les across the anterior
extremity and is continued backwards on each side for about one-fifth of the length
of the body where it merges into the mottlings which cover the whole of the posterior
part of the dorsal surface.
Ventral surface mottled.
Pharyngeal opening 12 m.m. from posterior end. Genital opening 5 m.m. from
to} co) i co)
posterior end. Length in spirits 24 m.m.
Locality.—Lord Howe Island. (Coll. Mr. T. Whitelegge.)
LAND PLANARIANS FROM LORD HOWE ISLAND. 45
The sucker placed on the ventral surface at the anterior end is distinctive of
this genus and the somewhat spathulate form of the head together with the crenate
antero-lateral margins are characteristic of the species. It is possible that this
crenate appearance may be due to the presence of tentacular structures retracted in
the spirit specimens but Mr. Whitelegge makes no remarks in his notes of the
presence of any such nor do I think that they are anything more than crenations.
I have much pleasure in naming the first species of this genus described after
Mr. Whitelegge, and am now engaged in studying its anatomy an account of which
will form Part II. of this paper.
Coiyloplana punctata, sp.n. (PI. I., Figs. 5, 6, 7, 8.)
Body flattened with the posterior half somewhat wider than the anterior half.
Sucker on the ventral surface close to the anterior end, circular in outline, margin of body
smooth. Eyes two, placed close to the anterior extremity. Dorsal surface except
the lateral margins mottled with brown spots and with a median light line which may
or may not extend the whole length. Ventral surface mottled except along the lateral
margins and around the sucker, and somewhat lighter than the dorsal surface.
Pharyngeal opening 9 m.m. from the posterior extremity; genital opening
4 m.m. from the posterior extremity. Length whenalive 50 m.m. Length in spirits
42 m.m.
Locality.—Lord Howe Island. (Coll. Mr. T. Whitelegge.)
This species differs considerably from C. whiteleggei in the shape of its body
whilst both agree in their notably mottled appearance on both dorsal and ventral
surface though there is in the latter no indication of a median light line such as is
present in C. punctata. This varies slightly being in one of the specimens figured
(Fig. 5) continued to the posterior extremity whilst in the other (Fig. 6) it only
passes along slightly more than one-half of the length of the body and there are
indications of another light line on the left of the median one. ‘The margins of the
anterior part of the body are, in spirit specimens, curved over ventrally (Figs. 7 and 8)
making the dorsal, surface distinctly convex and the ventral concave.
The two forms agree however in the presence of a ventral sucker anteriorly
though in C. punctata the margins of this are not so strongly marked as in
C. whitelegget.
46 LAND PLANARIANS FROM LORD HOWE ISLAND.
Genus Rhynchodemus. (Leidy.)
Rhynchodemus fasciatus, sp.n. (Pl. V., Figs. 9, 10.)
Dorsal face marked by a median broad blackish-brown band tapering suddenly
anteriorly ; at either side of this are two dark bands separated by light lines the inner
of the two former being broader than the outer. The edges of the upper surface are
light coloured. The dark bands unite together some little distance in front of the
posterior end leaving a distinct light margin. Eyes two placed close to the anterior
end where the two outer dark bands unite together on each side.
Ventral surface of dun colour with broad darker band in the median line and a
narrow dark line on either side. Dark bands not nearly so prominent as on the
dorsal surface.
Pharyngeal opening 12 m.m. from the posterior end. Genital opening 9 m.m.
from the posterior end. Length when alive, 50 m.m. In spirits 29 m.m.
Locality.—Lord Howe Island. (Coll. Mr. T. Whitelegge.)
The bands as described above are probably much more distinctly marked in
spirit than in living specimens. Mr. Whitelegge notes the colour as blackish brown
on the dorsal surface with two light lines on the laterals, the ventral surface being dun
colour. There are seen very clearly in spirit specimens five dark bands as shown in
the figure (10) dorsally and three bands on the ventral surface though the latter are
faint when compared with the former. A characteristic feature is the way in which
the broad median land suddenly narrows at the anterior extremity whilst posteriorly
it is broad and merges into the lateral ones.
Rhynchodemus laterolineatus, sp.n. (Pl. V., Figs. 11, 12, 18.)
Body much flattened; pointed at both extremities. Hdges of body with
strongly marked dark bands tapering at both ends. Two dark lines along the body
leaving between them a median narrow line of ground colour.
Ventral surface with two dark lines one on either side of the median line and
uniting anteriorly and posteriorly before reaching either extremity. Close to the
two lateral dark bands is a dark line on each side; the two unite together anteriorly
but posteriorly merge into the lateral bands. The dark bands on the ventral surface
may be broad and as many as six be present.
Pharyngeal opening 11 m.m. from post end. Genital opening 6 m.m. from the
posterior end. Length of animal when alive 88 m.m. Length in spirit 25 m.m.
Locality —Lord Howe Island. (Coll. Mr. T. Whitelegge.)
LAND PLANARIANS FROM LORD HOWE ISLAND. AT
This species is marked by its very distinct bands of dark colour along the edges
of the body. The latter is much flattened, broad in the middle and tapering to both
ends ; there is no difference in the thickness of the body at any part, and the lateral
margins are very distinctly marked. Whilst the dorsal surface is the same in all the
specimens the ventral varies. Figures 12 and 13 serve to show the two extremes.
The important dark bands consist (1) of two, one on either side the median line and
(2) of two, one close to each of the lateral margins each of these falling into the
lateral band before the posterior end is reached.
In addition to these there may be present two more bands one on each side
intermediate between those already described. In addition to varying in number
ventrally the thickness of the bands may vary considerably, with the exception of the
outermost one which always remains thin. ‘Thus in figure 13 the specimen drawn
has four thick bands and the two lateral thin ones, whilst in figure 12 the specimen
figured has only two thin median and the two thin lateral ones. In both cases the
dorsal surface resembled the one drawn in figure 11.
Rhynchodemus grandis, sp.n. (Pl. VI., Figs. 14, 15, 16, 17, 18, 19.)
Dorsal surface with the ground colour fawn-brown, marked with streakings and
spots of darker brown. The anterior end with the margins light-coloured and two
dark bands, one on either side, which gradually become less marked posteriorly.
They enclose between them at the front end a median light line, and the streaks
show a tendency to form two dark lines one on either side the median line which
remains light-coloured. Eyes two, one on either side in the lateral light lines close
- to the anterior somewhat bluntly pointed extremity. Ventral surface of a lighter
fawn-yellow colour, spotted with a prominent median line of dark spots. Pharynx
35 m.m. from the posterior end. Genital opening 16 m.m. from the posterior
end. Length when alive 6-7 inches, width 2-2 inches.
Locality.—Lord Howe Island. (Coll. T. Whitelegge.)
In spirit specimens the lateral edges of the body become curved towards the
ventral surface so as to produce a distinct: concavity ventrally, but this only extends
for a short distance.
The specimen figured (Pl. VI., Figs. 14, 15) represents the animal very much
contracted after the action of spirit, but this is probably one of the largest examples
of the genus yet known. Some two or three of our Australian land-planarians of
the genus Geoplana, notably G. dendyi and G. hoggii, reach a length when crawling
equal to that of R. grandis, but the width of the body when fully expanded is
nothing like so much as five-eighths of an inch.
48 LAND PLANARIANS FROM LORD HOWE ISLAND.
In Mr. Whitelegge’s collection are some specimens which are in all probability
young forms of the species. These are interesting as showing, what Professor
Moseley* has already drawn attention to, the curious fact that the markings are
often more definite in young than in old and full-grown examples. Two young speci-
mens are represented in figures 16,17,18, and 19. The first of these shows dorsally
three stripes of dark colour, (1) a median one and (2) two lateral ones. In the older
form these stripes, as it were, break up into a series of streaks and spots, but the
two lateral ones persist anteriorly, whilst the median one becomes more obscure and
represented by two lines of darker streaks. Ventrally also three dark but not so
prominent stripes are present each composed of closely placed spots.
The second shows dorsally the lateral stripes strongly marked only anteriorly,
the median one being much fainter, whilst ventrally the lateral ones are only seen at
the anterior end, and the median one is much as in the adult. In both of the young
specimens figured only the pharyngeal opening is visible, the genital opening not
being apparently developed presumably on account of the immaturity of the
specimens.
Rhynchodemus mediolineatus, sp.n. (Pl. VI., Figs. 20, 21, 22, 23.)
Dorsal surface with a median dark stripe which may or may not extend the
whole length of the body and on either side of which is a dark stripe anteriorly. The
lateral edges are marked by dark lines. Two eyes close to the anterior end.
Ventrally are two dark stripes one on either side the median line uniting anteriorly.
Pharyngeal opening 12 m.m. from the posterior end. Genital opening 6 m.m. from
the posterior end.
Locality.—Lord Howe Island. (Coll. Mr. T. Whitelegge.)
This species varies somewhat in the development of the stripe just as does the
species Geoplana mediolineata, which is distinguished by its median stripe. Two
specimens are figured. In the first of these (Figs. 20, 21) the three stripes are only
present at the anterior end dorsally whilst ventrally two extend for the whole length
and enclose the pharyngeal and genital openings uniting together at both ends.
In the second (Figs. 22, 23) the anterior dorsal end is darker than in the first and
the dorsal median stripe extends the whole length whilst ventrally the two stripes
are only present for not quite one-third of the length. The genital opening in the
latter was indistinguishable.
Rhynchodemus dubius, sp.n. (Pl. VI., Figs. 24, 25.)
Dorsal surface with ground colour of pale cream colour, anterior end with four
dark lines extending back not quite one-fourth of the length of the body the two
* Trans. R. 8. London. The Anatomy and Histology of the Land Planarians of Ceylon. Vol. 164, 1874, p. 110.
LAND PLANARIANS FROM LORD HOWE ISLAND. 49
middle ones being more prominent than the outer ones and enclosing anteriorly
a median light line of ground colour. The dark lines fuse at the anterior extremity
leaving the margins of the body light coloured. The two eyes are placed in the dark
lines where the four have fused close to the anterior end. The posterior three quarters
of the body are marked by irregular, undulating, disconnected, dark lines the more
prominent of which seem to forma broken up continuation of the two inner of the
stripes. A median portion nearly one-third of the width of the surface is light
coloured and there is present in this a median thin dark line made up of slightly
discontinuous parts. The streaks have a tendency to form four lines at the very
posterior end. Ventral surface pale cream colour with no streaks. Pharynx 36 m.m.
from the posterior end. Genital opening 12 m.m. from the posterior end.
Locality.—Lord Howe Island. (Coll. Mr. T. Whitelegge.)
The length of the animal when alive is 4-5 inches. The spirit specimens are all
much coiled and vary somewhat in marking especially in the distinctness of the four
stripes anteriorly and in the development of the thin dark median stripe. In one or
two this is very difficult to detect and it may be very irregularly developed. It is not
apparently at all prominent during life. The characteristic features are the lines
giving place to streaks posteriorly and the entire absence of markings on the ventral
surface.
Rhynchodemus fletcheri, sp. un. (Pl. VI., Figs. 26, 27.)
Dorsal surface with median light line and two broad black bands one on either side
of this. Lateral portions of body and ventral surface light coloured. Ventral surface
with two lateral dark lines and median broad punctated band not extending quite to
the anterior or posterior ends and showing indication of division into two lines.
Anterior extremity bluntly rounded with: eyes dorsal and close to the anterior edge.
Pharyngeal opening 8 m.m. from the posterior end; genital opening 3 m.m. from
the posterior end.
Locality.—Lord Howe Island. (Coll. Mr. T. Whitelegge.)
The animal when alive measures 25 m.m. in length the spirit specimen
measuring 19 m.m. In the latter the pharyngeal opening has the form of a transverse
slit with a light margin lying in the middle of the median dark. band; the genital
opening is distinct and more noticeable than the pharyngeal and has a light-
coloured circular margin the dark band being divided in its region into two by a
median light line.
The body in spirit specimens is somewhat flattened and band like the anterior end
blunt and rounded the posterior more pointed.
The median dorsal line swells out slightly on the head and does not quite reach
the posterior end.
50 LAND PLANARIANS FROM LORD HOWE ISLAND.
DEscRIPTION oF PuLaTEs.
Puate V.
Fig. 1.—Cotyloplana whiteleggei, dorsal view. x 8.
Fig. 2.—Cotyloplana whiteleggei, ventral view of the same. x 3.
Fig. 3.—Cotyloplana whiteleggei, enlarged view of the dorsal surface of the
anterior extremity to show the semicircular end with the crenated antero-lateral
margins and the two eyes. x 7.
Fig. 4.—Cotyloplana whiteleggei, enlarged view of the ventral surface of the
same specimen as figure 3, to show the sucker with its slit at right angles to the long
axis of the body and prominent margin. x 7.
Fig. 5.—Cotyloplana punctata, dorsal view. x 2.
Fig. 6.—Cotyloplana punctata, dorsal view of a larger specimen in which in
addition to the median light line which does not extend the whole length of the
body there is an indication of another light line at the side of the median one. x 1}.
Fig. 7.—Cotyloplana punctata, ventral view of the same specimen as figure 6,
showing the sucker anteriorly. x 1}.
Fig 8.—Cotyloplana punctata, enlarged ventral view of the anterior extremity
of the specimen represented in figure 6 to show the sucker. In figures 6, 7, 8, the
lateral margins are seen to be strongly contracted so as to produce at the anterior end
a very concave surface ventrally and convex dorsally.
Fig. 9.—Rhynchodemus fasciatus, ventral view. x 3.
Fig. 10.—Rhynchodemus fasciatus, dorsal view of the same. x 3.
Fig. 11.—Rhynchodemus laterolineatus, dorsal view. x 2.
Fig. 12.—Rhynchodemus laterolineatus, ventral view of the same. x 2.
Fig. 18.—Rhynchodemus laterolineatus, ventral view of another specimen, in
which the dark bands ventrally are strongly developed. x 3.
LAND PLANARIANS FROM LORD HOWE ISLAND. 51
Prats VI,
Fig. 14.—Rhynchodemus grandis, dorsal view. Actual size of spirit specimen.
Fig. 15.—Rhynchodemus grandis, ventral view of the same. Actual size of spirit
specimen.
Fig. 16.—Rhynchodemus grandis, young specimen, dorsal view. Actual size of
spirit specimen. |
Fig. 17.—Rhynchodemus grandis, young specimen, ventral view of the same.
Actual size of spirit specimen.
Fig. 18.—Rhynchodemus grandis, young specimen, dorsal view. Actual size of
spirit specimen.
Fig. 19.—-Riynchodemus grandis, young specimen, ventral view of the same.
Actual size of spirit specimen.
Fig. 20.—Rhynchodemus mediolineatus, dorsal view. x 38.
Fig. 21.—Rhynchodemus mediolineatus, ventral view of the same. x 3.
Fig. 22.—Rhynchodemus mediolineatus, dorsal view of another specimen in
which the median line passes along the whole length of the body. x 3.
Fig. 23.—Rhynchodemus mediolineatus, ventral view of the same, the two lines
only being present at the anterior end of the body.
Fig. 24.—Rhynchodemus dubius, dorsal view. x1}.
Fig. 25.—Rhynchodemus dubius, ventral view of the same. x 1}.
Fig. 26.—Rhynchodemus fletcheri, dorsal view. x 3.
Fig. 27.—Rhynchodemus fletcheri, ventral view of the same. x 3.
Norrt.—The blue tint in figures 14, 15, 16 and 18 should not be present.
Vol. Hl. Plate 5.
Trans. RS. Victoria
= a naeemaneniaenel storia tlio Soo re
Seer pndedann prt Manan simanern ene te wree n=
W.B.Spencer del
i
LORD HOWE ISLAND - LAND PLANARIANS
19
Voll. Plate oO.
18
Trans. RS. Victoria
19
OI
NI
LORD HOWE ISLAND - LAND PLANARIANS.
WB. Svence
TRANSACTIONS.
VOLUME Il.
Part I.—6s.
THE ANATOMY OF MEGASCOLIDES AUSTRALIS (THE GIANT
EARTHWORM OF GIPPSLAND), sy Prorssson W. Baupwin Spencer, M.A.
Plates 1 to 6.
Part I1.—12s. 6d.
1.—RECORDS OF OBSERVATIONS ON SIR WILLIAM MACGREGOR’S ~
HIGHLAND PLANTS FROM NEW GUINEA, sy Baron Von Muzzuzr, F.R.S.
2.—THE PREPARATION OF AULKYL-SULPHINE, SELENINE, AND
PHOSPHONIUM SALIT'S, sy Prorgssor Orme Masson, M.A., D.Sc., ann J. B.
Kirxuanp, F.C.S.
3.—IHE ANATOMY OF AN AUSTRALIAN LAND PLANARIAN, sy
Artuur Denpy, D.Sc. Plates 7 to 10. .
4—ON THE ORGANISATION OF AUSTRALIAN TRIBES, sy A. W.
Howirr (with Map).
5.—THE ANATOMY OF AMPHIPTYCHES URNA, sy Proressor W.
Batpwin Spencer, M.A. Plates 11 to 18.
VOLUME Il.
Part I.—17s. 6d.
1.—THE TRANSVERSE SECTIONS OF PETIOLES OF EUCALYPTS
AS AIDS IN THE DETERMINATION OF SPECIES, sy D. McAupine anp
J. KR. Rempry. Plates 1 to 6a.
2.—THE VICTORIAN LAND PLANARIANS, sy Arraur Denny, D.Sc.
Plate 7.
3.—THE EUCALYPTS OF GIPPSLAND, sy A. W. Howirr. Plates 8 to 16.
4.—A NEW FAMILY OF HYDROIDEA, TOGETHER WITH A DES-
CRIPTION OF THE STRUCTURE OF A NEW SPECIES OF PLUMULARIA,
BY Prorgessor W. Batpwin Spencer, M.A. Plates 17 to 23.
VOLUME Ill.
Part U.—7s. 6a:
MONOGRAPH OF THE VICTORIAN SPONGES, sy Artuur Denpy, D.Sc.
Part I., THE ORGANISATION AND CLASSIFICATION OF THE
CALCAREA HOMOCGILA WITH DESCRIPTIONS OF THE VICTORIAN
SPECIES. Plates 1 to 11.
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BOUND JAN 1973
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