JOURNAL OF
THE ROYAL SOCIETY
OF
WESTERN AUSTRALIA
INCORPORATED
VOLUME 45 (1962)
PART 1
PUBLISHED 30TH MARCH, 1962
REGISTERED AT THE G.P.O., PERTH FOR TRANSMISSION BY POST AS A PERIODICAL
THE
ROYAL SOCIETY
OF
WESTERN AUSTRALIA
INCORPORATED
COUNCIL 1961-1962
President
Past President
Vice-Presidents ....
Joint Hon. Secretaries
Hon. Treasurer .
Hon. Librarian ....
Hon. Editor
J. E. Glover, B,Sc., Ph.D.
N. H. Brittan, B.Sc., Ph.D.
W. D. L. Ride, M.A., D.Phil.
C. F. H. Jenkins, M.A.
R. W. George, B.Sc., Ph.D.
C. V. Malcolm, B.Sc. (Agric.).
L. J. Hollis, B.A., Dip. Com.
G. G. Smith, M.Sc.
J. E. Glover, B.Sc., Ph.D.
J. G. Kay, B.Sc.
R. J. Little.
D. Merrilees, B.Sc.
R. T. Prider, B.Sc., Ph.D., M.Aust.I.M.M., F.G.S.
R. D. Royce, B.Sc. (Agric.).
L. W. Samuel, B.Sc., Ph.D., F.R.I.C., F.R.A.CJ.
D. W. Stewart, B.Sc. (For.).
W. R. Wallace, Dip. For.
Journal
of the
Royal Society of Western Australia
Vol. 45 Part 1
1. — Variation, Classification and Evolution in Flowering Plants —
with particular reference to Thysanotus
Presidential Address, 1961
By Norman H. Brittan, B.Sc., Ph.D.*
Delwered — 17th July, 1961
The sources of variation in flowering plants
are briefly discussed and illustrated by examples
from the genus Thysanotus. In two species—
T. patersonii and T. tuberosws— evidence of
variation is presented which supports the estab-
lishment of sub -specific taxa. The occurrence
of Intra-speciflc polyploidy is reported and com-
pared from the point of view of geographical
distribution with other selected Australian
genera.
The world distribution of the tribe Asphodelae
in terms of its constituent genera and the dis-
tribution of the species of Thysanotus are
considered in the light of Willis’ Age and Area
theory and of a recent hypothesis of Smith
White. A hypothetical evolutionary history of
the genus Thysanotus is presented.
This address will be concerned with plant
systematics and will indicate the various lines
along which research into what may in general
be called “the species problem” has been con-
ducted and how the results from such investi-
gations have affected the science of flowering
plant taxonomy and thrown light on pathways
in evolution.
In systems of classification of flowering plants
there had grown up from the time of Linnaeus
(1753) the idea that each individual species was
an invariable entity and out of this concept
arose the use of the “type” — a single plant
specimen located in some particular herbarium
which provided a fixed reference point and
which, taken in conjunction with the written
description of the species, enabled subsequent
\vorkers to identify plants of the particular
taxon. This may be an unfamiliar word to
some of you — it means a particular taxonomic
entity at whatever level is under discussion, for
example — species, genus, etc.
Over the last thirty years concurrent with the
rise of genetics as a science there has been a
marked increase in the interest taken in varia-
tion in plants and as a result there has
developed what has variously been called the
“new systematics” (Huxley 1940), “experimental
taxonomy” (Clausen, Keck and Hiesey 1941) and
“biosystematy” (Camp and Gilly 1943). The
results from such investigations have not so far
resulted in a change in the orthodox taxonomic
* Dept, of Botany. University of Western Australia,
Nedlands. Western Australia
methods nor in a rejection of the type system,
although various authors as for example Gil-
mour and Gregor (1939) and Gilmour and
Heslop-Harrison ( 1954) have suggested a
nomenclatural system which would assist in
codification of the various categories among
plants recognised as a result of experimental
taxonomic investigation.
The point of departure between what has
been called “classical” or “alpha” taxonomy
and experimental taxonomy is the attention
paid to the variability of the plant species as a
result of intrinsic and extrinsic factors. The
interrelationship of the two approaches is shown
in Table I modified from Heslop-Harrison
(1953).
In order that we may better understand the
relationship between these two approaches let
us look first at seme of the sources of variation
in the flowering plant. Most people are fami-
liar with the basic cell of which the complete
structure of the plant is made up. It is a
complex assemblage of inorganic and organic
chemical compounds in an aqueous medium
contained within a membrane enclosed by a
cellulose cell wall. Among the important
organic chemical compounds are proteins,
ribose nucleic acid (RNA) and desoxyribose
nucleic acid (DNA). The proteins are import-
ant because of their part in building the fine
structure of the cell and their association w'ith
the enzyme system. Together with the nucleic
acids just mentioned they form nucleoproteins
of which DNA-proteins make up the genetic
units of the cell — the chromosomes — which can
be made visible by suitable methods, and RNA-
proteins make up microsomes in the cytoplasm
and also form other so far invisible (except in
the electron microscope) cytoplasmic bodies.
Genetics has established that the chromosomal
part of the genetic system is distributed in a
regular fashion by meiosis prior to the forma-
tion of embryo sacs and pollen grains and that
at this stage in the life history there is the
possibility of variation in the genetic constitu-
tions of the various products of meiosis. Here
then we have the first method by which varia-
tion can occur.
1
TABLE I
The relationships of ^"classical' and ''experimental taxonomy (after Heslop-Harrison 1953)
AIMS
UNIT OF STUDY
CLASSICAL TAXONOMY
DescHptlon, classification and nam-
ing.
Individuals, usually dead.
SYSTEM OF
CLASSIFICATION
SOURCE OP DATA
TESTS OP CHARACTERS
METHODS OF
DESCRIPTION
CONCEPT OF NATURAL
VARIATION INVOLVED
Basic unit the species, fitted into
a hierarchical system.
Morphology, anatomy plus notes on
flowering time and geographical
distribution.
Intuition and trial and error.
Individuals and “type” concept.
Essentially static, assumes continu-
ity of form within species.
EXPERIMENTAL TAXONOMY
Identification of evolutionary units, determina-
tion of genetical relationships.
Breeding population, or a sample from it,
living.
Classification not the primary aim. basic units
vary.
Morphology, anatomy, karyology, life cycle de-
tails. effects of habitats, genetical tests.
Statistical methods.
Populations or samples of populations.
Essentially dynamic, internal variability of
sexual populations is recognised, adaptive
nature of much population differentiation
acknowledged and made deliberate object of
study.
Whether the process of meiosis always results
in variation depends upon the genetic state of
the organism, whether it is homozygous or
heterozygous and also on the breeding system
of the plant. With regard to the latter there
are three possibilities, (1) inbreeding (or self-
pollination*, (2) outbreeding, or cross-pollina-
tion t and (3) apomixis — a single term covering
a number of possible variants of the sexual pro-
cess. To deal with the last of these first —
apomixis is a mechanism which results in the
elimination of variation resulting from meiosis
and conversely the maintenance of variation
occasioned by means other than meiotic re-
combination. An extreme form of apomixis is
the replacement of sexual reproduction by vege-
tative propagation.
To return to the first two. the choice between
in- or out-breeding may be controlled by either
the floral morphology of the plant or a specific
part of its genetic make-up, or possibly both.
As an example the pea flower is anatomically
designed for self -fertilisation since the stigma
is receptive at the time of anthesis, and also in
close proximity to the anthers. The cleisto-
gamous flower of the violet — since it never opens
—also ensures self-fertilisation. On the other
hand distylous flowers— types of flowers which
are found with two possible lengths of style
alternating with two possible anther positions —
are spatially arranged to encourage cross-polli-
nation by insect visitors as for example the
primrose (Primula^ and loosestrife (Lythrunv.
In some of these cases the anatomical mecha-
nism is reinforced by a genetical system of
incompatibility genes.
Of anatomical mechanisms for ensuring cross-
pollination may also be mentioned the system
in some members of the Compositae (the daisy
family) in which pollen is shed into a central
tube and released to visiting insects as a result
of the style growing up through the tube. When
the style has grown beyond the limits of the
tube the divided apical parts of the style re-
curve and expose the inner receptive surfaces
of the stigma which then becomes receptive to
other pollen. Genetic systems for ensuring
cross-pollination are based on the “S” gene
series which, by means of interaction between
pollen carrying a specific allele of the series and
diploid stylar tissue carrying the same allele
results in retardation or complete inhibition of
the growth of the pollen tube or even inhibition
of pollen germination upon such stigmas. Other
methods are known which achieve the same end
as for example in the evening primrose
f Oenothera f in which both ovules and pollen
grains containing particular genetic combina-
tions (the so-called Renner complexes) do not
develop. This is a system of gametic lethals;
alternatively where particular zygotic genetic
combinatiens do not develop, a system of
zygotic lethals is found.
What then can be said from the point of view
of variation about the results of the various
breeding systems just mentioned? Apomixis will
tend to reduce variation, or if variation results
from mutation it will perpetuate it without
change — it is therefore a system which imposes
restrictions upon the variability of the plant
possessing it. Obligate self-pollination will re-
sult in homozygosity after relatively few genera-
ions. since from inbreeding a heterozygote fol-
lowed by self-pollination of all the progeny
(neglecting any form of selection for a particu-
lar genotype) the proportion of heterozygotes;
homozygotes becomes progressively 1:1, 1:3, 1:7,
etc. Put in another way the proportion of
heterozygotes in such a population would be
represented by ( 2 )^ where n number of
generations of inbreeding. We see in this a
very rapid reduction in the number of heterozy-
gotes in such a population. Only with outbreed-
ing is there the possibility of segregation and
1 ‘ecombination and thus the persistence of
heterozygosity. We therefore expect that the
self -fertilised species will possess little or no
genetic variation within its populations derived
from a single individual whereas the out-
breeding species will show marked variability.
So far we have been concerned with genetic
variation, which is controlled by the chromo-
somally-borne genes. The whole chromosome
complement of the plant may be altered by the
incidence of polyploidy— in the simplest case the
duplication of the chromosomal set to produce
a tetraploid. This increase in chromosomal
material may or may not show itself in the
2
morphology or physiology of the plant. There
is a great deal of literature now available on
this topic and one may generalise to the extent
that increase in size of various parts, for
example flowers, frequently coupled with reduc-
tion in fertility, often occurs following poly-
ploidy.
Variatioii may also arise from environmental
factors. This was thought at one period to be
the reason for the identity of allied species and
claims have been made that by transplanting
individuals from their normal habitat to an-
other they could be shown to be transformed
Into another species typical of the new habitat.
The experiments of Bonnier (1895) were claimed
to show the transformation of lowland species
into alpine ones. Due to lack of careful culti-
vation methods and keeping of accurate records,
the results from these experiments are now
viewed with suspicion. Subsequently the
transplantation technique has been used by
such workers as Turesson in Sweden, Gregor in
Scotland and Clausen. Keck and Hiesey in Cali-
fornia. Not only have reciprocal transplants
been used, but the concept of growth under
uniform environment of plants from diverse
ecological situations has been extensively used.
The use of this approach under most critically
controlled conditions has been in the use of
what have been termed “phytotrcns*’ — each
consisting of a collection of rooms with either
natural or artificial lighting in which strict
control of day and night temperatures, humidity
and day length is possible. Under such condi-
tions it has been shown by Went (1953) in the
case of the Earhart Laboratories in California
that variation due to environment can be vir-
tually entirely removed and that as a result
genetical experiments which would normally re-
quire many replications to ensure statistical
accuracy can be conducted with very few plants.
This type of apparatus provides an ideal site
for researches into physiological differences be-
tween races of plants as for example the work of
Hiesey (1953) on Achillea.
The sources of variation have now been men-
tioned, but what of their effect upon the plant
species? Genetic variation may provide sharply
discontinuous morphological changes as for
example in flower colour. A very common
example of this is the mutation to white flowers
in a species normally possessing coloured flowers.
In Thysanotus this has been found in several
species, e.g., T. triandrus (Shearer, personal
communication), in T. tenuis near Tinkurrin,
Western Australia, and T. tuherosus at Virginia,
Queensland. In most cases the change to
albinism is complete and affects all parts of the
flower. In the case of the Tinkurrin plants
however, there seem to be two stages each pre-
sumably controlled by a separate gene. In one
of these the anthocyanin pigment is absent from
the perianth parts but remains in the anthers
< which in this and most other Thysanotus
species are normally purple coloured) ; in the
other, anthocyanin is absent both from perianth
and anthers, the latter then appear green.
Mutation of the normal pale purple flower colour
to a salmoir pink has also been seen in a popu-
lation of T. tuherosus in southern Queensland.
A presumed mutational effect which concern.s
the morphology of the perianth in T. vdtersonii
is reported from localities in northern South
Australia and southern Northern Territory. In
this case the fringed margin to the inner
perianth segments is absent and led Black
(1943) to use the manuscript name of var.
exfimbriatus for such plants. Live material has
recently been obtained from a population of this
plant thi*ough the courtesy of Mr. Paiol Wilson
of the South Australian State Herbarium and
it is hoped in the forthcoming season to be able
to determine whether environment changes the
expression of this character. By experimental
cross-pollination with normal fringed types it
is hoped to be able to determine the genetical
relationship between the normal and the ex-
fimbriate foi*m.
Genetic variation may also result in con-
tinuous as opposed to discontinuous morpholo-
gical variation giving the so-called quantitative
variation shown mostly in characters such as
the overall size of plant parts. This aspect of
plant variation has been extensively studied by
the Californian workers Clausen, Keck and
Hiesey (Clausen 1951 >. The results shown in
Figure 1 are a sample of the type of results ob-
tained when plants from various localities on
a transect across California are grown under
uniform conditions at the Carnegie Institute
experimental plots at Stanford, California. Each
plant shown in the figure represents the mean
of the population from that locality and the
histograms show the variation within that popu-
lation. If individual plants from such popula-
tions are selected and vegetatively propagated
by means of ramets and grown under identical
conditions at each of three localities, results
such as those shown in Figure 2 can be ob-
tained. This shows that the responses of
genetically different plants are quite different
at the three localities and that among the ver-
tical lines of plants which represent the ramets
derived from a single individual, there are
marked differences in the response of the plant
to the environment at the three localities used
in the experiment. At any one particular habi-
tat the sum of the variation adds up to the
total variation shown by the histograms in the
previous figure.
As a result of these approaches together with
experimental crossing of races from different
geographical and ecclogical habitats it has been
possible to elucidate the genetical and physio-
logical bases for the observed morphological
differences among members of a particular
species.
As yet no extensive experiments of this sort
have been carried out with Thysanotus — some
transplanting to a more or less uniform en-
vironment in a Perth glasshouse has been
undertaken and progeny have been raised from
seed derived from selfed parents. Prom the
latter the indications are that there is little
variation in such characters as size of plant
w'ithin the progeny. This lack of variation in
sexually produced progeny would lead one to
suspect that the plant was most likely to be
normally self-fertilised in nature.
3
Fig. 1. — Altitudinal climatic races of Achillea from a west-to-east transect across central California. The fre-
quency diagrams indicate variation in height within the populations in the Carnegie Institution plots at
Stanford, California. The plant specimens represent the means, (after Clausen, Keck and Hiesey 1948.)
4
Fig. 2. — Responses
planted to the
of seven cloned individuals taken from a population at Mather (alt. 4,600 ft.) when trans-
environments at Stanford, Mather and Timberline, (after Clausen, Keck and Hiesey, 1948.)
5
Here we may make a digression to look at the
floral morphology of the plants. The flowers
are either three or six stamened and typically
the anthers are curved and tend in the open
flower to lie with their apices near the terminal
stigmatic part of the style. In the majority of
the species the anther opens by a terminal pore
and it is thought that there is a "pepper pot”
pollination mechanism in these cases. Such
a mechanism does not provide large quantities
of pollen loose in the flower. In a few species
the anthers open by longitudinal slits and in
these cases there is much more pollen freed at
any one time. In these latter cases also there
is close proximity of style and anther and pol-
lination by direct mechanical transfer of pollen
is suspected. Insects are hardly ever seen visit-
ing the flowers and emasculated non-bagged
nowers in a glasshouse in the vicinity of other
open non-emasculated flowers of the same
species do not set seed. The evidence therefore
suggests self-fertilisation as the normal condi-
tion for the plant.
If instead of considering variation in height
on an intrapopulation basis we look for inter-
population variation within the species we find
examples of this in T. trimidrus and T. 7tiulti~
fiorus. In the former species the variation was
such that Dornin <1913> erected a series of
varieties of which the smallest — from the
vicinity of Youndegin (35 miles east of York,
Western Australia) had been given specific rank
by Ewart and White ( 1908 ) as T. bentianus. If
these variations in height are plotted geographi-
cally a positive correlation appears between
decrease in height and decrease in annual rain-
fall. It would seem that in this case we have
a dine — defined as a gradual transition in a
characteristic associated with geographical or
ecological change. A similar pattern emerges
from a study of height in T. multifiorufi which
shows reduction in size correlated with progres-
sively south-western distribution. These varia-
tions may be merely plastic responses to the
environment, this can be proved or disproved
only by determining whether the height typical
of the original population is maintained when
plants are grown under more or less uniform
conditions. If changes of environment do not
bring about changes in the height of the plants
the genetical control of the character is demon-
strated. If it is possible to select types morpho-
logically different from others in the dine the
term ecotype of Turesson could be applied to
such plants. Ecotypical variation implies that
genetic modification has occurred in response to
the environment with mcdification in some
characters as for example size, but not to the
extent that the plants showing this variation
should be placed in different taxonomic groups.
The work of Silsbury and Brittan (1955) demon-
strated the existence of ecotypical variation in
Kenncdya prostraia.
As a general rule it is to be expected that if
a species covers a wide geographical range,
which is therefore likely to include several dif-
ferent ecological conditions together with the
possibility of isolation into small populations
there is greatei* likelihood of the development
of infraspecific categories. In ThysanoUts the
two most widely ranging species are T. tuberosum
and T. patersonii. The former has a distribu-
tion which ranges from Cape York and northern
Northern Territory through Queensland, New
South Wales, Victoria and to the east of the
Mount Lofty Ranges in South Australia. The
typical form over this range has flowers with
perianth segments 11-19 mm long, six stamens
— of which three have short (3-3.5 mm), more
or less straight anthers and three with long
)6-9 mm» curved anthers. The longer ones are
comparable in length to the style. This floral
morphological pattern is constant over most of
the range but may be accompanied by variation
in overall size and vigour of the plants. In the
area around Brisbane and possibily south into
New England there occurs a distinct form with
perianth parts 7. 5-9.5 mm long, six stamens
almost equal in length — three with anthers
2.5-3 mm long, three with 3-5 mm anthers —
arranged more or less parallel to the open
perianth parts. The differences were first re-
cognised in live material sent from Brisbane
through the courtesy of Dr. Blake and Mr. Coal-
drake. The morphological differences were
found to persist in cultivation. In an attempt
to elucidate the relationship between these types
artificial cross-pollinations between them have
been attempted. The results to date show that
both types are self fertile and that crossings in
which the large flowered form is the female
result in no seed production, whereas the re-
ciprocal (small flowered female) sets seed. So
far plants have not been raised from such ex-
perimental crosses.
During an extended collecting trip in 1959 I
was able to examine several habitats for T.
tuberosus near Brisbane. Plotting the occur-
rence of the two types gave a distribution in
which the types replaced each other without
obvious reason at intervals in a north-south
direction (the sampling having been carried
mostly along the Brisbane-Maryboi*ough i*oad
which runs more or less north-south). The
area is mostly occupied by “wallum” — coastal
plain country on which grows Bayiksia aeviula
and Melaleuca leucadendra (sensu lato). There
are slight altitudinal variations of the order of
100 feet, the lower parts tending to be water-
logged. From the study of the distribution of
the two types the pattern emerges that on the
lower wetter parts the smaller flowered type is
found and that with increase in elevation it is
replaced by the larger flowered type. In an
area of c. 300 X 10 yards at Beerburrum both
forms were found in ratios varying from 8:0
( large :small) to 1:9 over the area. Again the
higher proportion of the smaller flowered type
was found on the lower parts. The sampling
was obviously confined to the plants in flower
at the time and does not represent a complete
sample of the population. It demonstrates that
the two types are distinct and are able to main-
tain their identities in a mixed population. It
is proposed on the basis of the evidence pre-
sented to separate the types at the subspecifle
level, the large flowered one to be known as T.
tuberosus subsp. tuberosus and the smaller
flowered type as T. tuberosus subsp. blakei. In
the same tuberous-rooted, paniculate inflores-
cenced group Robert Brown (1810) described
T. elatior — a taller, more robust plant with
6
larger flowers collected from the north coast of
Queensland and islands in the Gulf of Car-
pentaria. Similar plairts have been found by
me in Inland New South Wales and although no
experimental work has been done it is thought
that the morphological variation does not merit
specific rank and that until the results of in-
vestigations are known such plants should be
included within the subspecies tuherosus.
By comparison with the distribution of T.
tuherosus which shows a mainly north and
south trend, the distribution of T. patersonii
shows largely an east-west trend occurring as
it does in scuthern New South Wales, Victoria,
Tasmania, South Australia, southern Northern
Territory and extra-tropical Western Australia.
It is worth noting that on present knowledge
it is the only widespread extra-tropical
Thysanolus species recorded from eastern and
western Australia, in addition it is the only
.species so far recorded from Tasmania, whence
the type specimen was collected. Kunth in his
Enumeratio G861) delineated the species 7’.
manglesianus based on a specimen collected
from New Holland by Captain Mangles, in the
course of the description mention is made that
a specimen collected by Gaudichaud from the
Vasse River (near Busseltcn. Western Austra-
lia) is identical with the type. Subsequent
workers as for example Baker (1877) have sub-
merged T. ma^ujlesianus within T. patersonii.
One of the distinctions made by Kunth was the
larger flower size in his specimens compared to
Brown's type specimens from Tasmania. My
researches have shown that in addition to larger
flower size the anthers of the inner and outer
whorls of stamens are dissimilar in size, whereas
those in T. paterscnii (sensu stricto) are nearly
equal in length. An additional morphological
character is the form of the ripe capsule which
in the small flowered forms has an overall
length of c. 5 mm and has the persistent
perianth parts free at the tips whereas in the
large flowered form the overall length is c. 13
mm of which c. 8 mm is made up of the per-
sistent perianth parts closely appressed and
.'ilightly twisted together. The size of the cap-
sule itself in the two forms is almost the same.
The two forms are found growing in nearby
localities in Western Australia, but so far no
evidence has been obtained of the presence of
intermediates. As mentioned above some species
of Tliysariotus possess anthers which dehisce by
longitudinal slits, T. patersonii is one of these
and as a result of the larger amounts of pollen
shed it is less likely that cross-pollination be-
tween types would occur in this species. I there-
fore propose to re-establish manglesianus as an
epithet at the subspecific level to describe the
large flowered, western form of T. patersonii.
The small flowered plant as exemplified by
Brown’s Tasmanian collection becomes T.
patersonii subsp. patersonii.
In T. patersonii it has also been possible to
demonstrate the existence of polyploidy — the
occurrence in plants of chromosome numbers
greater than the typical diploid number which
for Thysanotus is 22. A chromosome count of
c. 80 was reported on Tasmanian material
(Jackson, unpublished data) and subsequently
in association with Jackson I was able to show
that this was 88 — the plant would therefore be
termed an octoploid. There is in this case no
morphological difference from other plants of
T. patersonii, we therefore have an example of
a chromosome race whose identification is only
possible from microscopic examination rather
than from macroscopic examination — the latter
the more normal technique in herbaria. So far
no chromosome numbers intermediate between
diploid and octoploid have been found, one
would expect that — unless they had been lost in
the evolutionary process — there would be
tetraploid types in existence. Live plants have
been obtained from various localities in Tas-
mania and northward on the mainland towards
Newcastle, N S.W. — -which seems from herbarium
records to be the northernmost limit for the
species in eastern Australia. Cytological in-
vestigation of these plants may, it is hoped,
provide evidence of the existence of the tetra-
ploid race. A similar situation of chromosome
polymorphism without morphological differen-
tiation has been found by James (unpublished
data) working cn Western Australian popula-
tions of Isoloma.
Hagerup (1932) stated that in Uie Northern
Hemisphere more northerly populations po.sscss
a higher proportion of polyploids. Subsequent
wci'kers who analysed the Spitsbergen flora were
however not able to substantiate this point.
Manton (1937> working on a crucifer Biscutella
laevigata which had been found to possess
diploid and tetraploid races in Europe, showed
that the distribution pattern was such that the
tetraploid occupied the ai*eas which had most
recently been freed from ice and that the
diploid remained in the lower altitudes. This
was held to show that the more I'ecently evolved
tetrapoid possessed the ability of more rapidly
colonising new habitats. Other workers have
subsequently found evidence from other plants
which agrees with this interpretation. It may
well be that in the case of T. patei'sonii an
ancestral diploid (or possibly tetraploid' form
migrated aci’oss the then existing Bass Isthmus
and that under glacial conditions in Tasmania
these forms were replaced by higher polyploid
forms and themselves failed to survive. That
the octoploid is not recent in origin is sug-
gested by the regularity of its meiotic division
and the possession of high fertility. It would
seem that the process of “diploidisation” had
gone on with the production of regular meiosis
and the restoration of fertility. The possible
alternative would be that another species of
Thysanotus was able to hybridise with the an-
cestral form and that from a putative sterile
hybrid the present octoploid was developed by
chromosomal doubling. The possible objection
to this latter theory would be that some trace
might be expected of introgression (Anderson
1949) of characters from the other parent and
one would expect a diffex'ence in morphology
between the octoploid and its mainland rela-
tives. This has not so far been observed.
The last example I wish to quote from Thy-
sanotus is the case of the two species T.
triandrus and T. multi fior us. They are both
solely Western Australian in distribution and
both belong to the section of the genus possess-
ing only three stamens. They are similar in
7
habit, they have fibrous, non-tuberous rooted
stocks surrounded by leaf bases both current and
past, the leaves are without petioles and erect.
The scape is simple and has a single terminal
umbel of flowers. The distinguishing charac-
ters of the two species are T. triajidrus — leaves
linear-lanceolate, channelled, covered with a
more or less dense tomentum: T. multi fiorus —
leaves linear, fiat, glabrous. In the latter species
luxuriant specimens may develop a second
sessile or shortly stalked inflorescence some dis-
tance below the terminal one. Experimental
cross-pollination between the species produced
seed and subsequently a mature plant. The
hybrid is somewhat intermediate between the
parents: it has the flat linear leaves of multi-
fiorus which develop a tomentum similar to that
of triandrns. Only a single plant reached
maturity and rather than use the buds for
cytological investigation, experimental self- and
back-cross pollinations were carried out. No
seed was obtained from either of these experi-
mental crossings. From results so far one can
tentatively conclude that in spite of the produc-
tion of a viable hybrid plant there is eitlier in-
sufficient homology in the two chromosome sets
or that breakdown occurs in the development
of endosperm, leading to the failure to produce
viable seed. If this is the true picture it means
that should intercrossing take place in the field
the production of hybrid individuals does not
automatically mean the production of hybrid
swarms and the subsequent possible breakdown
of the species boundaries. Such an intercross-
ing is not very likely under natural conditions
.since the parents are ecologically isolated —
T. triandrus is found on the sandy soils of the
ccastal plain and the inland sand plain, whereas
T. multifl.orus is distributed through the
eucalypt forest on the lateritic soils of the Darl-
ing Range. There are areas in south-west
Western Australia where T. multifioTus tends
to occur on sandy soils but in these areas 7'.
triandrus is absent. If future experimental self-
pollination of hybrids between the two species
results in the production of mature seed and
subsequently plants, one would then be .iustified
m reducing the two parental species to the
status of sub-species of a single species, since
the possibility of their being able to exchange
genes would have been demonstrated.
A second case of polyploidy has been found in
the genus — within T. multi fiorus. here at the
tetraploid level 2n 44. Two localities are
known for plants which on morphological
grounds would be referred to T. multiflorus. but
which occur on the coastal plain associated with
swampy conditions, one near Welshpool and the
other near Pearce. In each locality T. triandrus
occurs in the vicinity. The tetraploid plants
are fertile and possess a regular meiotic divi-
sion. Taking the cytological and morphological
evidence together it could be postulated that
polyploidy here is a case of amphidiploidy —
the doubling of the chromosome number in c
sterile diploid hybrid, possibly similar to the
one between T. trumdrus and T. multiflorus to
which reference has already been made. So far
attempts to obtain seed from experimental
crosses between the tetraploid and its putative
parents have been unsuccessful. Another line
of attack is the possibility of artificially doubl-
ing the chromosome number of the experimen-
tally produced hybrid and then checking the
product against the naturally occurring 2n 44
plant. This would be a similar method to that
used by Mlintzing (1930> who was able to syn-
thesise from the two diploid species of Galeopsis
an amphidiploid which agreed very closely with
the naturally occurring G. tetrahit whose origin
had been hypothetically attributed to a cross
between the ancestors of the two diploid species
used in the experimental cross. Under such
experimental conditions one cannot expect
exact duplication of the existing form because
of the time lapse which occurs between the origi-
nal and the experimental cross which allows of
possible evolution in the parental genotypes.
The cytological situation as it affects the
evolutionary pattern of the Angiosperms in Aus-
tralia has been recently reviewed by Smith
White <1959* who commences by considering
separately the woody and non-woody types of
plants with Au.stralian and extra-Australiaii
distributions. After study of the available
chromosome numbers of hardwood genera of
families such as Myrtaceae, Proteaceae. part of
Rutaceae, Epacridaceae and Casuarinaceae he
finds that within Australia there is marked
diversity in chromosome numbers compared with
extra-Australian members of these groups and
that this diversity is at generic rather than
specific level. At the species level polyploidy
is rare and number variations are unknown
within species.
In softwood or herbaceous types he finds it
difficult to obtain groups of comparable size to
those .just mentioned, his choice is the families
Goodeniaceae and Lobeliaceae and the genera
Danthonia and Nicotiana. Within the two soft-
wood families polyploidy is frequent, reaching
levels of hexaploidy and octoploidy in the
Gccdeniaceae where many cases of intraspccific
polyploid series are found. In the genus
Lobelia different base numbers but no poly-
ploidy are found in the Australian species,
whereas extra-Australian species possess a single
base number and show polyploidy both at intra-
and interspecific levels. T^icotiana shows a
similar situation to that pertaining in Lobelia,
there is chromosome number diversity but little
polyploidy in the Australian compared to the
extra-Australian species. An entirely different
situation holds in the grass genus Danthonia
where, as shown by the data of Brock and
Brown (1961) high polyploids up to decaploid
level with 2n 120 are found in a series of
species which are considered to be primitive as
shewn by the possession of hairs on the lemma
< Table II). Associated with reduction in the
number of hairs there is a corresponding rise in
the proportion of diploid species and reduction
in the maximum level of polyploidy to the
hexaploid level.
The other monocotyledons mentioned by
Smith White are the grass genus Themeda and
the liliaceous genera Sowerbaea and Bland-
ferdia. The two former genera differ markedly
in number of species Themeda having many and
Sciverhaea only two. They are similai' in the
possession of polyploidy, although its geogra-
phical occurrence is reversed in the two genera.
8
TABLE II
Polyploidy and evolutionary series in species oj Danthonia Walter Brock and Brown 1961 f
Lemma hairs
2n = 24
2n :: 48
2n = 72
2n = 96
2n - 120
S
Hnkii
linkii & var. fulva
Imkii var. fulva
induta
mduta
C
Ic nqifolia
longijolia
induta
pallida
6%
A
carvhoides
clelandii
pallida
13%
T
seviiannularia
geniculata
T
cccidentalis
richardsonii
E
31 '7.
SlTo
R
E
D
T
caespitosa
caespitcsa
caespitosa
procera
W
o
setacea
acerosa
setacea
var. l>reviseta
hipartita
purpurascens
7%
alpicola
eriantlia
2k;.
R
auriculata
21%
O
duttoniana
W
monticola
S
50^;
R
laevis
laevis
laevis
E
pilosa
pilosa &
n%
D
var. paleacea
var. paleacea
U
nivlcola
22'o
c
penlcUlata
E
racemosa
D
66%
In Sowerhaea the eastern species is tetraploid
and the western diploid. Themeda australis
according to the data of Hayman (I960) has a
diploid distribution ranging from Tasmania to
southern Queensland. In the same region there
occurs a tetraploid. occasional triploids. penta-
plcids and three localities for a hexaploid. In
the central, western and northern part of the
continent the diploid is absent, the tetraploid
is widely represented and a single locality for
the hexaploid is reported.
The genus Blandfordia extends into Tasmania
and it is reported that this species is tetraploid,
whereas the three species in eastern New South
Wales are diploid. This situation parallels to
seme extent that already reported above for
Thysanotus patersonii and is the reverse of that
reported in Themeda australis where only the
diploid is found in Tasmania.
Smith White (1959) compared the chromo-
some numbers of the eastern and western Aus-
tralian members of the families already men-
tioned under hardwood types above and found
almost exact agreement between them (Table
III), This fact he uses as the basis for a theory
that the Angiosperms arose outside Australia
and then migrated as an already differentiated
body from the north. Let us look at the disti'i-
tautional and variational pattern in Thysanotus
in the light of this theory. A single species is
found in the Philippines, China, Siam, Malaya,
New Guinea and also in tropical Australia. In
New Guinea the pi'esence of T. tuberosus is also
reported. Of the two species with distributions
outside Australia the one with the wider distri-
bution, T. chinensis, is a plant with little or no
adaptive morphology to life in drier conditions,
it has thin mesophyllous leaves and a fibrous
root system without tubers. This habit is shared
with T, triandrus, T. multijlorus, T. asper and
T. glaucus — all of which are western in distri-
bution. T. tuberosus possesses a root system
with tubers and although the leaves are meso-
phyllous they die back and do not have to sur-
vive summer conditions. The other Australian
species possess either a tuberous root system or
an underground rhizome. Of these two types
all those with tuberous roots possess aerial
parts which die back and are replaced annually,
whereas the rhizomatous forms have perennial
above-ground parts which may be either leafy or
somewhat rush-like.
If we accept Smith White’s hypothesis it
would require that T. chinensis would have been
the original form developed outside Australia
and that it was waiting — as he puts it — “at the
bridgeheads” when migration began. It would
seem unlikely that a form with tuberous roots
and an annual cycle of leaf renewal would be
evolved under tropical conditions in New Guinea;
this statement being based on the adaptive value
to dry conditions of the root system and also
on the lack of the colonisation of the Malay
Archipelago by T. tuberosus. On the other
hand if we require that T. tuberosus or some
ancestral form of this species evolved in Aus-
tralia the time during which this could occur
is limited by the necessity for a land connection
for the northward migration to New Guinea.
This theory of the origin of Thysanotus con-
flicts with the view recently expressed by Bur-
bidge (1960, p. 194) that “it would appear that
Arthropodium and Thysanotus may be Austra-
lian elements and that they should be recognised
as such where they occur outside the region.”
If the distribution of the tribe Asphodelae
(to which Thysanotus belongs) is plotted in
terms of the areas occupied by its constituent
genera the following pattern emerges — the two
biggest genera Anthericum and Chlorophytum
have a wide tropical and sub-tropical distribu-
tion, the next largest genera occur in South
Africa and Central Asia, Thysanotus is fifth in
order. If the geographical distribution is looked
9
at in terms of number of genera in a given area
then the Australian continent with a total of
eleven genera has the highest concentration,
these are distributed as six occurring in both
east and west, three in the east only and two
occurring only in the west. South Africa has
only four genera, although one of these is the
large genus Bulbine. Only a few of the genera
within the Asphodelae which are represented
in Australia are also found beyond it; for
example Arthropodium is found in New Zealand
and New Caledonia, Herpolirion in New Zealand.
Chlorophytum generally distributed in the
tropics and Caesia in South Africa. If the
genera with Australian distributions are plotted
on a basis of rough known localities it is found
that the largest number of species occur in the
south-east and south-west of the continent. If
one invokes Willis’ Age and Area hypothesis
(Willis 1949) the evidence from geographical
distribution just presented favours a southern
rather than a northern origin for the group.
TABLE III
Comparison of basic chromosome numbers in
hardwood geiiera of eastern and south-western
Australia (after Smith White 1959
Family and genns Basic numbcr.s
South-western Eastern
Myrtaceae
Actinodhi m
6
—
Darwinia
6. 9
6
Homoranthns
-
6. 9
Verticordia
6. 8. 9, 11
—
Doronieae
Zieria
18
Boron ia
valvatae
16
terminales
9
9
pimiatae
11
11
Erio.'itefno?}
14
14
Phehalium
le
16
Proteaceae
Persoonia
7
7
Grevillea
10
10
Hakea
10
10
Cojiospermuw
11
11
Isopoyon
13
13
Petrophila
?
13
AdenantJiOS
13
Lambertia
14
14
Banksia
14
14
Dryaridra
14
—
Epacridaceae
Styphelia
4
4
Astroloma
4
4, 7
Conostephnnn
8
—
Melicli rifs
—
8
Lcucopogon p.p.
4
4
Lcucopogon p.p.
6
6
Lcucopogon p.p.
11
11
Brachyloma
7?
9
Acrotriche
9
9
Lif-santhe
—
7
Monotoca
—
12
Andersonia
12
—
Sprengelia
-
12
Lysinema
12
—
Evacris
—
13
Woollsia
—
13
Rich ea
—
13
DracophyUum
—
13
Sphenotovia
6, 7
Casuarinaceae
C. glauca group
9
y
C. distyla complex
11
11
If we look at the Australian distribution of
Thysanotus species we find that in common
with such genera as Banksia and Hakea the
genus is predominantly western, twenty-six
species being found in south-west Western Aus-
tralia compared to five in south-eastern Aus-
tralia. The section triandrae with the reduced
number of three stamens is restricted to West-
ern Australia. Of species which occur in both
east and west mention has already been made
of T. patersonii and T. chinensis, in addition
T. tenellus although mainly western is found in
South Australia. There are however examples
of what may on further investigation prove to
be vicarious species pairs as, for example T.
tiiberosus and T. thyrsoideus: T. juncifolius and
T. pseudo junceus.
Evidence for a much wider distribution of
Thysanotus, at least in Western Australia is
provided by the occasional occurrence of very
disjunct populations of the two species T. dicho-
tomus and T. sparteus. Both species are nor-
mally restricted to the Eucalyptus marginata-
E. calophylla forest (the Jarrah forest) of the
Darling Scarp. T, sparteus has a slightly wider
distribution and is also found on the coastal
plain with E. gomphocephala (Tuart) and
Banksia jnenziesii. Isolated occurrences of
these Thysanotus species are found associated
with granitic outcrops in what is now the West-
ern Australian wheat belt. None of these locali-
ties has specimens of E. marginata and curiously
the one locality (Jilakin Rock) with Jarrah
has not so far been found to contain Thysanotus.
Gardner (1944) in “The Vegetation of Western
Australia” makes a special category of plants
associated with the habitat provided in the
vicinity of these outcrops, which provide a
higher level of soil moisture as a result of the
runoff from the rocks.
If we attempt to reconstruct the history of
Thysanotus we could — following Smith White —
postulate an invasion from the north by T.
chinensis which did not go very far south in
eastern Australia but reached further in West-
ern Australia: secondly the evolution of tuberous
rooted forms either simultaneously in east and
west or at a single locality followed by migra-
tion: thirdly migration of a tuberous rooted
form to New Guinea: fourthly the development
of plants with rhizomatous rootstocks and the
lack of formation of root tubers (this again
needs two centres of origin or migration from a
single centre ) : fifthly the development of climb-
ing forms and the migration of these to Tas-
mania- — the absence of other species from Tas-
mania requires either a limitation of southward
migration or the complete destruction of im-
migrants during the period postulated above
for the production of the Tasmanian polyploid
T. patersonii: sixthly the evolution of the three
stamened species in Western Australia followed
by: seventhly further speciation in Western
Australia to account for the larger number of
species in that region. Future work on rela-
tionships of somewhat similar — morphologically
speaking — eastern and western species may help
to decide between the alternatives given in the
second and fourth stages, further work with
10
populations of T. patersonii should help solve
the problems mentioned under the fifth head-
ing.
In summary then we may say that a study
of variation in Thysanotus provides evidence of
variation at single gene mutation level (flower
colour); the occurrence of intraspecific poly-
ploidy (T. patersoTiii) ; the occurrence of sub-
specific variation maintained by partial genetic
barriers and by the pollination mechanism (T.
tuberosus and T. patersonii); the presence of
ecologically isolated species which have diverged
to the extent of being able to produce sterile
hybrid progeny, with which is associated the
possible production of an amphidiploid fT.
triandrus and T. multi florus ) and also the pre-
sence of ecotypes in clinal form (T. triandrus
and T. multifiorus ) .
A postulated pattern of invasion and specia-
tion is suggested based on Smith White’s hypo-
thesis, posing however the question as to the
possibility of a migration at a later date of one
species in the reverse direction. Much work
still remains to be done on this and many other
similar and related problems before the picture
of the evolution of the Australian flora will be
in any way complete.
Acknowledgments
I wish gratefully to acknowledge the receipt
of Research Grants from the University of West-
ern Australia from which the travelling in
connection with this work has been financed.
References
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(Wiley: New York.)
Baker, J. G. (1877).— Revision of the genera and species
of Anthericae and Eriospermae. J. Linn.
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Black, J. M. (1943). — “The Flora of South Australia.”
2nd Ed. (Govt. Printer: Adelaide.)
Bonnier, G. (1895). — Recherches experimental es sur
I’adaptation des plantes au cllmat alpin.
Ann. Sei. Nat. (Bot.) (7me.s.) 20: 217-358.
Brock, R. D. and Brown J. A. M. (1961).— Cytotaxonomy
of Australian Danthonia. Aust. J. Bot. 9:
62-91
Brown. R. (1810). — “Prodromus Florae Novae Hollan-
diae.” (London.)
Burbldge, Nancy T. (I960)— The phytogeography of the
Australian region. Aust. J. Bot. 8; 75-212.
Camp. W. H. and Gilly C. L. (1943).— The structure and
origin of species. Brittonia, N.Y. 4: 232-385.
Clausen. J. (1951). — “Stages in the Evolution of Plant
Species.” (Cornell Univ. Press: New York.)
Clausen, J.. Keck, D. D. and Hiesey. W. M. (1941). —
Experimental taxonomy. Yearb. Carneg.
Instn. 40: 160-170.
Domin, K. (1913). — Additions to the flora of western
and north-western Australia. Prcc. Linn.
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Ewart. A. J. and White, J. (1908).— Contributions to the
flora of Australia — 10. Proc. Roy. Soc. Viet.
(n.s.) 21: 540-548.
Gardner. C. A. (1944).— The vegetation of Western Aus-
tralia. J. Roy. Soc. W. Axist. 28: xi-
Ixxxvii.
Gilmour. J. S. L. and Gregor, J. W. ( 1939).— Demes, a
suggested new terminology. Nature, Lond.
144: 333-334.
Gilmour. J. S. L. and Heslop-Harrison. J. (1954). — The
deme terminology and the units of micro-
evolutionary change. Genetica 27: 147-161.
Hagerup. O. (1932). — Uber polyploidie in Beziehung zu
Klima. okologie und Phylogenie. Hereditas,
Lund 16: 19-40.
Hayman. D. L. (1960). — The distribution and cytology of
the chromosome races of Themeda australis
in southern Australia. Aust. J. Bot. 8: 58-68.
Heslop-Harrison. J. (1953).— “New Concepts in Flower-
ing Plant Taxonomy.” (Heinemann: Lon-
don.)
Hiesey, W. M. (1953). -Comparative growth between and
within climatic races of Achillea under
controlled conditions. Evolution 7: 297-316.
Huxley, J. S. (1940). — Toward the new systematics. in
“The New Systematics.” (Ed. J. S. Huxley.)
(Clarendon Press: Oxford.)
Kunth, O. (1861). — “Enumeratio Plantarum.” 4: 616.
Linnaeus. C. ((1753). — “Species Plantarum.” (Stock-
holm.)
Manton, I. (1937). — The problem of Biscutella laevigata
-ii. Ann. Bot., Lond. (n.s.) 1: 439-462.
Muntzing, A. (1930). — uber Chromosomenvehrmehrung
in Galeopsis Kreuzungen und ihre phylo-
genetische Bedeutung. Hereditas, Lund 14:
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11
2. — The Largest Known Australite and Three Smaller Specimens from
Warralakin, Western Australia
By George Baker*
Manuscript received — J9t/i Dece7?iber. J960
A large oval australite core recently discovered
near Warralakin, Western Australia, is incom-
plete because of artificial fracturing and rela-
Uvely severe natural weathering but it is
nevertheless the largest australite so far brought
to the notice of the scientific world. Even with
the fracture fragments missing, the specimen
weighs 20 grams more than the heaviest austra-
lite recorded to date. It weighs 238 grams; its
weight at the time of landing upon the earth's
surface has been estimated as approximately
280 grams. Reconstruction of the approximate
primary shape of this australite reveals that
about 35 per cent, of its original bulk was lost
by ablation from aerodynamic friction during
passage through the earth's atmosphere at high
velocities.
Three smaller australites subsequently found
in the same general area, six to nine miles south
of the Warralakin-Warrachuppin railway line,
are briefly described.
Introduction
The largest australite so far discovered was
unearthed during post-hole digging operations
which penetrated to a depth of 20 inches in soil,
approximately 16 chains north of the south-
west corner of Block 301, 9.6 miles S.S.E. of
Warralakin Siding, Yilgarn Area, Western Aus-
tralia (Fig. 1). The geographical position of
this locality is 31° 08^ *25" S. and 118° 41' 21" E.
The specimen was found in August, 1957 — pre-
viously to which no other australites had been
observed in this area.
Warralakin is 175 miles E.N.E. of Perth.
Western Australia, and is on the Wyalkatchem-
Southern Cross railway loop-line.
The specimen was recovered in a slightly chip-
ped and weathered condition by Messrs. D. S.
and A. V. Poole on the property “Coppin Rock’'
near Warralakin. The slight chipping resulted
from damage by the post-hole digger. Screen-
ing of the soil from the post-hole failed to re-
cover the detached fragments. Unfortunately a
large piece was subsequently fractured from the
end of the specimen with a cold chisel and lost.
Even without this fragment, however, the speci-
men constitutes the largest australite known
to science. In the unbroken condition it would
have been approximately 50 grams heavier than
the previously recorded (Fenner 1955 » heaviest
australite.
This australite is lodged in the geological
collection (Reg. No. 8925) of the School of
Mines of Western Australia, Kalgoorlie, to
* C.S.I.R.O., Mineragraphic Investigations, c/o Geo-
logy Department, University of Melbourne. Parkville,
N.2. Victoria.
which institution it was donated by Mr. R. W.
Poole (of Gold Mines of Kalgoorlie), a brother
of the discoverers.
Fig. 1. — Locality map of the area south of the
Warralakin-Warrachuppin railway line. Yilgarn Area.
Western Australia, showing sites (full circles) of recently
discovered australites. The large figvires are the
registered numbers of the australites in the geological
collection of the School of Mines of Western Australia.
Kalgoorlie.
The precise depth and orientation of the
specimen in the soil could not be noted at the
time of discovery, but detailed inspection of all
surfaces and of the clay lodged in pits on them
leads to the conclusion that the posterior sur-
face lay downwards in the soil. The clay on the
posterior surface was redder in colour due to
more ironstaining, whilst that on the anterior
surface was biscuit-coloured to almost white due
to leaching, and the redder coloured clay on the
posterior surface was rather more firmly
cemented into pits and narrow flow lines. In
12
the vicinity of the find, the area is relatively
well- timbered, the surface is sandy with a few
pebbles brought up by the roots of fallen trees.
The soil in which the specimen was found is
of the “sand plain” type. In Western Australia,
this is often a leached “A’' horizon but, irres-
pective of the initial origin of the sand, it has
usually been wind-drifted to a greater or lesser
extent. It can be quite confidently stated that
this large australite occurred at shallow depth
in wind-blown sandy soil.
The deduced position of rest of the austra-
lite, i.e., with its anterior surface upwards, is
the reverse to its position of aerodynamically
stable orientation during earthward flight.
Nevertheless this is the stable position of rest
on the earth’s surface, just as for the majority
of australites exposed on the surface for which
this position was observed on discovery. The
stable position of rest was either attained im-
mediately after striking the ground, or resulted
subsequently when specimens were moved by
transporting agents after landing.
PLATE I
The largest known australite (Reg, 8925, geological collection. School of Mines of Western Australia, Kalgoorlie).
From south-west corner of Block 301. 9.6 miles S.S.E. of Warralakln Siding, Western Australia.
Pigs A to D showing etched ‘bruise-marks", a few of which resemble “hofchen" and "tischschen". on an oval
australite core A — end-on view across shortest diameter; posterior surface uppermost: rim poorly preserved
(XI 16) B = side view across largest diameter; posterior surface uppermost: equatorial zone at right-hand end
' relatively well preserved (Xl-10). C = anterior surface (XI. 10). D posterior surface (Xl.lO).
(Photographs by K. L. Williams.)
13
Description of Specimen
As submitted for examination, the weathered
specimen revealed a small quantity of light
buff-coloured to reddish-brown lateritic sandy
soil partially cemented and partly jammed into
shallow^ pits and lunate to circular shallow
grooves on all surfaces, except the newly ex-
posed (artificially fractm*ed) surface. It was
necessary to remove all secondary material prior
to determining the specific gravity of the austra-
lite.
The weathered surface has a dull lustre and
reveals occasional poorly pronounced “hofchen”
and “tischschen” structures brought out by dif-
ferential solution-etching by soil solutions.
Rare, fine flow lines occur on parts of the sur-
face. while only a few straight, slightly deeper
gi coves with parallel walls are present in one
place. Many of the markings on the posterior
and anterior surfaces of the specimen resemble
chatter-marks brought about by collisional
bruising, but since the australite was found
in a milieu where natural agencies likely to
have caused these features are apparently want-
ing. the possibility arises that the marks may
have been due to aboriginal activities (e.g. use
as a pounding stone, etc.), although this is
difficult to prove.
The broken surface reveals the highly
vitreous lustre and conchoidal fracture with
subsidiary ripple fracture pattern that is so
characteristic of freshly fractured tektite glass.
Up to three dozen minute internal bubbles can
be detected on the fractured surface with the
aid of a 10 X hand lens. The area of the frac-
tured surface is approximately 12.5 cm'-, and if
the internal bubbles are maintained in this dis-
tribution throughout the interior of the large
australite. they could partly account for the
specific gravity of the tektite glass as a whole
being lower than usual for australites from the
western portions of the Australian tektite
strewnfield. These bubbles are more or less
spherical in shape, and range from 0.25 mm
to nearly 0.75 mm in diameter.
A rim (cf. Baker 1959, p. 39) is just discern-
ible around most of the periphery of the speci-
men <see Plate I, Figs. A and B), and is suffi-
ciently pronounced to aid in discriminating
between the anterior and posterior surfaces of
the australite
The tektite glass is jet black and opaque in
reflected light for the specimen in bulk, but
yellowish bottle-green and translucent in trans-
mitted light on the thinner edges of the frac-
tured surface and in small splinters detached
for refractive index determinations.
Dimensions, AVeight and Specific Gravity
The specimen is 42 mm in depth < thickness)
and 62.5 mm in width as determined from the
non-fractured portions of the australite. Its
present length is 65 mm, but the original
length (on reconstruction of the fractured
form) was approximately 70 to 72 mm. It is
thus an oval australite core. It weighs
238.00 grams and has a specific gravity
value of 2.409 as determined in distilled water
(Th^o 12.8° C.) on a Mettler K-type balance.
The specific gravity is approximately that of
the mean specific gravity value determined for
1,086 specimens cf australites (Baker and
Forster 1943. p. 403), but is significantly lower
than the general run of specific gravity values
for australites from the western half of Austra-
lia, evidently because of its content of small
bubbles.
The volume of the cleaned specimen is 98.8
cm\ Reconstruction of the unbroken (but
weathered) form reveals that approximately one
ninth was removed by artificial fracturing, so
that the specimen as found would have weighed
about 265 grams. It has not been possible to
reconstruct the specimen accurately enough to
ascertain its size as it would have been on first
landing upon the earth’s surface from an extra-
terrestrial source, but an approximate estimate
of the amount weathered from this large oval
australite core indicates that in the perfectly
preserved state, the original weight would have
been in the vicinity of 280 grams.
Australites weighing over 200 grams are ex-
tremely rare, and only two others are known.
One. weighing 218 grams, was found at Lake
Yealering. Western Australia (Fenner 1955), the
geographical position of which is approximately
32° S. and 118° E., the other came from
Karoonda, South Australia (Fenner 1955), at
35° S. and 140° E. and it weighs 208.9 grams.
The Lake Yealering specimen is lodged in the
collection of the Western Australian Museum in
Perth, Western Australia, and the Karoonda
specimen is in the South Australian Museum
collection in Adelaide. South Australia.
Only eleven specimens are known that weigli
over 100 grams and under 200 grams. Seven
of these are recorded by Fenner (1955, pp. 90-
91); the other four are: — a boat-shaped form
of 141.63 grams weight from Port Campbell.
Victoria (Reg. No, 11402, National Museum of
Victoria collection), a round core of 135.16 grams
weight from Gymbower near Goroke. Western
District of Victoria (National Museum of Vic-
toria collection), a round core of 111.25 grams
weight from Lake Wallace near Edenhope, Vic-
toria (Reg. No. E1986, National Museum of Vic-
toria collection), and a boat-shaped form of
107.46 grams weight from near Narembeen.
Western Australia (Reg. No. 8950, geological
collection, School of Mines of Western Austra-
lia, Kalgoorlie).
Refractive Index and Estimated Silica Content
The refractive index of the glass varies ac-
cording to the chemical composition of different
internal schlieren. but does not show a wide
range in the small fragments examined. Values
obtained by the Immersion Method using mono-
chromatic (Na) light and employing micro-
scopic fragments removed from the freshly
fractured surface of the specimen by light pres-
sure flaking, showed a range from: —
n^j, 1.504 to 1.506
The specific refractivity (k) of the glass,
determined from the relationship k n — l^d
(where n the refractive index, and d- the
specific gravity of the specimen), ranges from
0.2092 to 0.2100, according to the composition
of the different schlieren.
14
From the Silica-Specific Gravity and the
Silica-Refractive Index graphs for tektites
generally (see Barnes 1940; Baker 1959', the
silica content of the specimen is estimated to
be approximately 74 per cent. In view of the
possibility that the content of small internal
bubbles lowers the specific gravity of this speci-
men, however, both the silica content and the
specific refractivity may be slightly different
from the estimated values, but the silica con-
tent is likely to be no more than one or two
per cent, lower.
Curvature of Surfaces
Silhouette traces of the weathered specimen
reveal that arcs of curvature across the shortest
diameter of the australite are:
Rh 34.5 mm. and Rv 31.5 mm,
where Rn the radius of curvature of the pos-
terior (back) surface and Rv the radius of
curvature of the anterior (front) surface. Across
the longer diameter, the radii of curvature arc
Rij -40.9 mm and Rk ^ 44.1 mm, but con-
structed circles do not fit as accurately to the
arcs of curvature in this direction as to the
surface curvature across the shorter diameter.
The original form, prior to modification by
(i) ablation arising from aerodynamic friction
during atmospheric flight, (ii) weathering by
subaerial agents, and (iii) artificial fracturing
after the specimen was collected, is estimated
to have been an ellipsoid of revolution measur-
ing approximately 7 cms X 7 cms X 8 cms in
size. The depth of ablation in the stagnation
point region (i.e. at the front pole of the speci-
men as aligned in aerodynamically stable
orientation) was approximately 2.4 cms, and the
overall amount ablated from the surface pro-
jected in the line of flight U.e. the anterior
surface) totalled about 60 cm'* in volume. Some
35 per cent, of the original form was thus lost
during transit at ultrasupersonic speeds through
the earth’s atmosphere, by the processes arising
from aerodynamic friction. This is a relatively
low percentage loss compared with many
.smaller australites for which the range _ in
amount of glass removed by ablation and fusion
stripping plus flange shedding is from 32 per
cent, to 98 per cent., with an average loss <for
65 specimens) of 50.5 per cent. (cf. Baker
1961a, Tables 4, 9 and ID. It is also much
lower than the percentage loss range <61 to 96
per cent.) by ablation of ten larger, well-pre-
served australite cores from Port Campbell,
Victoria (cf. Baker 1961a, Table 13).
Smaller Australites
Of the three smaller australites subsequently
found some three and a half miles west and
four and a half miles north-west of the site
of the largest australite. two are boat-shaped
and one oval-shaped. They anci the largest
australite were recovered from within an area
of approximately nine square miles (Fig. 1).
Boat-shaved Forvi
The largest of the three smaller forms was
found late in 1958 by Mr. D. S. Poole on Block
309 on H. M. Poole’s property “Devon.” This
is 7.2 miles east of south from Warralakin Sid-
ing, or 3D 06' 36" S, and 118*^ 38' 34" E. Here
the surface consists of residual gravel resulting
from partial exposure of the laterite horizon.
The specimen was presented by Messrs. D. S.
and R. H. Poole to the School of Mines of West-
ern Australia, Kalgoorlie (Reg. No. 9021). Its
dimensions are 38.5 mm long, 27.5 mm wdde, and
16.5 mm deep ( thick). It weighs 22.227
gmms, and its specific gravity as determined in
distilled water (Tii^o - IS*^ C.) on a chemical
balance is 2.425.
The specimen reveals a fairly well-defined
rim separating the pitted and flow-lined pos-
terior surface from the finely etch-pitted
anterior surface. All structures on the austra-
lite, however, have been considerably modified
by solution-etching, although the specimen is
generally rather better preserved than the large
australite described above. The boat-shaped
outline of the specimen from front and back
aspects is somew^hat irregular (Plate II, Figs.
A and B) due to erosion.
Portions of the flow^-swdrled central region
of the posterior surface (Plate II, Fig. B> have
been partly accentuated but some areas have
been partly destroyed by solution-etching. Its
general characteristics, however, are similar to
the primary flow' swirls on the posterior surfaces
of better preserved australites.
In side- and end-on aspects, the outline is
approximately that of a biconvex lens wdiich
has one diameter greater than the diameter at
right angles. The radii of curvature across
the shorter diameter ( wddth) are Rh 16.8
mm and Rr 17.2 mm. wdth the arcs of curva-
ture fitting those of constructed circles with
these radii. The radii of curvature along the
larger diameter ( length) are approximately
Rh 32.9 mm. and Rt- 25.9 mm, but these
two arcs of curvature do not conform accurately
to the arcs of curvature of constructed circles
with these radii, being rather flatter in the polar
regions and steeper towards the rim of the
.specimen.
Oval Core
An eroded oval australite core was found by
Mr. A. T. Miles with its anterior surface upw^ards
on 9th October, 1960. It was exposed on the
gravelly surface of an area that slopes gently
northw'ard near the site of Reg. No. 9021 (Fig.
1). The specimen is now- in the collection of the
School of Mines of Western Australia, Kalgoor-
lie (Reg. No. 9050).
It weighs 7,102 grams and its specific gravity
is 2.431 (T(i,o 15.5® C.). The longer dia-
meter measures 22 mm, the shorter 18 mm and
the depth ( thickness) is 14 mm.
All surfaces reveal the effects of solution -
etching and the sculpture pattern consists
principally of small pits 0.2 mm across, rang-
ing up to a few’ larger pits 3.0 mm across, mostly
on the posterior surface. The larger nits reveal
a few smaller pits and etched out flow lines on
their walls. Where the larger pits are crow’ded
together, sharp ridges separate one pit from its
neighbour, and these parts of the surface re-
semble that of hammered metal.
15
V
PLATE II
A boat-shaped australite found 4^ miles south-east of the large oval australite core shown in Plate I (Reg.
No. 9021, geological collection, School of Mines of Western Australia, Kalgoorlie).
A = anterior surface showing fine etch pitting ( X 2.57). B = posterior surface with weathered and etched flow
swirl surrounded hy bubble pits modified by etching (X2.57).
(Photographs by K. L. Williams.)
16
A flaked equatorial zone is detectible but
rather poorly preserved. It shows a few com-
plexly contorted flow lines that have been ex-
posed and accentuated by solution-etching.
These are the surface expressions of an internal
schlieren structure brought into prominence on
the level to which weathering has advanced.
Flaking around the equatorial zone has resulted
in the specimen appearing like a conical core
in side aspect.
Smaller Boat-shaped Form
Like all the other specimens from this area,
the smaller boat-shaped form reveals no evi-
dence that would indicate the former existence
of a circumferential flange. If ever present, the
flange has been completely removed, either by
shedding during flight, or subsequently by sub-
aerial erosion, and erosion has further modified
the equatorial region and the posterior and an-
terior surfaces of the specimen.
It was found with the anterior surface facing
upwards by Mr. D. S. Poole on 10th October.
1960, on the roadside between Block 303 and
Block 304, approximately 22 chains from the
south-west corner of Block 303 (Fig. l.>. Its
registered number is 9051 in the collection of
the School of Mines of Western Australia, Kal-
gcorlie, and it was discovered on top of the
ground in gravel country similar to that on
vrhich Reg. Nos. 9021 and 9050 were found. The
geographical location of the specimen is appro-
ximately 31“ 08' 21" S. and 118° ZT 58" E.
Its weight is 5.9115 grams and its specific
gravity is 2.433 (Tji,o 15.5° C.>. The measure-
ments of the specimen are 26 mm long, 17 mm
wide and 10 mm deep.
An cld fracture surface at one end of the
specimen is as prominently sculptured by
weathering (mainly solution-etching t as are the
posterior and anterior surfaces.
The rim separating the posterior from the
anterior surface is detectible in places around
the periphery of the specimen. Both surfaces
reveal flow lines which represent the “outcrops”
of an internal schlieren pattern that trends
generally parallel with the cutline of the form.
Some of the smaller pits on both surfaces are
evidently etch pits resulting from w’eathering.
but several deeper rounded pits 1 mm across
and elongated pits 2X1 mm in size are either
overdeepened, oi'iginally superficial bubble pits
or else internal bubbles exposed during the pro-
cess of weathering.
Summary and Conclusions
The four australites recovered from the area
south of the Warralakin-Warrachuppin railway
line in Western Australia have all been modi-
fied by subaerial erosion to such an extent that
their volumes on first landing on the earth’s
surface cannot be accurately determined. Their
sculpture patterns are largely a result of
weathering, more particularly by the process of
solution-etching in sandy soils.
In borrow pits and the banks of dams in the
district, the soil profile shows a variable thick-
ness of sandy soil passing downward into a zone
of discrete laterite nodules, and thence, in a
depth of a foot or so. into compact laterite. The
variable thickness of the sandy soil is partly
attributable to wind, and. in some places, partly
to rainwash. The largest specimen (Reg. No.
8925 1 was found w'here the w'ind-drifted sand is
thicker than usual, and at this Iccality the 20"
deep postholes barely entered the nodular
laterite zone. This specimen cculd w’ell have
been exposed at one time and reburied under
drifted sand. By contrast, two of the smaller
specimens (Reg Nos. 9021 and 9050* that were
found within four chains of each other, occurred
right at the surface (gibber-plain type) in an
area where partial stripping of the finer soil
constituents had exposed nodules of laterite,
among which the two australites were exposed
to view. The fourth specimen (Reg. No. 9051)
occurred on a gravelly, similarly stripped area.
On gravelly slopes in this area, more thorough
soil stripping down to the duricrust surface
cculd readily result in removal of both lateritic
nodules and australites and their transporta-
tion to fiat-lying areas where re-burial in sandy
soil is not improbable.
There is little doubt that these australites are
post-laterite in age and their recovery from the
overlying soils points to their recent age on
earth (cf. Baker 1961b >. Their surface sculp-
ture is largely a result of weathering, whereby
subaerial agents have reduced the sizes of the
original specimens and developed structures that
are a manifestation of their usually complex in-
ternal streakiness.
Acknowledgments
The author is indebted to W. H. Cleverly, B.A.,
B.Sc., head of the Geology Department, School
of Mines of Western Australia, Kalgoorlie, for
kindly submitting the australites for examina-
tion, and for obtaining details of their mode of
occurrence and precise localities of discovery.
I am also grateful to Professor Rex T. Prider
for communicating the paper through his mem-
bership of the Royal Society of Western Aus-
tralia.
References
Baker, G. (1959). — Tektites. Mem. Nat. Mus. Viet. 23;
5-313.
Baker. G. (1961a). — Volumenbeziehungen von wohler-
haltenen Australit-Knopfen. -Linsen unci
-Kernen zu ihren prlmaren Formen. Cliem.
d. Erde. 21 (3) (in press).
Baker, G. (1961b). — Australite von Wingellina. West-
Australien. Cliem. d. Erde. 21: 118-130.
Baker, G.. and Forster, H. C. (1943). — The specific
gravity relationships of australites. Amer.
J. Sci. 241: 377-406.
Barnes, V. E. (1940). — North American tektites. Univ.
Tex. Publ. No. 3945; 477-582.
Fenner, C. (1955). — Australites Part VI. Some notes on
unusually large australites. Tra7is. Roy. Soc.
S. Aust. 78: 88-91.
17
3.— The Flora of Granite Rocks of the Porongurup Range, South Western
Australia
By G. G. Smith*
Manuscript received — 18th April, 1061
The occurrence of the vascular and non-
vascular flora on granite outcrops of the
Poronerurup Range in south Western Australia
is described in relation to the environment. A
systematic list of the algal, lichen, bryophyte.
pteridophyte an angiosperm species of this
flora is given.
Introduction
The Poiongurup Range, t twelve miles east
of the town of Mount Barker, consists of
granitic rocks rising from the surrounding plain
to a maximum altitude of 2,200 feet at the peak
known as the Devil’s Slide. Other peaks in the
range include Castle Rock (altitude 1,870 feeti,
Twin Peak East (2,080 feet). Twin Peak West
(2,040 feet). Gibraltar Rock (2,100 feet). Nancy’s
Peak (2,140 feet), Hayward Park Peak (2.000
feet), Angwin Park Peak (1,780 feet) and Collier
Park Peak (2,080 feet).
The vegetation type of the range is an outlier
of the Karri forest to the west. The trees in-
clude Karri, Eucalyptus diversicolor, the Marri.
E. calophylla and to a less extent, E. cornuta
and E. megacar pa. The under-storey of this
forest includes the tall shrubs, Tryinalium
spathulatum. Acacia pentadenia, Albizzia dis-
tachya, Oxylohium lanceolatum, Mirbelia dila-
tata and numerous species of smaller sclero-
phyllous shrubs.
This paper describes the flora of granite out-
crops in this forest. Collections and observa-
tions were made at Castle Rock, Nancy's Peak,
Devil’s Slide, Gibraltar Rock and at several
outcrops towards the base of the range.
One large slope about 80 feet above the Bol-
ganup Dam (988 feet) has been called by the
author, Rain Gauge Rock, after the automatic
rain gauge set up on it since 1957 by the
Hydraulics Branch of the Country Water Supply
Department in connection with the Bolganup
Dam.
Climatic Data
Rainfall
The existence of an automatic rain gauge at
Rain Gauge Rock provides some evidence of
the amount and monthly distribution of pre-
cipitation on the site of some of the lithophyl-
lous communities described below. Table I
Department of Botany. University of Western Austra-
lia. Nedlands. Western Australia.
t Althorigh commonly written Porongorup, the name
Porongurup was given by Surveyor-General J. S.
Roe in 1835, and is followed by the Geographic
Nomenclature Committee.
shows the annual rainfall for three consecutive
years as taken from August, 1957, when the
gauge was set up.
TABLE I
Annual rainfall for three consecutive years at Rain
Gauge Rock (Bolganup Dam) Porongurup Range, West’
ern Aiistralia
Rainfall
Year
In Inches
August, 1957-1958
20.16
August. 1958-1959 . .. ..
29.91
August. 1959-1960
52.03
Average Annual Rainfall
34.03
The average yearly rainfall for the nearby
town of Mount Barker is 30.23 inches and that
of Albany to the south of the Porongurups is
39.67 inches. A study of rainfall isohyets for
this region indicates that the normal annual
rainfall at the Porongurups could be about 34
inches, as both Mount Barker and the Poron-
gurups are between the 30 inch and 40 inch
isohyets, but much closer to the former.
An analysis of the monthly rainfall at Rain
Gauge Rock into total winter and summer pre-
cipitation gives some idea of the seasonal
desiccation to which the lithophyllous communi-
ties are subjected. In Table II the winter rain-
fall is taken as a six month period from May.
when the winter rains start, to the end of Oc-
tober. and the summer rainfall as the precipita-
tion from November to the end of April. The
table shows that the lithophyllous communities
are subjected to a wet half and a relatively dry
half of the year. This evidence of summer
desiccation supports the general climatic picture
in the south-west of Western Australia where
shallow-rooted plant communities are subjected
to drought from about November to April.
TABLE II
Wmter and summer rainfall at Rain Gauge Rook
(Bolganup Dam), Porongurup Range, Western Australia
Winter rainfall in inches. Summer rainfall in inches.
May Ist-Oct. 31st Nov. Ist-April 30th
At Rain Gauge Rock
22.59 (1958) 3.75 (1957-58)
23.56 (1959) 8.78 (1958-59)
27.05 (I960) 16.62 (1959-60)
Mt. Barker (Average)
21.75 8.48
Temperature
The yearly average of daily maximum tem-
perature for Mount Barker is 67.7° F, the
monthly average of daily maximum tempera-
ture being greatest in February at 78.3'’ F. At
Mount Barker the average of daily minimum
18
temperature is 48.3*" F. the monthly average of
minimum temperature being lowest in July at
42.0^ F. These figures give some indication of
the temperature extremes likely to obtain at the
Porongurups, where unfortunately there is no
recording station. The climatic data for Albany
and Mount Barker were taken from Climatic
Averages of Australia — Bureau cf Meteorology.
Melbourne, 1956.
Flora on the Granite Outcrops
The lithophyllous flora of granite rocks in
south Western Australia is composed chiefly of
bryophytes, lichens and sub-aerial algae. Small
herbaceous vascular plants are also common in
moss swards on granite slopes, while larger,
perennial vascular plants are limited to the
humus-filled crevices and depressions, or to the
shallow soil at the border of outcrops.
The bryophytes and lichens of south Western
Australia are reasonably well-known systemati-
cally from early collections made by visiting
botanists, but there are few published records
of these plants in relation to their habitats.
Diels <19061. in his account of the vegetation of
south Western Australia, did not describe the
cryptogamic flora of rock outcrops. Likewise.
Gardner (1942i described only the vascular flora
in very general terms. Willis (19531. in his
descriptions and analysis of the land flora of
the Archipe'ago of the Recherche gave con-
siderable ecological information on the litho-
phyllous flora of the igneous rocks of these
islands. This flora shows some affinities with
the lithophytes of the Porongurup Range, such
as the wide occurrence in the Recherche of the
mosses Campylopus bicolor and Bryum billar-
dieri, and of the lichens, Parmelia rutidoia and
P. conspersa. The two floras differ strikingly,
however, in the paucity of Cl(tdo7iia species at
the Recherche, whereas four species of this
genus are common components of moss swards
at the Poi'ongurup Range. On the other hand
the cushion plant, Borya nitida is common on
granite slopes of the Recherche as it is on
granite rocks of the Darling Range near Perth,
but is conspicuous by its absence in the Poron-
gurup Range.
Crustose Algal and Lichen Communities
The surfaces of granite tors and other steeply
sloping outcrops which, to the casual observer,
are bare rock surfaces, bear extensive communi-
ties of minute crustaceous lichens and blue
green algae (Plate II, Pig. 2).
This type of community consists of the crus-
tose lichen. Caloplaca aurantiaca and at least
five other species of crustose lichens which to
date it has not been possible to determine. The
sub-aerial algae, Calothrix sp. and Chroococcus
sp. occur in black, stain-like stands, mainly
along seepage depressions over the outcrops.
The more conspicuous lichens, Parmelia
rutidota, P. imitatrix, P. conspersa, P. perlata
and Sticta crocata, also occur on steep rock
faces whether exposed to full sunlight oi pai-
tially shaded by the canopy of the Karri forest
(Plate I, Fig, D. Sticta appears to reach
maximum vegetative development on gentle
.semi-shaded slopes where water seepage is high.
The moss. Grimmia trichophylla. is the only
bryophyte found on the summits of granite tors
and domes where it grows in small cushions in
depressions and crevices resulting from weather-
ing of the rock.
Moss Sward CommuiiUics
Low angled granite slopes throughout the
Porongurup Range bear extensive moss swards
which form mosaics with encrusting algal com-
munities or crustose lichen communities (Plate
I. Fig. 1 and Plate II. Fig. 1>. Rarely is much
of the rock surface devoid of plant life. Where
recent exfoliation of the rock has occurred this
is so, of course.
The association of species in these moss
swards is variable but from examination and
sampling of numerous slopes up to an angle of
approximately 45^ and of varying exposures to
all points of the compass, but not shaded by
trees or shrubs, the following species assessment
may be given.
Two mosses. Campylopus bicolor and Breutelia
affiJiis are the most extensive sward components.
Other mosses in this sward include Semato-
phyllum hoviomallum, Rhacocarpus humboldtii,
Brytan argenteum, B. billardieri, Tortella
calycina, Hedwigia imberbis. The sward rarely
exceeds three inches in depth where it overlies
fairly smooth granite slopes and its surface is
remarkably even considering the several moss
components.
The high water-holding capacity of the mosses
enables several herbs and geophytes to colonise
the sward. The chief vascular plants are: —
Opliioglossum coriaceum
Triglochin centrocarpa
Aira caryophyllea
Brizula muelleri
Centrolepis glabra
Pritzelia pygmaea
Bulbine semibarbata
Chamaescilla ecry7ubosa
Tribonanthes variabilis
Hypoxis glabella
H. occidenialis
Pterostylis nana
Calandrinia pygmaea
C. calyptrata
Spcrgularia rubra
Drosera glanduligera
Crassula pcdicellosa
C. 7nacraniha
C. sieberia7ia
Trifoliu7n dubhon
Jiydrocotyle callicarpa
II. diantha
H. blepharocarpa
Trachymenc pilosa
T. anisocarpa
Ho7nalosciadiu7ti verticillatiun
A7iagallis fe7nina
Mit7'asac7Tie paradoxa
Erythraea australis
Bartsia latifolia
Styliduwi calcarahim
S. corymbosu7n var. prolijeruvi
Vero7iica calyciinis
Polypompholyx te7ieUa
GalUi77i murale
Levenhookia dubia
Rutidosis muUiJlorus
Helipteruin cotula
A7igia7ithus te7iellus
A. hmnifusus
Quinetia urvillei
Cenia turbmata
Hypoclioeris glabra
Podolepis lesso7iii
PLATE I
Fig. 1 .— Moss sward over granite with Trackymene ariisocarpa in flower and Pe/ar.oo?itiAm druTTunondii Granite
surface in the background with Parmelia rutidota. The shrubbery in the right background consists of
Thryptomene saxicola and Stypandra grandifiora.
Fig. 2.— Granite boulder with Veronica calycina in shaded recess. Right foreground with moss sward and
Villarsia calthifolia in flower.
20
PLATE II
wirr 1 —Mnq-? «?ward on eranite slopes with Pelargonium drummondii, C}ieilanthes tenuifolia in centre and back-
eroimd Thrvvtomene saxicola and Stypandra grandifiora shrubbery towards edge of slope. Eticalyptus diversicolor
gruunu, ifLiypt, calophvlla in the background.
Pig. 2. — Moss sward on granite slopes.
amongst boulders, ilcacta sp.»
Slopes and boulders with crustose lichens. Pelargonium drummondii
Lepidosperma cUusum and Thryptomene saxicola in background.
21
The lichen component of the sward includes
the foliose forms, Cladonia furcata. C. aggregata,
C. chlorophaea, C. retipora, Siphula caesia,
Sticta crocata and Collema sp. These lichens
grow embedded in the moss turf, but Siphula,
also grows in pure stands in exposed humus
of shallow rock crevices. The only liverwort
collected from the sward was Fimbriaria cono-
cephala.
A few species of perennial vascular plants
occur on granite slopes where either a crevice
or the moss sward over depressions provides
sufficient depth of humus for their root systems
(Plate I, Fig. 1 and Plate II, Figs. 1 and 2>.
The fern, Cheilanthes tenuifolia occurs in ex-
tensive stands in the deeper parts of the moss
swards. Stypandra grandifiora. Pelargonium
drummondii, Carpobroius aequilaterus. Gera-
nium pilosum. Lepidosper7ua effusum, Thrypto-
viene saxicola. Danoinia citriodora and Agonis
juniperina are conspicuous perennials of the
shrubbery of these crevices and depressions.
CommuniUes of Shaded Crevices
Along the ridge of the Porongurup Range the
exposed granite core of the range forms a mas-
sive system of tors with numerous deep crevasses
and shallow caverns or recesses resulting from
the sphaeroidal weathering of the granite. In
these dissections, leaf litter and the products
of exfoliation accumulate to form a shallow
loam. Wherever this loam is sufficiently shaded
by the ad.iacent rock a distinctive microflora
develops (Plate I. Fig. 2).
The principal bryophyte and lichen elements
of this microflora are Lophocolea heterophyl-
loides and Lepidozia parvistipa. both of which
form extensive mats over either loam or granite
substrates. Frullania latogaleata is common in
drainage depressions of the overhanging rock.
Sematophylhini lionwmalluvi forms extensive
mats. Sticta^ cracaXa is most common in shaded
exfoliation depressions. Marchantia cephalo-
scypha is a rare but striking liverwort on humus
of shaded undercuts.
Commonly associated with these bryophytes
and lichens in the recesses of rock dissections
are the ferns, Asple7iium fiahelUfoliu7n, Adian-
tum aethiopicuin and A7iogra7nma leptophylla,
the latter invariably accompanied by large
numbers of its long-lived prothalli. Asplenium
prae7norsu77i occurs in more exposed crevices,
although this fern is not common in the Poron-
gurup Range.
The most common angiosperms in this habitat
are Coryhas dilatatus, Cryptostylis ovata, Poa
caespitosa. Pelargonium drmrmondii, Oxalis
corniculata, Hydrocotyle hirta, Villarsia calthi-
Jolia (a handsome species endemic to the
Porongurup Range) and Veronica calycina
(Plate I. Fig. 2). In rock dissections of more
open aspect the following shrubs are common; —
Thrypto77iene saxicola, Sollya fusiformis, Sty-
pandra grandifiora and Sola7ium nigrum.
Systematic List of the Flora on Granite Out-
crops of the Porongurup Range
Specimens of the following species are pre-
served in the Herbarium of the Botany Depart-
ment of the University of Western Australia.
Algae
Chroococcaceae
Cfiroococcus sp.
Rivulariaceae
Calothrix sp.
Bryophyta-IIopaticae
Lepidoziaceae
Lepidozia parvistipa Tayl.
Harpaiithaceae
Lophocolea hcterophylloides Nees
Fnillaniaceae
Frullania latogaleata Herz.
Marchantiaceae
Marchantia cephaloscypha Steph.
Operculatae
Fimhriaria conocephala Steph.
Bryoph.vta-Musci
Grimmiaceae
Grirnmia trichophylla Grev.
Dicranaceae
Campy fopus hicolor (Hornsch.) Hook.f.
Pottiaceae
Tortella calycina (Schwgr.) Dixon
Bryaceae
Bryum argemteum Hedw.
B. hillardieri Schwgr.
Biirtramiaceae
Breutelia affi7iis (Hook.) Mitt.
Hedwlgiaceae
Hedwigia imberbis (Sm.) Spreng.
Rhacocarp7is hum.boldtii (Hook.) Lindb.
Sematophyllaceae
Sematophyllmn homomallum (Hpe.) Broth.
Licheiies
Collemaceae
Collema sp.
Stictaccae
Sticta crocata (L.) Ach.
CUidonlaceae
Cladoiiia furcata (Huds.) Schrad.
C. angregata (Sw.) Eschw.
C. chlorophaea (Flk.) Spreng.
C. retipora (Labill.) Fries
Parmellaceae
Parmelia rutidota Hook.f. and Tayl.
P. iniitairix Tayl.
P. conspersa Ach.
P. perlata (L.) Ach.
Usneaccae
Siphula caesia Mnell. Arg.
Caloplacaceae
Caloplaca aurantiaca (Lightf.) Fries
Pteridophyta
Ophioglossaceae
Ophioglossuvi eoriaceum A. Cunn.
ci c 0 Et ©
Clieila7itfies tenuifolia (Burm.f.) Swartz
Anogramma leptophylla (L.) Link
Adiantum aethiopicum L.
Aspleniaceae
Asplenium ftabellifolium Cav.
A. praemorsum Swartz
Angiospermae
Scheuchzeriaceae
Triglochin centrocarpa Hook.
Gramineae
Aira caryophyllea L.
Poa caespitosa Forst.
Cyperaceae
Lepidosperma effusinn Benth.
Centrolepidaceae
Brizula muelleri Hieron.
Centrolepis glabra (F. Muell) Hieron.
Philydraceae
Pritzelia pygmaea (R. Br.) F. Muell.
Lillaccae
Bulbine semibarbata (R. Br.) F. Muell.
Ch-ainaescilla corymbosa (R. Br.) F. Muell
Siyjmndra grandifiora Lindl.
22
Amaryllidaceae
Trihonanthes variabilis Lindl.
Hypoxis glabella R. Br.
H. occidentalis Benth.
Orchidaceae
Corybas dilatatus Rupp and Nicholls
Pterostylis nana R. Br.
Cryptostylis ovata R. Br.
Alzoaceae
Carpobrotus aequilaterus (Haw) N. E. Br.
Portulacaceae
Calandrinia pygmaea F. Muell.
C. calyptrata Hook.f.
Caryophyllaceae
Spergularia rubra (L.) J. and C. Presl
Droseraceae
Drosera glanduligera Lehm.
Crassula pedicellosa (F. Muell.) Ostenf.
C. macrantha (Hook.f.) Diels and Pritzel
C. sieberiana (Schultes) Druce
Pittosporaceae
Sollya f^LSiformis (Labill.) Briq.
Papilionaceae
Trifolium dubium Sibth.
Geraniaceae
Geranium pilosum Forst.
Pelargonium drummondii Turcz.
Oxalidaceae
Oxalis corniculata L.
Myrtaceae
Thryptomene saxicola (A. Cunn.) Schau.
Agonis juniperina Schau.
Darwinia citriodora (Endl.) Benth.
Umbelllferae
Hydrocotyle callicarpa Bunge
H. diantha D.C.
H. blepharocarpa F. Muell.
H. hirta R. Br.
Trachymene pilosa Sm.
T. anisocarpa (Turcz.) B. L. Burtt
Homalosciadium verticillatum (Turcz.) Domin
Primulaceae
Anagallis femina Mill.
Longaniaceae
Mitrasacme paradoxa R. Br.
Gentianaceae
Erythraea australis R. Br.
Villarsia calthifolia F. Muell.
Solanaceae
Solanum nigrum L.
Scrophiilariaceae
Veronica calycina R. Br.
Bartsia latifolia Sibth. and Sm.
Lentibulariaceae
Polypompliolyx tenella (R. Br.) Lehm.
Rubiaceae
Galium murale (L.) All.
Stylidiaceae
Levenhookia dubia Send.
Stylidium corymbosum R. Br. var. proliferum
Benth.
S. calcaratum R. Br.
Compositae
Helipterum cotula (Benth.) D.C.
Quinetia urvillei Cass.
Rutidosis multiflorus (Nees) B. L. Robinson
Angianthus tenellus (F. Muell.) Benth.
A. humifusus (Labill.) Benth.
Hypochoeris glabra L.
Cenia turbinata (L.) Pers.
Podolepis lessonii (Cass.) Benth.
Acknowledgments
The author gratefully acknowle(iges the assist-
ance given by the following colleagues: — Mr.
J. H. Willis of the National Herbarium of Vic-
toria for his identification of several bryophytes
and lichens: Mr. R. D. Royce of the State Herb-
arium of Western Australia for identification of
several angiosperms; Mr. Roger Carolin of the
University of Sydney for his identification of
Pelargonium drummondii: the Superintendent
of Mapping, Mr. J. Ryan, Geodetic Section of
the Department of Lands and Surveys, Perth,
for providing the altitudes of the peaks of the
Porongurup Range; and Mr. D. Herlihy of the
Hydraulics Branch of the Public Works Depart-
ment, Perth, for providing rainfall data from
the Bolganup Dam, with the permission of the
Hydraulics Engineer, Mr. D. C. Munroe.
References
Bentham, G. (1863-1878).— "Flora Australiensis” (Reeve:
London).
Bibby, P.. and Smith, G. G. (1954). —A List of Lichens
of Western Australia. J. Roy. Soc. W. Aust.
39: 28.
Diels, L. (1906).— "Die Vegetation der Erde" VII. Die
Pflanzenwelt von West Australien. (Engel-
mann: Leipzig.)
Gardner, C. A. (1942).— The vegetation of Western Aus-
tralia with special reference to the climate
and soils. J. Roy. Soc. W. Aust. 28: 11-87.
Sainsbury, G. O. K. (1955). — A handbook of the New
Zealand Mosses. Bull. Roy. Soc. N.Z. 5.
Willis, J. H. (1953). — The Archipelago of the Recherche
3a. Land Flora. Rep. Aust. Geogr. Soc. No.
1: 1-35.
23
4. The Subterranean Freshwater Fauna of Yardie Creek Station, North
West Cape, Western Australia
By G. F. Mees*
Manuscript received — 22nd August. 1961
The fauna of the wells of Yardie Creek
Station consists of four species: a Synbranchid
eel which is described and named in this paper,
the Eleotrid fish Milyeringa veritas, and two
species of Atyid shrimps. All these species
show the usual characters of cave fauna; loss
of eyes and of pigment. The fact that all
species represent endemic genera suggests a
considerable antiquity of this fauna. A dis-
cussion of the geology of the area is given, but
no conclusion as regards the actual age of the
fauna is reached.
Historical Review and Introduction
The occurrence of a specialised subterranean
freshwater fauna at North West Cape was first
made known by Whitley (1945). In October
1944, Mr. Whitley visited Yardie Creek Station,
and was told by the station owner. Mr. E. Payne,
of the occurrence of blind fishes in the Milyer-
ing Well, one of the wells on the station. Mr.
Whitley and Mr. Payne paid a visit to the well,
and Mr. Payne climbed down into it and
managed to scoop up in his hat one specimen
out of about a dozen present at the time. Sub-
sequently this specimen became the type of
Milyeringa veritas Whitley.
Whitley (1945) gave some particulars about
the Milyering Well, and suggested that a sub-
terranean river might seep through its lime-
stone walls. In subsequent years Whitley men-
tioned the species Milyeringa veritas a few
times in publications, and in recent years addi-
tional specimens were received by the Western
Australian Museum. Some years ago live
specimens were on show in the annual Wildlife
Show in Perth, organised by the Western Aus-
tralian Naturalists’ Club.
Nothing essential was added to knowledge of
this subterranean fauna until 1958 when Mr.
Alf Snell, who had visited Yardie Creek Station
as maintenance man of a shearing team, found
amongst a sample of Milyei'inga’s he had col-
lected some shrimps. He presented these to
the Australian Museum. Sydney, whence they
were forwarded for identification to Dr. L. B.
Holthuis of the Leiden Museum (cf. Anon. 1959 h
Additional specimens collected by Mr. Snell in
May 1959, were donated to the Western Austra-
lian Museum, and were also sent to Dr. Holthuis.
During one of his visits to the Western Aus-
tralian Museum Mr. Snell also mentioned the
observation of ^‘blind" eels in one of the wells
at Yardie Creek Station, of which, notwithstand-
ing many efforts, he had not managed to catch
a specimen.
♦ Western Australian Museum, Perth, Western Australia.
It was mainly with a view to obtaining
material of this eel. but also to get larger series
of the known animals and to gain a general
idea about their habitat, that from the end of
July 1959 onwards Mr. A. M. Douglas and I
spent ten days on Yardie Creek Station. The
results of this stay were satisfactory, a specimen
of the eel was obtained, and series of shrimps
and Milyeringa veritas. Besides, collections of
birds, reptiles, and insects were made.
In May 1960 another short visit was made and,
thanks to Mr. Douglas’s nocturnal activity, a
second eel was collected.
In the meantime Holthuis (I960) described
the shrimps, which he found to belong to two
different, but related species. Hence, the fauna
of the wells, as at present known, consists of
four species: two shrimps, a goby and an eel.
Though one must always remain prepared for
surprises, I consider it unlikely that more species
occur and believe that the macrofauna of these
subterranean waters is now completely known.
In this paper, besides descriptions and notes
on observations, the results of investigations into
the following three problems are given: —
1. How long has the habitat where this
fauna is found been in existence?
2. How different, morphologically, are
the cave forms from their presumed an-
cestors living outside the caves?
3. What are the chemical and physical
properties of the habitat?
A correct answer to these questions would
mean a knowledge of how long it has taken the
cave fauna to develop from its presumably eyed
ancestors into what it is now, in other w^ords.
of the speed of evolution, or better, morpholo-
gical change, that has occurred under these
very specialised conditions. It is perhaps as
well to add here that I have failed because no
certainty could be obtained that this fauna has
evolved in the place where it is now living.
Description of Habitat
The area under discussion is the north-west-
ern part of the North West Cape Peninsula
(Fig. Ih The Cape Range, which reaches up
to the lighthouse at Vlaming Head, forms the
backbone of the peninsula. On the outside of
the range is a platform of between li and 3
km wide, and along the sea coast is a narrow
line of sand dunes. It is evident that on many
places fresh sand has been blown quite recently
over the older coastal platform.
24
Fig. 1.— Map of the North West Cape Peninsula. The area in which the subterranean fauna is known to
occur is dotted.
25
It is this coastal plain or platform that is of
interest, because that is where the wells ai*e
sunk. The whole platform is very flat and
stony, and in many places small and shallow
sink holes occur w'hich show, as do the artificial
wells, that the base of this plain consists of
almost pure coral rock, brownish white in
colour, mixed with fossil shells, etc. The sur-
face of the platform is about two metres above
water level.
This w'hole layer of coral rock is apparently
traversed by crevices, small holes, and connect-
ing corridors so that an extensive subterranean
netw^ork of wateiwvays exists, which forms the
habitat of the fauna under di.scussion. More
arguments for regarding the structure as this,
and not. for example, as a subterranean river
<as Whitley 1945. thought), follow' below.
The platform extends along the whole out-
side of the peninsula, but to the south I have
only been as far as the Tulki Well, and I do not
know' if the structure farther south is identical.
However, there is a length of at least fifteen
miles of coral-rock platform.
In this platform, at distances of five miles
apart, sheep wells have been sunk, which are,
from north to south: Five Mile Well (Chugorie
Well on the Admiralty Chart), Kudumurra Well
and Homestead Well (these tw^o are only a few'
hundred metres apart', Tantabiddi Well, Milyer-
ing Well. Tulki Well . Holthuis’s statement
that the Milyering and Kudumurra Wells are
20 miles apart is a .slip, for their actual distance
is 10 miles.
We visited and climbed down all these wells,
but only in three of them cave fauna was found,
and descriptions of these three follow here.
The Milyering Well (Fig. 2» is hacked out in
the coral stone; the upper rim is cemented, but
low^er down the coral stone was apparently con-
sidered solid enough by the builders and re-
mains uncovered. This makes climbing down
easy because everyw'here there are small dents
and holes in the sides which allow' one to get
a good grip. Normally the w'ell is covered by
a few' sheets of corrugated iron. Though the
cemented rim has a diameter of not over a
metre, low'er dowm the well is somewhat wider.
The w'ater surface is about tw'o metres down,
and the water in the w'ell is not over about 50
cm deep, on a cup-shaped bottom. But in the
sides of the walls, on and just below the water
surface, are some crevices, w'ater channels, the
places through w'hich visiting Milyeringa’s enter
and leave the w'ell.
A mill is cn top of the well, and pumps the
water in a biggish reservoir from where two
pipes lead on to tw'o drinking troughs. The
w'ater is very tasteless, w'hich is partly caused
by the high temperature, and partly by a fairly
high salinity (slightly brackish). The tempera-
ture. measured several times a day in the first
w'eek of August, 1959. and again on 16th May.
1960, was about 27° C.
* The Kudumurra Well is called Kiirumuru Well on
the Admiralty Chart No. 3187: Mangrove Islands
to North W'est Cape, new ed. 1915. Kurumurii may
well be the more correct spelling, but the people
who gave these names have gone for ever and their
meaning has been forgotten.
-r
Pig. 2.— Mr. Douglas at the Milyering Well. 16.V.1960.
Note the numerous blocks and pieces of coral stone
that cover the platform.
A further discussion, viz. of the water move-
ments and of the composition of the mud on
the bottom follow's after.
The Tantabiddi Well is a natural cavity that
either w'as already open, or of w'hich part of
the roof has been removed. Now it is a little
pod with several square metres of w'ater sur-
face, partly vaulted over. In tw'o directions one
sees deep tunnels or shafts, which, however,
lead upw'ards, and therefore are dry and cannot
be used by aquatic animals. The pool has a
greatest depth of about half a metre and the
bottom is covered with mud. There are no
open connecting channels; this is important in
connection with the distribution of the fauna
as I shall explain later on. There is the usual
Southern Cross windmill which, however, is not
straight above the waterhole, but some little
distance away. Probably because of its being
open, the temperature of the water is slightly
lower than in the other wells, about 24 ’ C. in
the first week of August, 1959. As the analysis
(Table I) shows, the water is definitely less
brackish than that of the Milyering Well, which
is doubtless caused by the fact that rainwater
has free access to this well.
The Kudumurra Well is the most prosaic
looking of the three, as it is entirely cemented.
The water-level is at about 1.75 m depth, and
about 20 cm lower down the cemented tube
26
which forms the well ends, and rests on some
horizontal wooden joists. The water is not over
50 cm deep and, just as in the Milyering Well,
crevices and channels seem to run in various
directions. The Southern Cross windmill is not
above the well, but at its side, some metres
away. Inscriptions on the rims of this and other
wells show that they were constructed in 1913.
and not in the early 1920’s as Whitley (1945)
would have it. The temperature of the water in
this well, in August, 1959, was 29-30^ C (84-86*"
F). on 17th May. 1960. it was 28.3 C (83" F>.
TABLE 1
Certificate of Analysis
Date received: 1st September. 1959.
Result of Analysis :
Marks:
Tantabiddt Well
Milyering Well
Reaction
Neutral
Neuti'al
pH
7.5
7.1
Mineral Matter
Parts per million
Calcium. Ca
113
126
Magnesium, Mg
141
188
Sodium. Na
999
1414
Potassium. K
36
53
Bicarbonate, HCOa
336
316
Carbonate. COa
nil
nil
Sulphate. SOi
240
358
Chloride. Cl
1810
2550
Nitrate, NO;i
3
3
Silica, SiOi
15
1 1
Iron oxide. Fe^O:}
less than 0.1
less than 0.1
Aluminium oxide,
AloO:; 5
10
Phosphate. POi
0.4
0.2
3698 5029
The mud on the bottom of all three wells
is very rich in organic matter. Particularly
the Milyering Well and the Kudumurra Well,
both of which are normally covered with sheets
of iron, act as traps for all kinds of animals
that seek shelter under the covering sheets. The
first time I descended the Kudumurra Well, for
example, I found a small python. Liasis child-
reni, swimming in it: if I had not saved it it
would doubtless have died and added to the
nutritive value of the debris at the bottom. In
the mud of this same well I found bones and
teeth of small mammals, and in all the wells
great numbers of exoskeletons of woodlice.
Woodlice live in some numbers under the covers
of the wells, and fall in the water regularly.
It is likely that, because of these favourable
factors, the concentration of fish and shrimps
in the wells is greater than it is elsewhere in
their subterranean domain.
Fauna
Family SYNBRANCHIDAE
Anommatophasina genus novum*
Superficially very similar to the eel from
Hoctun Cave near Chichen Itza, Yucatan, de-
scribed by Hubbs (1938) as Fluto infernalis.
hence to the genus Synbraiichus, but differs
from that species as well as from all other
members of the suborder Synbranchoidei by the
pocition of the anus, which is in the anterior
half of the body.
Assumed combination 07i evaporali07i at N.T.P .
Calcium carbonate. CaCO;i
Magnesium carbonate, MgCOj
Sodium carbonate, NaaCO.{
Calcium sulphate. CaSOt
Magnesium srilphate, MgSOt
Sodium sulphate. Na->SOi
Magnesium chloride. MgCU
Potassium chloride. KCl
Soduim chloride. NaCl
Sodium nitrate. NaNO»
Total hardness
Bicarbonate (temporary)
hardness
Non-carbonate (permanent
hardness
Calcium hardness
Magnesium hardness
276
259
nil
nil
nil
nil
9
75
293
382
nil
nil
320
434
69
101
2537
3592
4
4
um carbonate.
862
1089
276
259
586
830
282
315
580
774
Sgd. R. C. GORMAN.
Deputy Government Agricultural Chemist.
Water samples were taken from all three
wells, but the bottle from the Kudumurra Well
unfortunately lost its contents on the way down
to Perth. The result of a very full analysis of
the water cf the other wells is given in Table I.
There is no trace of current in the water of
any of the wells, which supports my opinion
of the nature of the subterranean water.
It is interesting that all three wells show
tidal movements: the water moves up and down
about 15 cm each day. According to my geo-
logist friends this does not necessarily mean
that an open subterranean ccnnection with the
sea exists, but rather that there are permeable
layers of sand, etc., which allow the tidal in-
fluences to be felt some way inland.
Type species:
AnommatopbasiTUi caiididum species nova
A slender Synbranchid adapted to life in total
darkness in caves. Head comparatively shoit,
its height about half its length; upper surface
of head behind snout swollen, with a shallow
longitudinal groove in the middle; head deeper
than any other part of the body and theiefoie
fairly well defined; mouth comparatively large,
cleft in lateral view about two-fifths of length
of head; lips thick, especially anteriorly; teeth
fairly strong, laterally in both jaws in a single
row anteriorly in a band of three or four rows
deep; a narrow band of teeth, parallel to the
jaws, on a ridge on each side of the palate:
tongue well developed, anteriorly free and
rounded to slightly truncate, without teeth; one
pair of very small nostrils at the tip of the
snout in the upper lip, a second much larger
pair of nostrils on upper surface of snout just
before elevation of forehead, roundish, each one
covered by a dermal flap that is attached on its
anterio-lateral rim: eyes absent, but anteiio-
ventrally of the second pair of nostrils is a sub-
cutaneous concentration of what appeals to be
nerve tissue, and which may well represent
vestiges of eyes; several pairs of mucous pores
are present on the head but they are difficult
to see, one fairly conspicuous pair is on the
snout half way between the first and the second
pair of nostrils; throat with some longitudinal
dermal folds; gill opening rather wide, trans-
verse, the covering skin lunate in shape; four
pairs of gills, well developed; body long and
♦From avofifxaT07 without eyes, and <paaua =
apparition, phantom.
27
slender, roundish, the last few centimetres of
the tail compressed; anus in anterior half of the
body: lateral line distinct and continuous to
near the tip of the tail, but I have been unable
to find pores in it: no fins except a thin rayless
fin membrane near and round the tip of the
tail, in which I have been unable to detect even
the vestiges of rays, but at the tip of the tail
four or five hypurals are present.
Material. Two specimens.
P 4917 Tantabiddi Well. Yardie Creek
Station. North West Cape,
31.VII.1959. Type.
Total length 370 mm. length of
head and body <to anus) 150 mm.
length of head from tip of snout to
gill opening 20 mm, cleft of mouth
from tip of snout to posterior
border of maxilla 9i mm.
P 4918 Tantabiddi Well, Yardi? Creek
Station. North West Cape,
17.V.1960.
Total length 316 mm. length of
head and body ito anus) 121 mm.
length of head from tip of snout to
gill opening 16i mm: cleft of
mouth from tip of snout to pos-
terior border of maxilla 7 mm.
Colour. The colour in life is a very striking
pure white. In a captured specimen we noticed
that in some parts a faint pinkish tone appeared
that was not originally present: presumably
this was caused by damage of small bloodvessels
as a result of its capture. Microscopic examina-
tion revealed the presence of dispersed pigment
in the skin which is irregularly but more or less
evenly distributed over the whole body and does
not show any particular pattern.
Abundance. Besides the Tantabiddi Well,
where our two specimens were obtained, one
example was observed in the Milyering Well.
Though we have not seen specimens in the
Kudumurra Well the species has been reported
10 occur there by one of the stationhands. The
limited number of observations conclusively
shows that this eel is far less plentiful than
Milyeringa though there is no reason to regard
it as particularly rare.
The explanation of the fact that in the Tan-
tabiddi Well only eels were found, and no other
cave fauna, is pi’obably that this well, contrary
to the other two wells, has no open connections
with subterranean waters. Such connections as
exist are filled with mud and debris, through
which the eels are able to move, but not the
free-swimming Milyeringa’s and shrimps.
Discussion. From the preceding description
it will be deal* that in all characters, except
those which are evidently connected with life
in darkness and which it shares with S.
infervalis*. this species fits very well in the
genus Synbranchtis, but for the anterior posi-
tion of the anus.
The position of the anus in the Syntaranchi-
formes has hitherto been regarded as of great
systematic importance, but it seems now that
the value of this character has been overesti-
• Reasons for placing Pl-uto" infernalis in the semis
Synbranchua are given in a later section of this
paper.
mated. Reference to the position of the anus
in the diagnosis of the suborder Synbranchoidei
as given by Regan (1912) and of the family
Synbranchidae as given by Weber & de Beaufort
(1916). to mention but a few authors, should be
eliminated. It is interesting to note that Berg
(1955) did not mention the position of the vent
in the diagnoses of any of the subdivisions of
the order.
On the other hand, it seems unlikely that the
anterior position of the anus in Anomviato-
phas 77 ia would be advantageous in cave life,
hence be adaptive (more about this will be said
in the section on convergence and nomen-
clature). Perhaps, however, food conditions and
the predator-free cave life may have been re-
sponsible for a shortening of the digestive tract.
Anatomical work and a study of feeding habits
and food may in future cast new light on this
problem. Until these points have been cleared
I feel perfectly justified in attaching generic
importance to the character. This point of view
finds mild support in the fact that both the
other fish and the shrimps inhabiting this
habitat have developed into indigenous genera.
As far as the relationships of Anommato-
phasina candiduni are concerned, the obvious
species to consider is Syiibranchus bengale7isia
(McClelland), which is the only other Syn-
branchid eel known to occur in Western Austra-
lia^ .
The genus Sy7ibra7ichus is in need of a re-
vision, and the various nominal species are very
similar to each other. I have compared speci-
mens of S. viar 7 noratus and S. bengaleiisis and
found that S. 7 narrnoratus apparently differs
from S. bengalensis by having a much narrower
gill-opening which is concealed in longitudinal
skin-folds, in having relatively larger eyes, and
more vertebrae. One specimen, however, is very
near S. bengalensis but differs by having a
slightly larger number of vertebrae; the anus
is slightly more posterior in position*. A?io?n-
matophas 77 ia has a wide gill-slit, as has S. ben-
gale 7 isis, but unfortunately S. inferiialis has, as
Hubbs's figure shows, also a wide slit though it
is supposed to have been dei’ived from S. mar-
vioratus.
' Wliitlev 11948. 1960) has resurrected the name Syn-
braiicluis gutturaHs Richardson for specimens from
Australia. ' As he has apparently never given reasons
for this I do not follow him. I do not belong to
that group of zoologists who regard the mere fact
of the occurrence of an animal in Australia in
itself as sufficient reason to separate It nomen -
claturally.
^ This specimen bears on its label the name Syn-
branchufi cJiilensis. Chili. Prank 1849. R.M.N.H. no.
3899. S. chilensis is apparently a manuscript name
that has never been published. The only Syn~
branchus at present known from the west coast
of South America is S. ynarmorahis which, how-
ever. has not been recorded from Chili, Peru being
Us known southern limit. In view of the somewhat
obscure origin of the specimen (localities supplied
by dealers are notoriously untrustworthy; as far as
Frank is concerned, see Gijzen 1938. p. 180), it
seems best not to attach much importance to this.
Though the specimen seems to show some slight
differences from S. 7narnioratus. the fact that it has
the same number of vertebrae points to Its be-
longing to that species. The mention of the name
S. cliilensis in this discussion must not be inter-
preted as a validation of this name for use in
zoological nomenclature (cf. Copenhagen Decisions
on Zoological Nomenclature. 1953. p. 63, § 114).
28
The differences in position of the anus be-
tween Synbranchus and A. candidum are quite
striking:
S. viarmoratus r•c/a^e?Ks^s”^
preanal length: postanal length 10 : 3
S. 7uar7Uorafws.
preanal length: postanal length 8 : 3
S. hengalensis,
preanal length: postanal length 8 : 3
A. candidum.
preanal length; postanal length 2 ; 3
Some of the specimens were X-rayed. The
number of vertebrae as counted from X-ray
photographs is:
body
tail
total
S. mar7norat7is
(R.M.N.H. no. 3899)
84
-hc.54
C.138
5. 7nar7noratus
(R.M.N.H. no. 16352)
87 + 48' 135'
(tip of tail damaged )
S. bengalensis
(R.M.N.H. no. 7146)
75
+ 54
129
S. bengale7isis
(same regd. no.)
77
+ C.53
C.130
A. candidtnn
(W.A.M. no. P 4917)
51
+ 111
162
A. candidu7n
(W.A.M. no. P4918)
54
+ 109
163
These figures show that the number of ver-
tebrae is a useful systematic character in the
group. Because I have been unable to examine
this character in all species of the genus Syii-
branchus and related genera. I have not men-
tioned the large number of vertebrae of
Anom7natophas7ua in the diagnosis of that
genus though I note that according to Regan
(1912) Synbra 7 ichus has 127-137 vertebrae. It
is clear, however, that A. ca7ididum differs from
the species of Synbranchus that were examined
both in having a decreased number of vertebrae
in the body and an increased number in the
tail.
The description of Typhlosy7ibranchus boueii
Pellegrin (1922) shows that in cne respect, the
presence of only three pairs of gills, that genus
is more different from Synbranchus than is
A7iommatophas7nci, but it has the anus in the
posterior third of the body. From A7ioin7naio~
p 7 ^as? 7 ^a it further differs by the strong pigmen-
tation and the small gill-opening.
Comparison of Ano 7 nmatophasma with other
genera of the Synbranchidae is superfluous.
Family ELEOTRIDAE
IVIilyeringa veritas Whitley
Milveringa i^eritas Whitley. Aust. Zool. 11. 1945, p. 36.
Fig. i5--MilyerinK, Yardie. 20 miles south-west of
Vlamingh Head, North West Cape. Western Australia.
MilveriJiga: Whitley. Aust. Zool. 11. 1947 (June 20|.
p 146 (Western Australia): Whitley, in: Biogcogr. Ecol.
Aust.. 1959. p. 142. 146 (Pig. 3 no. 34). 147 (in a well in
the North West Cape area); Holtluus, Crustiiccana l.
1960. p. 48 (Kudumurra Well).
Milyeringa veritas: Whitley. W. Aust. Nat. 1. 1947
(Dec. 15). o. 53 (Greylan Fluvifaunula ) ; Whitley. W.
Aust. Fish. Dept.. Pish. Bull. 2. 1948. p.
Australia): Whitley. Aust. Mus. Mag. 10. 1951, p. 162
(Milyering); Whitley. Aust. Mus. Mag. 11, 1954. P-
153 Fie. (north-western Australia); Whitley. Proc. Ro>.
Zool. Soc. N.S.W. 1954/55. 1956. p. 42 (Australia): Holt-
huis. Crustaoeana 1. 1960 (Jan.), p. 47 (MUyeriug
Well): Whitley, Nat. Freshw. Pish. Aust.. 1960 (Nov.),
p. 121 (North 'west Cape): Mees in Ann. Rep. W. Aust.
Mus. 1959-60. 1961. p. 23 (North-west Cape): Ride ^
Serventv. in Little (editor); Oir. Yearb. W. Aust. 1960.
no. 2 (n.s.). 1961, p. 67 (wells and subterranean channels
in the North West Cape area).
This curious goby was described and figured
by Whitley, to whose notes I have little to add.
Instead of D IV-9. A 9. it would be better to
write D IV-8i, A 8J. The scales are reduced,
and entirely absent from the head; there are
about 28 rows in a longitudinal line. The varia-
tion in size in our scries of about 50 specimens
is from 30 mm to 42 mm in standard length.
Colour. Yellowish white in life and health.
The brain is visible through the upper surface
of the head as a more or less triangular daik
patch. The fins are white as the rest of the
body. The flesh colour of the fins recorded by
Whitley (1945. 1960 » must be due to the same
cause referred to under A 7 iommatophasma
candidum. I have been unable to detect any
pigment in the skin.
Abundance. At our arrival on 29th Jul>.
1959, we found about half a dozen specimens
of Milyeringa veritas in the Milyering Well.
We caught them. and. as the well is fed by
subterranean channels, the original number was
gradually restored, so that during the after-
noon and evening we could harvest several
times. However, in my notes of 5tli August, it
is stated that, whereas on the first day every
time about six specimens could be taken, now
we did not find more than one or tw'o during
each visit. It is out of question that, by taking
specimens from one single well, w-e would have
been gradually depleting the whole population,
but it does indicate that the fishes do not swim
far or fast, that because of our activities they
had become scarce in the near surroundings of
the well, and that replenishment from more
remote areas took place too slowly to counter-
balance the influence of our collecting. On 16th
May. 1960, the usual half-dozen was present
again. Whitley (1945) slated that during his
visit about a dozen specimens were present in
the Milyering Well and that three times as
many had been seen in the well, but we nevei
found such large numbers.
In the Kudumurra Well the species is much
scarcer quite often no specimens at all were
present, usually only one or two. and never more
than three at a time. No specimens were ever
observed in the Tantabiddi Well.
Discussion. When Whitley (1945) described
Milyeringa ‘veritas, he not only placed the
species in a new genus, but also in a new family,
though suggesting that it was; "perhaps evolved
from some gudgeon similar to Carassiops, which
is not known from Western Australia.” Sub-
sequently Whitley < 1947 ) recorded Carassiops
compress 7 is from a well east of Carnal von and
added that; . they indicate the probable
line of descent of the interesting Western Aus-
tralian blind gudgeon, Milyeringa . . . .”
It should be remarked that the creation of a
new family or other high systematic unit for
a new species is about the cheapest way to
escape from the trouble of finding its true alfl-
nities. In the present case one may wonder
29
why a new family had to be established when
Mr. Whitley already suggested that Milyeringa
was derived from Carassiop 2 \ in other words,
if the new genus Milyeringa can be said or is
believed to be close to Carassiops and not to
other genera of the Eleotridae, Milyeringa and
Carassiops are evidently closer to each other
than either is to other genera of the Eleotridae.
and therefore Milyerit^ga can certainly not be
separated fi’om Carassiops as a different family.
The genus Carassiops is nowadays regarded
as a synonym of Hypseleotris (cf. Koumans
1953. p. 324 >. As I wanted to have an expert
opinion on the affinities of Milyeringa, some
specimens were foi'warded to Dr. Boeseman who
gave me (in litt.. 13.1.1960) as his opinion that
the species Milyeringa is closest to PrionobiLtis
microps (M. Weber). Prio7iobutis is a genus of
Eleotrid fishes that is supposed to be not dis-
tantly related to “Carassiops." Prionobutis
( Pogoneleotris! ^nicrops occurs in New Guinea
and in north-western Australia (Daly River) in
fresh and brackish water. The similarity in
many respects is striking, according to Dr.
Boeseman. and includes general shape, particu-
larly shape of the head, D 1.8 or 9, A 1.8 or 9.
C. c. 14, V 1.5. sc. c. 30, shape of mouth and
tongue (rounded-truncate), dentition, papillae
on snout, etc.
On the other hand, the differences between
Prionobutis and Milyermga are certainly large
enough to keep the two genera separate. Mil-
yeringa may cr may not have been derived
from Prionobutis, at present it is morphologi-
cally sufficiently different to be regarded as a
fairly well-marked separate genus.
Family ATYIDAE
Stygiocaris lancifera Holthuis
Stygiocaris iancifera Holthuis, Crustaceaua 1. 1960.
p. 48 — Kudumurra Well. Yardic Creek Station. North
West Cape Peninstila, W. Australia.
Stygiocaris lanci/era; Mees in Ann. Rep. W. Aust.
Mus. 1959-60. 1961, p. 23 {North-west Cape).
This species seems to be very much the com-
moner of the tw*o, for the material sent to Dr.
Holthuis consisted of 147 specimens of this
species as against 15 of S. stylifera.
As the two species were not distinguished by
me the following notes may apply to either of
them, but doubtless mainly to S. lancifera.
Shrimps were common in the Kudumurra Well,
but scarce in the Milyering Well and not found
in the Tantabiddi Well.
They are entirely colourless transparent, with
the exception of the internal organs of the
thorax, which show as a yellowish mass. The
contents of the intestine is visible as a straight
black stripe, but many specimens have the in-
testine empty.
In the Kudumurra Well I often observed
shrimps. In daytime it was very difficult to see
them, but at night in torchlight they were more
visible. They appeared to be resting, probably
also feeding, on the upper surface of the wooden
joists, on the walls of the well, and also on the
mud of the bottom. An occasional individual
would swim round freely just under the water
surface, possibly obtaining food from the sur-
face film. There is no evidence that digging
or rooting in mud takes place to any extent.
Stygiocaris stylifera Holthuis
Stygiocaris stylifera Holthiiis, Crustaceana 1. 1960,
p. 54 — Kudumurra Well, Yardie Creek Station, North
West Cape Peninsula.
Stygiocaris stylifera: Mees in Ann. Rep. W. Aust.
Mus. 1959-60, 1961, p. 23 {North-west Cape).
As noted under the preceding species, the
relative abundance of S. stylifera as opposed to
S. Umcifera is about 1 : 10 in the material col-
lected. This slightly larger species seems there-
fore to be much less plentiful.
Geological Evidence
Largely as a preliminary to and a result of
the recent intensive oil exploration in the region,
the geological structure of the North West Cape
Peninsula is well known (Condon, Johnstone &
Perry 1953). The spine of the peninsula is
formed by the Cape Range, the exposed parts
of which are mainly of Tertiary age: these
same authors state that the coastal platform
is Recent, but they do not devote any particular
attention to it.
To Dr. Logan (in litt., 16.IX.1959* I am
greatly indebted for much additional informa-
tion, most cf which I quote verbatim:
“I examined the plain along the western
piedmont of the Cape Range some years
ago, and I believe that it is a wave cut
platform related to a former sea level of
five to six feet above present mean sea
level. In a few places in the area one can
observe undercut pedestals or stacks of
limestone standing above the level of the
platform which exhibits most of the
characteristics of the undercut on the
shoreward periphery of the contemporary
‘reef’ flat which Fairbi'idge and others have
described at Rottnest, Point Peron and else-
where on the Western Australian coastline.
The higher sea level stand at about two
metres can be substantiated by emerged
shell beds, coral reefs and wave cut plat-
forms which occur at about tw^o metres
above present sea level in the Shark Bay
area. Allowdng for somewhat more tidal
amplitude in the vicinity of N.W. Cape the
Yardie platform can be well correlated with
this level; meaning that the area along the
foot of the Cape Range was inundated by
the sea in sub-Recent times.
Evidence all along the W.A. coastline and
elsewffiere in the world suggests a higher
stand of sea level at about two metres above
present due to eustatic causes (advance and
retreat of the polar ice caps). The evidence
for this eustatic sea level high is imposing
and the constancy of the terrace level over
wide stretches of coastline must tend to
rule out local uplift as a genetic cause of
this feature. One may expect slight varia-
tions in height due to local energy factors
in the marine erosion of the coastline as
differences in exposure to waves and tidal
amplitude must inevitably cause slight
variations in height of the platforms. I be-
30
lieve that the above explanation of the
platform along the edge of the Cape Range
is reasonable although one must be cautious
for the Cape Range is a fold mountain
which shows evidence for upwarping in the
Pliocene, Pleistocene and possibly the
Recent.
The two metre plus sea level stand of
sub-Recent times has been dated in various
parts of the globe by C14 dating techniques
which gives ages of about 5,000 B.P.
The actual limestone outcrop along the
foot of the Cape Range is similar litho-
logically to the Coastal Limestone along the
western coast of W.A. between Geraldton
and Dirk Hartogs Island: this is mainly
an aeolian limestone formed by terrestrial
agencies, and it dates from Pleistocene to
Recent in geological age. Some of the
platform may also be cut in Pliocene or
Miocene limestones which occur on the
flanks of the Cape Range
The evidence that the coastal platform is not
more than about 5.000 years old is quite conclu-
sive and can be accepted without reservation.
On the other hand, everything known about the
speed of evolution or morphological change of
animals pouits to its being a very slow process.
Even under the extreme conditions under which
the fauna under discussion lives, it is very hard
to believe that morphological changes of a
magnitude that demands generic separation
from their presumed ancestors would have
taken place in only 5,000 years. However, I
quote a further paragraph from Dr. Logan's
letter:
•‘As was found in the petroleum explora-
tion of the Rough Range, the limestones in
the N.W. Cape region are very cavernous
down to about sea level, and it is not out-
side the bounds of possibility that the
fauna developed and is living in these sub-
terranean caverns which may have been
connected to the sea at one time (for that
matter they may still be in some connec-
tion) and the wells are now tapping water
from these caves.”
Therefore I regard it as likely that the sub-
terranean fauna has developed in late Tertiary
or in Pleistocene times, in w’hat are now the
hills, and that w’ith the retreat of the sea dur-
ing the last 5.000 years, and the subsequent
emergence of the coastal platfoi'm and the de-
crease in salinity of the water, this fauna has
been able to colonise the platform from the
cavern systems in the hills.
Problems of Convergence and Nomenclature
Attention has already been drawn to the
extraordinary similarity in general appearance
between Anommatovhasina candidum and Syn~
branchus iiifernalis.
However, whatever the true relationships be-
tween the two species may be — and the aberrant
position of the anus in Anonmatophasiiia sug-
gests that they are only distantly related — it is
likely that each of them has been derived from
a different species of Synbranchid. Nevertheless,
if morphological evidence only was considered,
the two species would probably be legarded as
very closely related. Yet. what we really have
is convergence because it is fairly clear that
S. infei'nalis has been derived from S. maryno-
ratus, whereas the possible ancestor of
AnomniatophasTna was S. bengaleyisis. When
applying morphological criteria without histori-
cal considerations, '‘Pluto'’ inferyialis and A/iom-
matophasnia candidum might well be united in
one genus, and their respective ancestors in a
different one. Hubbs was aware of the fact
that his "Pluto" infernalis has been derived
from S. jnarmoratus. and therefore is histori-
cally closer related to S. marynoratus than that
species is to the other species of Sytibrayichus,
but notwithstanding that, he allowed morpho-
logical facts to prevail when he created for his
blind eel a new genus.
Of the conflict between historical relationship
and morphological similarity in what we please
to call the ‘‘natural system.” most systematists
are doubtless aware, and I do not see how it can
ever be solved'. It seems appropriate, however,
to draw attention to the fact that in our whole
system authors use nearly always words like
"affinity” and "relationship.” for what is actu-
ally morphological similarity. Though the
jargon has changed with the times, the method
is still essentially the same as that used by
Linnaeus and his contemporaries, tw'o centuries
ago.
Personally I believe that ideally in a natural
system actual relationship should be expressed
rather than morphological similarity, but I am
well aware that only in a very few cases, as
that of "Pluto" it is possible to distinguish be-
tween the two. and even when the distinction
is clear, group names (families, genera, etc.)
are applied in an arbitrary manner. For
example, birds and mammals are probably
closer related to certain orders of reptiles than
the latter are to other orders of reptiles, yet we
retain the classes Aves, Mammalia and Rcptilia
— to do other than adopt this arbitrary (but
phylogenetically incorrect) procedure w'ould lead
to nomenclatural chaos. Therefore, subjective
judgment in which historical knowledge (in-
cluding the fossil record if available) is weighed
against morphological criteria, will continue to
be the basis of our system of classification.
It seems to me that Holthuis (I960' has not
found a solution of the problem. In his diag-
nosis of the getms Stygiocaris he wrote;
"The genus is closely related to Tyyohlopatsa
Holthuis from Madagascar perhaps
Stygiocaris should only be considered a subgenus
of Typhlcpatsa" I do not claim to have even
the slightest knowledge of shrimps, and there-
fore am unable to evaluate their morphological
characters. But I regard it as unlikely that
blind shrimps from fresh w’ater in Madagascar
and blind shrimps from fresh water in Western
Australia, separated by many hundreds of miles
of ocean, would be nearer related to each other
than either of them would ba to some eyed
species living along the coasts of Madagascar
and Western Australia respectively. Instead of
the w'ords “closely related” Holthuis should have
* Of course I am aware of the existence of a lari?e
philosophical literature on the subject.
31
written “moi-phologically closest.” for this mor-
phological similarity may well be due to con-
vergence.
Several times I have mentioned the fact that
I regard certain characters as not of generic
value because they are connected with life in
darkness. This point of view is likely to meet
with criticism because it may well be argued
that any character is adaptive. However, most
zoologists nowadays do not attach too much
systematic significance to characters that are
evidently connected with some particular w'ay
of life in animals that otherwise are morpho-
Icgically close to other species. This would
apply even more where a loss of characters is
involved as a consequence of the absence of
selective pressure for their retention.
It is on the basis of the preceding argumenU
that I prefer to place "Pluto’' infernalis in the
genus Synbranchiis. so that the species should
now be known as Synhranchus infernali!^
(Hubbs». The genera Pluto Hubbs and Fur-
mastix Whitley 1951 ( nomen novum for Pluto
Hubbs, preoccupied*, consequently enter into the
synonymy of Synbranchiis. It will be noted
that I have based a separate genus for the Aus-
tralian blind eel not on characters common to
all cave fishes, but on the anterior position of
the anus, a character that is unlikely to be
directly connected with life in total darkness
*as it is not found in S. inferjictUs. which lives
under similar conditions).
Orientation
Very little can be said by me about the sub-
ject. All the “blind” species have probably lost
their eyesight completely, which seems evident
from the complete absence of pigmentation in
the ocular region.
Nevertheless, some orientation must occur:
particularly Anommatophasma has a habit of
making straight for dark crevices when being
disturbed out in the open of a well, and it
seems likely that some kind of light pcrcepticn
exists. Tactile factors could hardly be involved
in this kind of orientation. On the other liand
it is well known that some other senses, like
smell, are extremely highly developed in .some
fishes. Without experiments and anatomical in-
vestigations it will be difficult to decide if any
perception of light still cccurs in the species:
light sensitivity 'which is not sight* may well
exist, it may be located in the pineal organ as
is the case in other eyeless fishes, or in the eye-
rudiments.
Acknowledgments
From the section on the geology of tlie coastal
platform of Yardie Creek Station, it will be
evident hew much I owe to Dr. B. W. Logan
'Agricultural and Mechanical College of Texas.
College Station. Texas*. Discussions with Dr.
J. E. Glovei* and with Messrs. C. W. Hassell and
E. W. S. Kneebone (all of the Geology Depart-
ment. University of Western Australia, Ned-
lands*, further assisted in giving me a clear idea
of the geology of the region.
The X-ray photographs were taken by courtesy
of Mr. R. W. Stanford. Department of Medical
Physics, Royal Perth Hospital.
To the authorities of the Government Chemi-
cal Laboratories, Perth, I am indebted for the
very complete chemical analysis of the collected
water samples which is reproduced in Table I.
Dr. M. Bceseman (Rijksmuseum van Natuur-
lijke Historie, Leiden*, examined specimens of
Milyeringa, and gave me his valuable opinion
cn their affinities; he also sent me on loan
specimens of various species of Synbranchid eels.
The manuscript was read by Drs. M. Boeseman.
L. B. Holthuis and W. D. L. Ride, to ;:.ll of w'hom
I am indebted for useful suggestions.
To the keen interest cf Mr. A. Snell we owe
the information that with the discovery of
Milyeringa veritas the subterranean fauna of
Yardie Creek was not yet fully known. This not
only led to the discovery by himself of the two
species of shrimps, but ultimately also to our
trip to the region and the capture of specimens
of the remarkable Ano7nmatophasma candiduni-
Finally, both Mr. Douglas and I want to ex-
press our sincere gratitude to Mr. and Mrs. W.
D'Arcy of Yardie Creek Station, who not only
allowed us to come and stay with them and col-
lect on the station but who also assisted in
every possible w-ay to make our trio the complete
success it became. The same thanks have to be
extended to Mr. E. Payne, owner of the station,
who. unfortunately, we did not have the pleasure
of meeting.
References
Anon. (1959).— New cave-dwelling shrimps ioimd. AusL.
Mus. Mag. 13: 86.
Berg. L. S. ( 1955).— Sistema ryboobraznykh i ryb. nyne
zhivushchikh I iskopajemykh. Trai\ Inst.
Zool. Acad. Sci. U.S.S.R. 20.
Condon. M. A.. Johnstone. D. & Perry, \V. J. (1953).—
The Cape Range structure. Western Austra-
lia. Ft. I. stratigraphy and structure. Bull.
Bur. Min. Resour. Ailst. 21; 7-42.
Oijzen. .A. (1938). — ” ‘s Rijks Museum van Natiuirlijke
Historie 1820-1915'' iBrusse: Rotterdam.)
Holllmis. L. B. (I960). — Two new species of Atyid
shrimps from subterranean waters cf N.W.
Australia (Decapeda Natantia). Crustaccana
1: 47-57.
Hubbs, C. L. 1 1938).— Fishes from the caves of Yucatan.
in A. S. Pearse: Fauna of the caves of
Yucatan: 261-295. Pub'.. Carneg. histn. no.
491.
Koumans. P. P. (1953).-- “The Fishes of the Indo-Aus-
tralian Archipelago.” X (Brill: Leiden.)
PeUegrin, J. (1922). — Sur un nouveau pcissou aveugle
des eaux douces de I’Afrique occidentale.
C.R. Acad. Sci. Paris 174: 884-885.
Regan, C. T. (1912). — The anatomy and classification
of the Symbranchoid eels. A7i7i. Mag. Nat.
Hist. (8) 9: 387-390.
Weber. M. <sz de Beaufort, L. F. (1916). — “The Fishes of
the Indo-Australiiin Archipelago.” Ill (Brill:
Leiden.)
Whitley. G. P. (1945). — New sharks and fishes from
Western Australia. Part 2. Aust. Zool. 11:
(1947). — New sharks and fishes from Western
Australia. Part 3. Aust. Zool. 11: 129-150.
(1948). — A list of the fishes of Western Aus-
tralia. W. Aust. Fisk. Dept., Fish Bull. 2.
(1951). — New fish names and records. Proc.
Rotj. Zool. Szc. N.S.W. 1949/50; 61-63.
(I960). — “Native Freshwater Fishes of Aus-
tralia." (Jacaranda Press: Brisbane.)
32
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Journal
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Royal Society of Western Australia, Inc.
Volume 45
1962
Part 1
Contents
1- Variation, Classification and Evolution in Flowering Plants — with particular
reference to Thysanotus. Presidential Address, 1961. By Norman H. Brittan.
2. — The Largest Known Australite and Three Smaller Specimens from Warra-
lakin. Western Australia. By George Baker.
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