FIELD NOTES ON RABBIT BANDICOOTS, MACROTIS LAGOTIS REID

(MARSUPIALIA), FROM CENTRAL WESTERN AUSTRALIA

BY D. R. SMYTH*} AND C. M. PHILPOTT*

Summary

The numbers and range of the rabbit bandicoot have greatly diminished in recent years. The results

of a natural history study are presented to act as a basis for future research, and to facilitate

conservation of bandicoot populations.

At least two bandicoots were living in one square mile of scrub containing mulga, spinifex and

tussock grasses, near the Warburton Range, Western Australia. Their burrows, scratchings, tracks

and faeces are described. Fifty-eight burrows were mapped and 55 of them found to be distributed

at random within two areas delineated by presence of bandicoot scratchings. The main food species

of the bandicoot were the termites Hamitermes (Drepanotermes) rubriceps (Froggat) and Eutermes

tumuli Froggat. On each of 8 consecutive nights from 2% to 27% of the burrows were used. Out of

35 representative burrows, 14 were not used on any of the 8 nights, and none were used on more

than 4 nights.

FIELD NOTES ON RABBIT BANDICOOTS, MACROTIS LAGOTIS REID

(MARSUPIALIA), FROM CENTRAL, WESTERN AUSTRALIA

By D, RB, Smyru*f and C. M, Paiwrorr*

[Read 2 October, 1967]

* Royal Zoological Soviety of South Australia, Frome Road, Adelaide, South Australia 5000;

f Present address: Research Sehool of Fhological Sciences. Australian National University,

Qynberra, A.CLT. 2600,

SUMMARY

The munbers and range of the rabbit bandicoot have greatly dirinished in

receut yeurs. The results of a natural history study are presented to act as a basis

for future research, and to facilitate conservation of bandivoot populations.

At Jeast two bandicuots were living in one square mile of sernh containing

mulga, spinifexs and tussock prasses, near the Warburton Range. Western Aus-

tralia, Their burrows, scratchings, tracks and foezes are desertbed. Fifty-eight

burrows were mapped and 55 of them found to be distributed at random within

two areas delineated by presence of bandicoot soratehings. The main food spe+

cies of the bandicnut were the termites IJamitermes (Drepanotermes) rnebriceps

(Froggat) and Hutermes tumuli Froggat- On cach of 8 consecutive nights from

2& to 27% of the burrows were used. Ont of 35 representative hutrows, 14 were

not used on any of the 8 nivhts, and none were used on more than 4 nights,

INTRODUCTION

Since the coming of European man to Australia, the range and population

size of many marsupials have been greatly reduced. In some species it is probable

that this reduction has reached its limit, and that the species are extinct, Calaby

(1963) suggests that there are six such species. If so, then further study of them is

restricted to museum specimens and fossils, Calaby also lists other marsupials

close to extinction. The biology of most of these is little-known, and opportunities

to record it imcommon. Calaby includes the rabbit bandicoot, Macrotis lagotis

Reid, shown in PI, 1, Fig, 1, amongst these species,

Recently, we studied some of the natural history of a small population of

rabbit bandicoots near the Warhurton Range, Western Australia. Previous work of

this uature in the genus Muerotis has mostly been in the form of brief observa-

tions. made during collecting trips. Wood-Jones (1924) made many such obserya-

lions. He diseusses notes recorded by other authors, and also describes aspects of

behaviour of captive rabbit bandicocts,

Our aim in recording and discussing our results is to provide a background on

which other studies on rabbit bandicoots may be based, or with which they may

be compared, If these handicoots were to become extinct, we hope our study will

give future workers some idea of how they lived.

METHODS

During 1866, we surveyed the present range of rare marsupials in parts of

arid Australia, the results of which have been reported (Philpott and Smyth,

Trans. Roy. Soc. $.A. (1968), Vol. 92.

4 D, RK, SMYTH ann C. M. PHILPOTT

1967). During this survey, we located an area near the Warburton Range where

rabbit bandicoots were living, and used it as our study arca. We carried out the

work recorded here during November and December, 1966.

We were taken to the areca by semi-nomadic aboriginal people of the Ngu-

‘nyatjara and Pitjantjatjara tribes, These people have an intimate knowledge of

the signs and habits of the rabbit bandicoot, because they still hunt it occasionally

for food. They were happy to communicate this knowledge to us. This communi-

cation was facilitated by use of the aboriginal names of the rabbit bandicoot. Near

the study area, they call it.the “ninu”. East, in the Musgraye and Everard Ranges,

it is called the “talku”, while south-west towards Laverton it is called “matura”, All

aboriginal names are spelt phonetically following Douglas (1964),

We mapped the burrows in the study area by triangulation using a prismatic

compass.

To record if burrows had been used overnight, we inspected them daily,

swept away fresh tracks from the mouth, and placed a twig in the single entrance

such that it would be dislodged by a bandicoot entering or leaving. We took care

to minimize our scent on the twig und around the entrance of the burrow. We

scored a burrow as being used by # rabbit bandicoot if the twig was knocked

down and characteristic tracks and tail marks left near the cntrance.

Faecal contents werc examined microscopically after crushing in water con-

taining a drop of detergent.

We identified plant species in the field, and checked several in the State Her-

barium of South Australia. Ants were identified at the C.S.I.B.O, Division of Ento-

mology, Canberra, and termites and insect larvae by staff at the South Australian

Museum.

RESULTS

(a) General description of the study area

The study area is about 7 miles (11 km) north of Warburton Mission, and its

latitude and longitude are 26°02’ S. and 126°34' E. respectively. Its position. is

shown in Fig, 1, It is an irregular shape, with a maximum length of 1-4 miles (2-3

km) and is 1-0 mile (1-6 km) across at its widest point. Fig. 2 is a sketch map of

the study area, which has a total area of 1-0 square mile (2-6 sq. km),

STUDY AREA : :

aXe Fig. 1, Location of the study area

near Warburton Mission, Central

Eth Australia. with the general area

shown om an indented map of

Australia.

FIELD STUDY OF THE RABBIT BANDICOOT

MULGA-TUSSOCK —

MULGA- SPINIFEX

OPEN SPINIFEX a

Fl OPEN ROCKY SAVANNAH + BANDICOOT BURROW #f |

OPEN MULGA = RABBIT ae | IG,

wet

- LIMIT OF

SCRATCHING

Vig. 2, Sketch map of the study wea, showing approximately the positions of 58 bandicont

burrows, the limits of bandivoot seratchings, and the areas covered by the 5 yegetation groups.

The following general description is based on observations and on informa-

tion from the Atlas of Australian Resources (1959).

The area has a dry continental climate, with irregular rainfall, but with the

maximum amount falling in summer. The average annual rainfall is 5-8 inches

(13-20 em). The average daily maximum température in January (summer) is

95-100" F (35-39° C), In July (winter), the average daily minimum is in the

range 40-45° F (4-7° Q).

The study area is bounded by lew rocky ridges on the north and north-west.

From the low pass between them, the land slopes gradually to the south-west.

Skeletal soils cover the hills, while the flats and plains are of deep, desert loams,

with small rocks and pebbles distributed through all Jevels. Permanent surface

water is absent from the study area, and uncommon nearby.

Tor the purpose of description, the vegetation of the arca has been divided

into five classes. These are not discontinuous, and are based on subjective observa-

tions.

(i) Open rocky savannah. This type occurs on the skeletal soils of the ridges

and hills. Perennial hard-leafed grasses and occasional shrubs such as. Cassia and

Piiletus obovatus provide a sparse coyer. Mulga (Acacia. aneura) also occurs at

very low density,

(ii) Open mulga scrub, Pl. 3, Fig. 1—Mulga occurs in stands of low to

medium density in certain areas. There are no shrubs or perennial herbs under the

open mulga canopy. Ephemeral crucifers, grasses, composites and other herbs

spring up on the bare ground after rain, The soil is often shallow.

6 D, A. SMYTIT ann C. M. PHILPOTT

(iii) Mulga-tussock scrub, Pl, 3, Fig. 2—This is similar to the second type,

except that the mulga is usually more dense, and the ground has a moderate cover

of perennial tussock grasses, The soil is probably deeper than in the previous

class,

(iv) Mulga-spimifex scrub, Pl. 3, Fiy, 2—On the deep desert loams, mulza

and spinifex (Triodia bysedowii) often occur together at medium densities. Many

other specics of trees and shrubs grow as scattered solitary individuals. Such

trees as the bloadwood (Eucalyptus terminalis) and the corkwood (Hakea sub-

erea) occur, The shrubs present include the dead finish (Acacia tetragzonophylla),

the witchetty bush (Acacia kempeana), aud various Cassia and Eremopliila

species. Some herbs and grasses also occur,

(v) Open spinifex glade, Pl. 3, Fig. 4—In some areas, the major plant cover is

spinifex, Occasional individuals of mulga and witchetty bush occur. Ephemerals

grow in the spaces between spinifex clumps after rain,

The approximate limits of these classes in the study area are shown in Tig. 2.

(by) Signs

We did not see any rabbit bandicoots above the surface of the ground during

daylight hours, They are probably strictly nocturnal. Their presence in an area

seatched during the day can only be proved by capturing a specimen. However.

various characteristic signs of their presence can be readily secn.

(i) Burraws—The most obvious sign is the hurrow. We located 58 bandicoot

burrows in the study area, and completely dug out six. A sketch diagram of two

representative burrows is shown in Fig, 3.

Fig. 3. Sketch diagrams of two

burrows. which were dug ont com-

pletely. (1) and (2) are the side

elevation and plan respectively of

«burrow near the unrth-east

corner, (3) and (4) are the side

eleyation and plan of a burrow an

the central western edge, X indi-

cates the position from which 2

temale rabbit bandivoot was taken.

ENTRANCE

A small cireular mound of soil was present outside the entrance of all bur-

rows. The tunnel cross-section was usually cirenlar or slightly oyal, and about 6

inches (£5 cm) in diameter. Burrow entrances were often partially hidden by

dense spinifex, tussock grass or juvenile mulga plants. Occasional burrows were

underneath or near fallen logs. Eight of the burrows were conspicuous by being

dug into the large mounds of loose soil frequent in the area. Two representative

lurrows are shown in Pl, 2, Figs. 2 and 3.

FIELD STUDY OF THE RABBIT BANDICOOT '

In two burrows which contained a solitary rabbit bandicoot, the tunacls were

blocked in from one to three places along their length by loose, frequently dug

soil. There were no faeces in, or close to the burrows. In one of the burrows con-

taining a bandicoot, an area of the floor near the end was damp, possibly from

recently voided urine. The two bandiceots were at the extreme ends of their res-

pective burrows, und were rapidly lengthening them when captured.

Rabbit bandicoot burrows were different from other types of burrows present

in the area. Rabbit ( Oryctolagus cuniculus Line) burrows are often built in war-

rens, are of a different plan, and are often marked by facees, We located two bur-

mows which were probably used by rabbits in the study area, Goanna (Varanus

sp.) bnrrows which were also present, were smaller, semi-circular in cross-section,

and much shallower.

(ii) Scratchings—Diggings and scratchings covered the soil over inuch of the

study area, They were of distinct types. (1) The most frequent type was a shal-

low, cylindrical pil, 2-8 inches (5-20 cm) deep, and 2-6 inches (5-15 cm) wide, an

example of which is shown in PI. 1, Fig. 2. The soil from them was piled up in all

directions around their mouths. (2) There were also smaller seratchings of irregn-

lar shape, and areas of looseued topsoil, often near mulga roots. (3) A further

tyne was a conical pit under certain tussock grasses, as illustrated by Pl. 1, Fig. .3,

Chit of a sample of 107, 62% were under Eragrostis erivpoda and EF. luniflora, 18

under Denthonia bipartita, and the rest under Aristida contorta. and Eriachne

mucronata. These pits were mostly in mulga-tussock scrub. They were about 4-8

inches (10-20 cm) deep, and descended at an angle,

We helieve that all these diggings were made by rabbit bandicoots. The evi-

dence which suggest this is: (1) the diggings were distributed in close association

with the handicoot burrows, as shown in Fig, 2, and were never more than 600

feet (180 m) From one; (2) the density of seratchings was frequently higher in

the vicinity of a burrow; (3) faeces, later shown to be closely similar to those of

recently captured bandicoats, were occasionally found on or near the freshly dug

soil; and (4) insect species, fragments of which were found in newly captured

bandicoot's faeces, (see Kesults, Part [d]), were often found in the seratch-

ings,

(iii) Tracks—Characteristic tracks were often left in burrow entrances and on

fresh scratchings. They were also left on the pebbly surface between some bur-

rows in the area. A sketch diagram of a set of tracks entering the mayth of a

rubbit handicoot burrow is shown in Fig. 4. Although we have only aboriginal

opinion and circumstantial evidence, we believe these charactcrishe tracks were

those of a rabbit bandicvot.

We observed charactcristic grooves on loose soil near scratchings and also

near burrow entrances, as shown in Pl]. 2, Fig. 3. Captive bandicoots left stmilar

Fig. 4. ‘Sketch diagram of a

series of presumed, rabbit byn-

dicont tracks leading to the

entrance of a burrow. The

probable direction of travel is

shown with an ariiow,

§ D, R, SMYTH asp C. M. PHILPOTT

4 Fig. 5, Frequency distribution

of 58 burrows, showing the dis-

tance of the nearest neighbour-

ing burrow to each of these

burrows. The mean distance is

249 feet with a standard error

4 of 14-1 feet,

NO. OF BURROWS

i) 100 200 300 400 500 600

DISTANCE OF NEAREST NEIGHBOURING BURROW (FEET)

Fig. 6, Frequency distribution

ie of 85 burrows, showing the

number of nights each of these

g burrows was used over a period

\ of 8 consccutive nights. The

~~.

mean number of nights is 1-14,

and its standard error is 0-036

nights,

OF BURROWS

No.

——— 2 « —__—s

0 1 2 3 4 5 6 7 2

NO, OF NIGHTS EACH BURROW USED

marks while digging in loose soil on the cage floor. The marks were left by the

tail, which was used as a strut while the hind legs were in use.

(iv) Faeces—I'aeces of captive bandicoots were similar in shape and size to

those found occasionally on or near scratchings. Four representative groups of pel-

lets are shown in Pl. 2, Fig, 1.

(ce) Distribution of burrows

The distribution of the 58 burrows over the study area is shown in Fig. 2.

Most burrows were in mulga-tussock scrub and mulga-spinifex scrub. Two were in

each of open mulga scrub and open spinifex glade. There were none in open

rocky savannah.

FIELD STUDY OF THE RABBIT BANDICOOT 4g

The burrows were distributed in two clusters bounded approximately by the

limit of bandicoot scratchinys. In order to test if the burrows were distributed at

random within the scratched arcas, a grid of quadrats with sides equivalent to 340

feet (105 m) was laid over a map of the area. We considered only those quadrats

within the scratched areas, and those intersected by the limit of scratchings. The

distribution of burrows within these quadrats was tested for goodness of fit with a

Poisson distribution, as shown in Table 1.

TABLE 1

Distribution of burrows m quadrats covering all of the seratchod area, showing goodness of fit

test Lo a Poisaon distribution.

No, of Observed Expected (O—f)*

burrows No, of No. of —--

per quadrats quadrats EH

qitartrat (QO) (BE) |

0 60 | 61-28 027 | Mean = 0-529

ot u2e41 | O07 | Wee = 0-458

2 , 9 $57 x o> 1d

, 1 1-5] L b-0n0 70%, p= 80%),

tor iuicre | i ; 0-25

Total | 14 104-00 ees

The result (x7, = 0-114, 70%4< p < 80%) strongly suggests that the burrows

were randomly distributed over the area, This indicates that the position in which

a bandicoot dug a burrow was independent of the position of other burrows in the

area.

It is probable that a rabbit bandicoot uses a burrow as a refuge into which to

flee when danger threatens. If so, then some description of distances between bur-

rows is useful. Fig. 5 shows a histogram of the number of burrows and the dis-

tance of the nearest neighbouring burrow to each of these 58 burrows taken in

turn. The spread of the nearest neighbour distance is small, with no burrow more

than 550 feet (170 m) from any other one.

There is a small peak in the histogram composed of burrows with nearest

neighbour distance greater than 350 feet (105 m). These burrows were distri-

huted over the whole study areca, and the presence of this peak is therefore per-

haps fortuitous.

(d) Food

We collected all the faeces voided by two bandicoots during their first night

in captivity, One pellet from each was examined microscopically after crushing,

They both contained more than 50% by volume of soil and grit. They also con-

tained a small number of rabbit bandicoot hairs, However, the most striking con-

tents. were the hard, keratinised mandibles and whole heads of certain insects, We

collected insects from the study area which were possible food species, dissected

their mandibles, and compared these mandibles and heads with those found in the

faecal pellets. By this method we were able to identify remains of both soldier

LO D, R, SMYTH snp C. M, PIITLPOTT

and worker castes of two termite species, Hamitermes (Drepanvtermes) rubriceps

(Froggat) and Eutermes tumuli Froggat, and workers of one species of ant,

Camponotus sp.

Counts of the numbers of mandibles, and in two instances twice the number

of heads (for comparative purposes), of these species are given in the first two

rows of Table 2, ‘

TABLE 2

Frequency of mandibles of yarivus insect specics in facenl pellets from recently captured rabbit

bandicoats and from near scratchings made by bandicoots in the study area,

Nowak. Termites | Ant Various

Origin of pellets Tridomyr- | unknown

pellots examined) | Hamitermes rubriceps Budermes tumuli ned apecins

{Total No. | Worker — Soldier Worker Soldier* — species*

Bandieont I 1/5 S 1 TO 34 { 0 OF

Baucheoot 2 1/12 4 0 60 1k 0 OF

xtreme 13 5/5 76 1 a. 4 V) cf)

odge 57 1 1 {) | i} )

72 2 ! £ H 0 fi

| 18 4 a 2 0 0

| 120 ih 6 2 j () } 4

SW Centre 2/2 70 4 0 0 2 1

59 ‘) H 12 0 ‘)

Extreme Ho! 2/2 87 i i) 2 174 i)

edge S85 f) 0) 2 140 Of

H eentre 1 1/4 221 ti) u 2A i) 0

Central E 0-5/4 HA ‘) 2 iE 4 i)

edge }

Centre O-a/2 26 0 i) 0 8 2

* No. of heads » 2.

F +1 mandible of Camponotus species.

~ +2 mandibles of Camponotus specios.

§ $2 manilibles of larval Lepidaptera.

The counts may not represent the relative numbers of individuals eaten. Cap-

tive bandicoots masticated their food very thoroughly, and it is possible that a dif-

ferent proportion of mandibles and heads from different food species were

crushed beyond recognition. However, most mandibles from pellets were whole,

and little changed in shape or colour. There is probably some simple relationship

between numbers eaten and fragments in faeces.

After working with faeces known to be bandicoots’, we analyzed contents

from presumed handicoot faeces collected near scratchings. These results are

given in the remainder of Table 2. They closely resemble those from. known ban-

dicoot faeces. In these other pellets, we identified remains of a small black ant,

Tridomyrmex sp., and a white Lepidopteran lJarya, as well as species found in the.

earlier analyses.

The two species of termites were abundant ever much of the study area.

They were found in vast underground colonies, with domes and mounds raised

above the soil surface in places. Bandicoot seratchings often intersected the under-

ground passages, bit there was no sign of interference with the mounds. From the

numbers of mandibles in the pellets, and from the numbers of scratchings inter-

secting colonies, it is likely that these termite species were a primary source of

food for bandicoots in the study area.

MIFIM STUDY OF TIE RABBIT BANDICOOT i

The species of Lridomyrmex identified from faeces was also extremely abun-

dant over the area, Being highly active and pugnacious, they readily attacked and

carried off termites when given access to their colonies, Because of this, and their

ubiquitousness (therefore being readily available where a bandicot was feeding

on termites), as well as the fact that their remains only occurred in large numbers

in one group of pellets, it is possible that they are ouly a subsidiary food sonree,

or perhaps even a “contaminant” of the usual diet.

The larger ant, Camponotus sp., was less abundant than Iridomyrmex, Ts

sporadic appearance in faeces suggests that this is also a secondary food species.

We found the large white grub (larval Lepidoptera) under fwo specees of

tussock grass, Eragrostis eriopoda aud FE, laniflora. Of the five grasses scratched

imder in the study area, these two were the most frequent. Although mandibles of

this particular larva were tound only twice in a safaple of 14 Keller the large size

of the grub (up to 20 mm long), and the large number of tussocks scratched

wnder, suggests that larvae of this species and perhaps other unidentified ones are

an impertant food.

Faeces oecasionally comtuined other materials. Small amounts of plant tissue

such as undigested seeds and plant hairs were sometimes present. Unidentified

thickened tissue also oceureed in moderate amounts. These objects were not eon-

sistently preseut,

(oc) Activity

For eight consecutive days we kept a record of the burrows that had been

used during the previous night, Thirly five burraws were scored on all eight

nights, and more were added ta tlic list as we found them, For the last two days

we scored 52 burrows, and considered thal we had located at least 90% of all the

lirrows present. Our method of scoring did not differentiate between a single

entry or exit by a bandicoot, nor did it dillerentiste either of these from inultiple

entries and exits during one night.

We must assume firstly that our working in the area did not have any effect

on the behaviour of the bandicoots present. Although we have no concrete evi-

dence to suggest that this is a valid assumption, we noted that recently captured

bandicoots showed little fear of hurnans or human scent, Secondly, we will assume

that the burrows we observed for the longest period were a representative sample

of all burrows with respect to use by a bandicoot. We could not detect any visible

difference between these burrows and the remaining ones.

For each night we calenlated the percentage of burrows which were used oul

pf all those we scored. The mean percentage and its standard errov were 15-5 +

2-6%, with a range of from 2-0% to 265%, Tho fact that only one burrow was used

om one night (2:0%), indicates that all the handicouts present did not forage every

night, but spent at least one underground. We recorded overnight minimum tem-

perature during the period, but could not deduce any relationship between it and

percentage of burrows used,

When considering the number of times each indinidiual burrow was used, it is

convenient to take only those hurraws scored for all eight nights. The frequency

élistribution of these 35 burrows showing the mumber of nights each of thein was

used is shown in Fig, 6.

Fourteen burrows (40%) were not used on any night. These burrows did not

appear to he distributed at random oyer the area, but a statistical test is impracti-

cal because of the small numbers involved: They tended to be on or near the east-

ern edge, where habitats with spinifex predominated, From Fig. 6, we can also

\2 Dy H. SMYTIT ann ©, My PHILPOUT

note that more burrows were used on two cecasions than on one. This can be

explained when we consider that a bandicvot sponding # day in a burrow would

use it on entering one night and leaving on a subsequent night, It would be

scored both times.

We know that there were at least two female bandicoots in the study area,

because we captured two from burrows about 1,400 feet (425 m) apart on the

western edge of the area, This was within two days of completing the observa-

tions on activity. Outside the study area, the nearest locality we found where

rabbit bandicoots were living was about 3 miles (5 km) away. Even assuming no

migration to or from the study area, it is impossible to estimate the total sumber

af handicoots present.

Recause there were no signs of activity beyond 600 feet (180 m) from the

nearest hurrow, we may assume that for both female bandicouts captured, their

home range was completely covered by the study area.

(f) Notes on -caplite specimens

The twe adult females we caught in the study area did not have pouch young

whon taken in mid-December, nor when examined 2 month later.

When kept unavoidably at an air temperature: of 100° F (39° C) for a short

period, they lapped the water which was supplied. There was no permanent water

available in or near the study arca, and it is milikely that bandicoots experience

such high air temperatures in the wild. At these temperatures. they slept on their

sides, with tail, hind-feet, and head all extended. However, at lower temperatures

they slept crouched ou all four feet, with the nose tucked under the chest. the tail

curled around the body, and the ears folded forward over the eyes,

‘The captives started to eat chopped, tinned beef and to drink milk about #

week after being taken into captivity. Later, they ate minced fresh meat und live

mealworns.

DISCUSSION

Where ony study overlaps. with those of other recent authors, the Beucral

results agree.

Marlow (personal communication) caught two rabbit bandicoats in an area

of similar habitat on April 5 and 6, 1965. At latitiide 20°49 S, and longitude

130°195 E., (about 25 miles (40 km) NW. of The Granites, N.T.), the bandicouts

were living in an area of spinifex-Melaleuca scrub, wilh sandy soil and numerous

termite mounds. One of the bandicoots, a female, is preserved in the Australian

Museum, registered number M8620.

A brief survey of the distribution of rabbit bandicoots neur Warburton Mis-

sion indicated thit they lived in small isolated groups over a large area. Newsome

(1962) says he found that rabbit bandicoots were living in at least four and per-

haps six localities in a rectangular area 50 miles long by 30 miles wide (80 km hy

‘50 km) in Central Australia. It is probable that rabbit bandicoots utilize only a

small percentage of the large homogeneous areas which could support them.

Tnsect larvae and small vertebrates have both heen the most frequently noted

types of food of the rabbit bandicoot, ¢.. Marlow (1962), Finlayson (1933)

found rodent fur in the gut of a closely related species, Macrotis minor (Spencer).

Although house mice, Mus musculus Linne, were present in the study area, their

fur did not appear in any of 50 pellets cursorily examined, Aboriginal reports

clearly indicated that the rabbit bandicoot eats the large witchetty grubs, (larvae

FLELD STUDY OF THE RABBIT BANDICOOT \4

of buprestid beetles}, when they are present in the roots of the witchetty lush,

Acacia kempeena, Gould (1645) made a similar observation in South-Westeru

Australia. At the time of our study, witchetty grubs were absent from the sludy

area, although ahoriginals elaim that they had been plentiful in former times.

The food of the rubbit bandicwot reported here is remarkably similar to that

of the marsupial, Myrmecobius fasciatus Waterhouse, (banded ant-eater), which

Calaby (1960) studied in South-Western Australia, He fommd that termites of

many species, all different from those recorded above, were the principal food,

and that remains of ants were also often Found in faeccs. Likewise, he argued that

ants were ingested incideatally while termites were being caten. However, he

fonnd no traves of Lepidopteran or other insect larvae,

There is little concrete data on the breeding of the rabbit bandicoct, Woaod-

Jones (1924) comments that breeding of Macrotis in Central Australia is regu-

lated by rains and abundance of food, an observation made om many species

which live ina climate without regalar seasonal variation. Two other species of

bandicoots haye heen shown to breed throughout the year, Lyne (1964) working

with Perameles nasuta Geoffroy, and Mackerras and Smith (1960) with Isoedon

macrourus (Gould), have heth recorded breeding during all seasons.

Observations of activity of the rabbit bandicoot are limited, but Wood-Jones

(1924) has noted that captives nsually emerged about an hour after dusk, Stodart

(1966) observed the long-nosed bandicoot, Perameles nusuta Geoffroy in a M-acre

enclosure. She found that they usually emerged from their vests at dusk, and most

of their activity was completed after about two hours. They spent nearly all of

this Lime searching for and,

Although we found that a group of rabbit bandicoots lived in a small propor-

tion of an area with the potential ta suppert them, we have no evidence concern-

ing territorial behaviour within this small area. Two male and four femule long-

nosed handicoots in am enclosure did not have territories (Stodart, 1966) and

were solitary feeders, with intentional contact between individuals restricted to

actual inating.

The rabbit banchicaot was moderately plentiful and widespread in temperate

and arid Australia during the last ceritury. owever, a rapid decrease in range

andl numbers has been the pattern im recent years, especially in Lhe more termper-

ate regions. It has not been recorded frown New South Wales since 1912 ( Marlow,

1988.) and from Victoria since at least 1866 ( Brazenor, 1950). In 1934, tt was said to

he widely distributed in Western Australia south of the Kimberley region (Glaucrt,

1934), but twenty years Jater he reported that there were “no reports of its pres-

euece anywhere (in Western Australia) within recent years”, (Glanert, 1954), In

arid South.and Central Australia, Finlayson (1961) claimed that it was rapidly

bein reduced loa rare form, but Newsome (1962) indicated that it had recently

been recorded from 36 loculities in the Northern Territory. Rabhit bandicnats

were ttken near Birdsville i South-Western Queensland during 1957 ta 1954

(Mack, 1961). but while in the area recently; we obtained good evidence that the

bandicoots disappeared there about five years later. Thus, a steady reduction 10

rattwe towards parts of arid Central Australia seems to be taking place, with no

evidence that this reduction is slowing down or stopping,

An attempt to conserve some natural populations of rabbit bandicants 14

probably necessary now if they are to continuc as one of Australia’s naturally

occurring species. Suecessfal canservation of a species depends largely on.a know-

ledge of its distribution and eeolugy. We hope our study will be useful to those

who plan to conserye the rabhit bandicoot in its natural habitat.

14 D. R, SMYTH ann C. M. PHULPOTT

ACKNOWLEDGEMENTS

This work was supported by a Nuffield Foundation Research Grant to the

Royal Zoological Society of South Australia, and was administered by the Society's

President, Dr. P. S$, Watts.

We wish to thank Professor IE. G. Andrewartha of the Department of Zoology,

University of Adclaide, for encouragement, guidance and for many useful discus-

sions. The Council and Staff of the Royal Zoological Society of S.A. gave us

unlimited help. We received excellent co-operation from the Western Australian

Fisheries Department, and Mr, A. J. Fraser, Chief Warden of Fauna. Finally, we

wish to thank the Western Australian Department of Native Welfare; and the

supeciteadent, stall and people at the Warburton Ranges Mission for generous

help.

REFERENCES,

Atlas af Australian Resources (1959), (Department of National Development: Canberra. )

Brazenon, C. W, (1950). “The Mainmals of Victoria”. (Brown, Prior, Anderson; Melbourne. }

Carapy, J, H. (1960), Observations on the banded ant-eater, Myrmecobius f, fasciatus Water-

Iynase (Marsupialia). with particular reference to its food habits. Proc. Zoul. Soc. Lond.

138; 183-207.

Carapy, J. A. (1963). Australia’s threatened mammals, Wildlife, Brisbane J: No. 1. 135-8.

Doucras, W. H, (1964), An introdnetion to the Western Desert Language. Oceania Linguistic

Monogr, No. 4 (revised). (University of Sydney: Sydney.)

Finnaysox, H, HL. (1935). On mammals from the Lake Eyré Basin: Part 11, The Peramelicdae,

Trans. RB. Soc, S, Aust. 59: 227-36,

Wincayson, FL. H. (1961). On Central Australian Mammals: Parl IV, The distribution and

status nf Central Australian species, Ree, $, Aust. Mus. 14: 141-91.

Gravenr. L. (1934), The distribution of inarsupials in Western Australian. J. Proe, R. Soc,

West, Aust. 19; 17-32.

Guavert, L. (1954). The recent increase of the rarer native mammals. If Records fron the

W.A, Museunt. W, Aust. Nat, 4: 129-82.

Goun, J. (1845). “The Mammals of Australia, Vol, 1° (The Author: London.)

Lyxz, A, C (1964), Observations on the breeding and growth of the mursupial Perameles

nasuta Geolltroy. with notes on other bandienots, Alist, J, Zool. 12; 322-39,

Mack, G, (1961). Mammals from South-Western Queensland. Mem. Od. Mus. 13: 213-29.

Macxerras, M, Josepnink, and Sacre, Ruta Uf, (1960), Breeding the short-nozed bandi-

coat, Isoodon macrourus (Gould). in captivity. Aust. J. Zool. 8: 371-82.

Mannow, B, J, (1958), A survey of the marsupials of New South Wales. C.S.1.R.0. Wildl.

Res. 3: TLld4

Mantow. B. J, (1962). “Marsupials of Australia.” (Jacaranda; ‘Brishanw.)

Newsome, A. E. (1962), Rabbit-cared. bandicoots or bilbies.. Aust. Nut. fist. I4: 7-8.

Parteotrr, CG. M., and Ssavru, D. R, (1967), A contribution to our knowledge of some natiye

mammals fram inland Australia, Trens, RA. Soe. S. Aust. 91: 115-34,

Stoparr, BE, (1966). Management and behaviour of breeding groups vf the marsupial

Perameles nasuta Geoffroy in captivity. Aust: J, Zool. 14: 611-23,

Weae Tones F. (1924), “The Mammal; of South Australia, Part IL” (Gavernment Printer:

Adelaide. }

D. KR. Smevier and C, M. Prowreorr

PLATE |

lig, 1. An immature female rabbit bandicoot from 120

miles east of the study area, Approx.

0-28 x natural size.

Vig, Cylindrical bandicoot scratching, with faecal pellets on the top edge. The measure is 1

foot (30 cm) long.

io]

Fig. 3, Conical bandicoot scratching under Eragrostis laniflora, The measure is 2 in. x UU in.,

(5-0 em x 8-5 em).

PLATE 2 D. BR. Ssivru and C. M, Pintrorr

Fs poe =) <=", =

esi aril

Piare 2

Fig. 1. Rabbit bandicoot faeces, showing groups of pellets voided together. The seale is in

inches and centimetres,

Fig. 2. Bandicoot burrow near a fallen mulga log.

Fig. 3, Bandicoot burrow with tailmarks on the mound of soil at the entrance, The measure is

1 foot (30 em) long.

D. R. Smeyvrn and C. M. Prirrorr PLATE 3

PLATE 3

Fig. 1. Open mulga serub, with a burrow at the base of « mulga in the foreground. This

vegetation type contained only 2 burrows.

Mulga-tussock scrub, with a burrow partially hidden by tussock grass in the fore-

ground.

3. Mulga-spinifex scrub, with bandicoot scratchings in the foreground.

Fig. 4. Open spinifex glade, showing low rocky hills in the background. There were only 2

burrows in this vegetation type.

“Tj

AN ANALYSIS OF VEGETATION ON STRANDED COASTAL DUNE

RANGES BETWEEN ROBE AND NARACOORTE, SOUTH AUSTRALIA

BY R. M. E. WELBOURN AND R. T. LANGE

Summary

The presence or absence of 80 species has been scored in 330 samples of sclerophyll forest on nine

dune ranges, and the data classified by association analysis.

The vegetation is a mosaic of groups, within which four main groups may be distinguished. These

roughly correspond to orthodox vegetation societies, lint key species are determined solely by

measured association with other species, without reference to physical prominence. For example,

the species with the highest degree of association in this study, Phyllota pleurandroides, is a small

shrub.

The potential value of the groups in vegetation mapping is indicated.

AN ANALYSIS OF VEGETATION ON STRANDED COASTAL DUNE

RANGES BETWEEN ROBE AND NARACOORTE, SOUTH AUSTRALIA

by R. M. KE. Wetvourn and BR. T, Lanon

[Read 9 November, 1967]

SUMMARY

The presence or absence of 80 species his been scored in S30 samples of

sclerophyll forest on nine dune ranges, and the data classified by association-

anilysis,

The veyetalion is a mosaic vt groups, within which four main groups may

he distinguished. These roughly correspond to orthodox vegetation societies, hut

key specics are determined solely by measured association with other species,

without reference to physical prommence. For example, the species with the

highest degree of association in this study, Phyllota pleurandroides, is 2 small

shrub,

The potential value vf the yroups in veyetation mapping is indicated,

INTRODUCTION

The purpose of this study was to show by measurement, the floristic composi-

tion of the dune range vegetation and to discuss factors controlling its disposilion.

A discussion of analytical methods used has been published clsewhere (Welbourn

and Lange, 1967), The study was an M.Sc, project with the University of

Adelaide, to whom, with the State Herbarium of South Australia, grateful acknow-

ledgment is made.

The habitats are distinct ranges considered to he successively younger toward

the present coast. In the area studied there are at least twelve ranges subparallel

to the. coast, about 2 km wide by 30 m high above the otherwise flat AR

(PI. 1). Each range consists of two portions. The core is more or less consolidated

caleareous beach sand, a rele of coastal dunes formed at various stages of the

Pleistocene from about 600 to 200 thousand years B.P. (Sprigg, 1952). This

material outcrops westward as acolianite limestone, parcnt material of the terra

rossa soil carrying open woodland. Oycrlying this core are siliccous sands, of

uncertain origin but considered to be windsorted, leached and of Recent age

(Blackburn ez al,, 1965), These sands form the bulk of each range, and are the

parent material of the podsol which supports the predominant vegetation. This

is largely the Eucalypius baxteri association within the dry selerophyl!l forest

formation (Crocker, 1944).

Pattern, the degree of non-randommness in the spatial distribution of indivi-

duals, was the vegetation feature inyestigated. Since pattern reflects habitat varia-

tion, the patterns of ecologically similar species tend to coincide, or associate, thus

indicating a vegetation group. ‘he variable commonly measured to reveal such

groups is species frequency, the proportion of sample quadrats occupied. Associa-

tins were determined with such data in this study.

Trans, Roy. Soc, 5.A, (1968), Vol. 92,

20 R, M, E. WELBOURN anp R. T. LANCE

330

+59 Phyllota plevrandroidas “27

*63 Mongkoca scoparia -208

ql. 120 * 107 Isopegon ceratophyllus “101 0

if 6

2 be

2 of

g i

9 5

a z

< GO

ie]

Fig. 2.

VEGETATION OF STRANDED COASTAL DUNES 21

METHODS

The arew bounded by Kingston, Naracoorte, Penola and Rohe was selected

for study because the dune ranges are most distinct here ( Fig. 1). Sampling areas

were located within the remaining estimated 750 sq. kin of vegetation so that the

variation of mean annual rainfall, about 200 mm, would be represented on each

range. Most of the vegetation was unnatural in some way, so that, haying

excluded areas obviousty disturbed, or burnt less than four years before, or

dominated by bracken, 33 areas remained. At each of these, ten 20 sq.m circular

samples were located randomly where feasible, otherwise systematically. All

species likely to occur in upland sites in the area studied were peat also

Banksia marginata and Calytrix tetragonda were divided into ad hoc torms to

render the data more sensitive to habitat variation. A reference collection of

relevant species was used throughout,

The method selected to reveal vegetation groups was association-analysis as

proposed by Williams and Lambert (1959), Them recommended association index

is for cach apecies the sum of all the chi-squared. values obtained from contin-

geney testing with each other species in turn, The procedure is to subdivide the

330 samples firstly into two groups, respectively with and without the appropriate

kev species, The key species is that with the highest association index at any stage

of subdivision, Within each of the groups thus formed, the process is repeated,

tractionating the original single group of 330 samples into progressively more sub-

slivisions, In this study, only species with a frequency greater than 1% were consid-

ered; data were computed clectronically, and subdivision was stopped at 16

SOUPS,

THE VEGETATION

‘Lhe analysis: ii Uhese terms reveals a serics of groups of deereasing

importance, conveniently represented as a hierarchy (Fig, 2). The validity of any

gcuup asa recognizable vegetation community is indicated by the range of associ-

alion index over which it persists undivided; this is supported by its containing

relatively few minor groups. For example, Group B subdivided on Monotnca is

such a community, since it persists fram Index 155 to Index 20, and embraces only

two minor groups as far as the data were analysed,

On this basis four such groups may he distinguished from the complex mosaic

of groups of yarying validity whicl: comprise the samples studied. These groups,

A to D, exsist simultancously over a larger range, 125 to 60, than any other number

of groups. Thus within the limitations of the analysis, they are communities

approximating societies within the Eucalyptus baxteri association, They are identi-

fied by 16 indicator species which vary in frequency between groups (Table 1),

For example, vegetation containing Monotaca, Persoonia and Acacia spinesevns,

birt nat Fudtonden, would be either Group A or B. lf furthermore it contained

Phyllota, Xanthorrhoea and Boronia, but not Ac. oxycedrus or Styphelia, it

would he Group A.

On the other hand, the remaining 64 species analysed are relatively insensi-

tive to the discontinuities that affect the key species, Of these, 22 have a fre-

vency greater than 30% (Table 2), their similar group frequencies indicating

their unbiased cistribntion throughout the area studied. The appearance of the

vegetation is dominated by the five species of frequency greater than 75%, vis, the

tree E. barteri, the small shrubs L. myrsinoides and X. australis, and the wider-

shrubs ff. sericea and L. virgatus. Superimposed upon this picture of four groups

R, M. E. WELBOURN ano R. T. LANGE

TABLE 1

INDICATOR, SPECIES

Species used to identify the groups; relatively high frequencies are in bold faced type.

| Frequency %%,

Species Overall | - - '

A B C dD

Monoloce scopuria (Sm.) R.Br. 22 4 100 0 0

Persoonia juniperina Labill. 8 17 21 5 i)

Aciteia spinescens Benth. 2 10 2 0 0

Pultenaea prostrata Benth, ex Hook. f. ) 3 0 13 8

Phyllota pleterandratdea Fo ov. M- i 18 i 100 0) ) "

Xanihorrhoen quadrangulota FL v. M. 6 | 82 0 {) (

Roroniu coerulescens F, v. M. 3 14 0 1 4)

Hibbertia virgata R, Br, ea DC. var, crassifolia

(Benth,) Black 4 5 27 3 () 0)

Lhotskya alpestris (Lindl.) Druce 7 25 0 2 tt

Acacia oaycedtrus Sieb. ex DO. 2 0 ll 0 0

Thomeasia petalocalys Ky. M- 15 29 2 15 14

Styphelin adacendens R. Br. 5 9 16 2 a

Bucalyplus obliqua L' Herit, @ 8 3 28 4

Tsupayon ceratophyllus R.Br. , 57 77 68 100 0

Dodonaea viscoza Jacq. 4 3 0 1 IL

Leucopogon collinus (Labill.) B.Br-. Nae ; 49 30 22 a

TABLE 2

COMMON SPECIES

Species of ovorall frequency greater than 30°, ; frequencies greater than 73%, are in bold faced type.

Frequency %,

Species Overall |-- :

A | 2B co) 6D

MYRTACEAE

Eucalyptus baxter: (Benth.) Maiden et | j

Blakely ex Black 89 55 i 73 74

Leptospermuin myrsinoties Schldl. 89 7 ts 4 7)

Calytrix tetragona Labill, (glabrous form) 30. 16 AG 28 a4

EPACRIDACEAK |

TLeucopogon virgatus (Labill.) R.Br. 80 61 86 82 44

Lstrolumu conoustephiailes (Sond.) Pov. M.

ee Bonth, | 74 TW ta) 70 ; 74

Brachiloma eiliatum Bonth, Gs 42 70 65 65

Leucopoyon erieoides (Sm_.) R.Br. 52 mo, 60 45 7H

Epacris impressa Labill. 5 47 | 4 58 BL

Aeratriche serrutata (Labill.) R.Br. +7 a 52 GL au

Astrolomn huminisum (Cav.) Ridr. 34 | 1 13 38 | fl

A, humifusum vor. dentieulatum (R,Br,) Black 31 34 2h 23 44

PROTEACEAR

Banksia ornata ¥. v. M. ex Meisn. AL 8&6 tb G4 BY

Isopogon cerataphyllus R.Br. (see Tably J) BT fox x x x

Banksta marginata Cay. (shrub form) ; 49 63 57 57 25

Bunksiu marginaia Cav. (tree form) a4 1 a2 55 47

LEGUMINOSAE j

Acnvin myrtifolia (Sm) Wild. 30 4] A 26 13

Kennedia prostrate R.Br. ex Ait. 30 20 35 ab 2a

OTHER FAMILIES | |

Hibbertia sericea (R.Br. ex DC.) Benth. §4 80 860002 2 RS

Nanthorrhoes musiralis R.Br. 80 Th sb 3 jeetil

Hibbertiu siricie (DC.) F. vy. M. 62 83 54 7 | Ad

Hypoluena fustigiata R.Br. 60 78 81 62 35:

Tetratheca elfiute Lindl. 48 25 | 7 55 as

Correa reflext (Labill.) Vont. var, refleace > 40 38 62 | 4g 18

VEGETATION OF STRANDED COASTAL DUNES BA

contrasted with common widespread species, aré 42 ypectes with similar frequency

to that of the Indicators, 2 to 29%, but occurring haphazardly in the cronps and

thus distributed unpredictably throughout the vegetation. Superficially, then, the

dey sclerophyll forest studied appears to comprise au open tree canopy oyer a

layer of small shruhs dominated by relatively few species. The overall impression

is of uniformity. [Mowever measurement revéals that, of the numerous Jow fre-

ucney species, some cunsistently associate in socictios or groups.

Since it is difficult to assess the validity of the groups other than by field

experience, it is necessary tu state at least seme sources of crror inherent in the

methods used, Firstly. there ure the hasic errors due to a sampling intensily ot

iubout 0-001%, and to operator fallibility in naming and scoring species. Secondly,

data based on frequency are liable to misrepresentation because quadrat size

affects the results, For example, a large quadrut overemphasizes uncommon

species, whilst a small quadrat underemphasizes the importance of species of

patchy distribution. This defect can be overcome only with additional data on

variables such as density, Thirdly, the analytical method was arbitrarily selected

from several which might have been used (Welbourn and Lange, 1967). Its mast

serious fault is that in subdividing on a single association index, this method does

not dicate the extent or nature of any subordinate association which may exist.

That is, there is no indication of the overall confidence with which a particular

subdivision is made, Furthermore the assuciation index itself falls short of thearct-

ical excellence. Finally, it should be recognized that any such method imposes

definite cut-off points to groups hetween which there is some continuity. Never-

theless such groups provide at least an abjective basis upon which to classify and

mup vegetation variation,

Fig, 3 is such a anap which reveals that the groups themselves are move or

less geographically restricted. For example, Croup A tends io the north-central

ares, B to the south-east, C to the west and south, and D to the cast and north.

Sinee these appear not to be chance distributions, there are likely to be enviran=

mental factors correlated with them. Several factors will be discussed, to illustrate

the sort of hypothesis which may arise from such vegetation mapping.

If time was the only factor involved, some groups could be expected to repre-

sent stages in a succession, and so to predominate on ranges of similar age. Such

trends are apparent; for example, between Naracoorte and Robe, Group D is

gradually replaced firstly by A and B, then by C on the younger ranges, However,

this evidence rests on the assumption that all other environmental factors aré held

constant. This was not so [we the case of soil and topography which were tore

Variable than expected, Thus the correlation of vegetation with habitat age canal

be tested under the sampling conditions of this study. Similar remarks apply to 4

correlation with aspects of climate such as coastal influence and annual rainfall,

From field descriptions of the sampling areas, it is apparent that variations in

soil and topogriphy are correlated to same degree sith vegetation variations, For

example Group A pceurs on sand ridges normal to the ranges, suggesting an

immature profile; the Fucelyntis baxteri here is stunted, and E. diversifolia

Bonpl, was observed nearby suggesting a gradation to the solodized solonctz soil

common to the north, Crewp B occurs on deep sands in the higher rainfall area:

the vewetation is profuse and undisturbed. Group C iy a diversified group which

tends to occur in flat, shallow areas, particularly to westward where the ranges

appear to have less siliceous sand covering the limestone. The next subdivision of

this group (Vig: 2) is on E, obligua, a species known to ovcur on shallower soils.

Group D occurs on the eastern sites of ranges well away from limestone, but with)

some seasonal watcrtable influence,

24 R. M. E, WELBOURN ann R, T. LANCE

Fig. 3.

Clearly, profile measurements and more intensive sampling are required to

substantiate any correlation of vegetation with soil and topography. Such correla-

tion, once established, will be of value in habitat classification and mapping.

REFERENCES

Biacksurn, G., Bonn, R. D, and Crarxe, A, L. P., 1965. Soil development associated with

stranded beach ridges in South-East South Australia. C.S.1.R.O. Aust. Soil. Publ. No, 22.

Crocker, R, L., 1944. Soil and vegetation relationships in the Lower South-Kast of South

Austrulia, A study in ecology. Trans. R. Soc. S. Aust., 68, pp. 144-172.

Sprica. R. C., 1952. The geology of the South-East Province, South Australia, with special

reference to Quaternary coastline inigrations and modern beach develoy:nents. Geol. Surv.

S. Aust, Bull. No. 29.

We tsourn, R. M. E. and Lancs, R. 'T., 1967, Subdividing vegetation on inter-specifie associa-

tion. Vegetatio Acta Ceobotanica, 15, pp. 129-136.

Witirams, W, T. and Lampert, J, M,, 1959. Multivariate methods in plant ecology. 1.

Association-analysis in plant communities. J. Ecol., 47, pp. 83-901.

R. M. I. Weteourn and R. T. LANGE PLATE |

PLATE 1

General view of the area studied, looking eastward towards Naracoorte, Three ranues. each

partly cleared of natural vegetation, are emphasized in the foreground and middle distance

with a line along their crests.

FURTHER TAXONOMIC NOTES ON THE SPECIES OF MILLOTIA

CASSINI (ASTERACEAE)

BY RICHARD SCHODDE*

Summary

One new species, Millotia inopinata, is described and chromosome numbers for it,

M. myosotidifolia (Benth.) Steetz, and M. tenuifolia Cass. are reported. Attention is drawn to the

reported tolerance to and accumulation of copper in M. myosotidifolia.

FURTHER TAXONOMIC NOTES ON THE SPECIES OF MILLOTIA

CASSINI (ASTERACEAE)

hy Ricwanp Scuoppe*

{Read 9 November, 1967]

SUMMARY

Gne new species, Millotia inopinate, is described and chromosome: rimbers

for it, AL. myosotidifolia (Beuth,) Steetz, and Af, tenuifolia Cass. are reported,

Attention is. drawn to the reported tolerance to and accumulation of copper in

M. miyosotidifolia.

Since the publication of my taxonomic account of the genus Millotia Cuss.

(Schodde, 1963), further material and information have become available. The

ulbbreviations AD, CANE, K, MEL, PERTH used im the text refer respectively to

the State Terbarinm of South Australia, Adelaide, South Australia; the herharium.

C.8.0H.0., Canberra, A.C.T.; the Herbarium, Royal Botanic Gardens, Kew,

England; the National Ierbarium, Royal Botanic Gardens, Melbourne, Australia;

and the Wester Australian Herbarium, Perth, Western Australia.

{, Millotia inopinata Schodde, sp. nov.—Fig. 1

M. myosotidifolia (Benth. ) Steetz proxima, cujus invelucrum, formam floscu-

lorum, et cypselas plerumque strigulosas habet, sed habitu latiore, foliis + asguste

lineati-oblanceatis vel puene fliformibus, capitulis + depressis globosis vel ali-

quantum turbinatis, bracteis involicralibus ad apices plerumdque ratundatis,

corollis aureo-flavis, cypselis aliquantum longioribus in rostris deflexis, papillis

cypselarum ad apices integris et + patentibus vix strigosis in rostris cypselarum,

et pappi setis 5-8 grosse barbellatis vel sub-phimosis differt,

flomer heads. Leaves relatively sparse and appearing confined towards stem hases

inear or linear-oblanceate to ulmost

* Division of Land Research, C.SJA.G.. Canberre, A.CT,

Frans. Roy. Soc. S.A. (1968), Val, 92.

SCHODDE

ant; B. mature Hower head; C, involucral bract; D, forets with maturing

A, whole pl

> and E, whole style and androecium.

ig. L,

psel:

TAXONOMIC NOTES MILLOTIA 20

Corollas rather natrow infundibuliform, 24-34 (— 34) mm Jong, het below the

throat and deflexed over involucre when mature (central corollas as detlexed as

ar somewhat more erect than peripheral corollas), bright golden vellow with the

tube becoming brownish after anthesis; corolla limbs of 3 (exceptionally 4)

spreading acute lobes $-3 mm long, Stanens 5, with apices becoming exserted to

or a little beyond rims of corolla limb; anthers with oblong thecae 3-1 mm lon,

the cannective tins extended }-3 mm beyond thecae, Style branches 4-1) mm

long, dilated broadly at the apices with conoidal acute appendages, Cypselne

lineyr, 6-9 (—10) mm Jong, sharply deflexed from near base of beaks over in-

volucre and protruding 3-4 (—5) mm beyond at maturity, variously dark hrawn,

sparsely hispidulous-strigillose; cypsela beaks (2k —) 3-5 mm Jong at maturity,

compressed and most broadly so on area of deflexion; cypsela papillae = terete

lo narrow cylindrical clavate, + 4 mm long, entire rather expanded chtuse ul

the apices, colourless more-or-less treuspurent, somewhat appressed and diffuse

on body of cypsela, rather spreading, denser, and confined to imuirgius of

compressed faces on beak. Pappi of 5-8 twisting more-or-less erect setaz + MX

as long as corollas, becoming sparsely coarse barbellate or semi-plumose towards

the apices. Chromosome number; n= 8 (fide i. L. Turner).

DISTRIBUTION AND ECOLOGY

Known presently from three collections: leg. A. S. George 7978: PERTH;

29 miles west of Mt. Magnet, comprising 12 plants, leg. ©. H, Giffins 1540:

PERTH, 7 miles south of Wannoo Roadhouse, comprising 7 plants, and the Lype

collection, of 14 plants, fram 82 miles north of Murchison river, All localities are

in the central west zegion of Western Australia,

According to Turner (pers. comm.), the species was looally common in the

type locality on coarse sandy soil in open areas with burned-over mulga (Acacia

sp.); George records il in sandy loam at a granite outcrop, Flowering: August-

September.

CHARACTERS AND AFFINITIES

M. inopinata is closely related to and sympatric with M. myosotidifolia. The

inerphological similarities lie in the long compressed beaks and cylindrical mostly

appressed papillae. of the cypselae, the form of the corolla, androvcium and style

branches, the narrowly midribbed bracts, and the wholly lanate indumentum.

The new species would be identified as M. myosolidifolia when my kev to the

species of Millotia (Schodde, 1963) is used but may he distinguished by the

following characters against which the contrasting characters of M. sayvsotidifolia

are bracketed: eypselae with heaks conspictiously ceflexed and papillae obtuse at

the upiccs (beaks erect and papillae minutely notched); corallas deep golden

yellow (creamy white rarely creamy yellow); pappus bristles 5-S, + ¢ as long as

corolla (bristles usually (15—) 18-25 (— 30), + as long as corolla, very rarely

fewer and shorter); mature flower heads depressed globose (cylindrical-clobose );

apices of involucral bracts broad rounded obtuse. often finely or obscurely fim-

briate, rarely wttenuated (caudate more-or-less entire, rarely to acuminate or

acute); leaves narraw linear-oblanceate to almost filiform (4 —) 1-2 (—3) em long

* (KM —) 41 (—1%) mm broad ( narrow to broad oblanceate or occasionally some-

what spathulate (%—) 2-4 (—G@) cm long * (1—) 2-5 (—8) mm broad); habit

broadly ascending (ascending to erect),

The chromosome numbers so tar recorded. n = § in M, inop/nala aud n = 10

dr 11 in M. myasotidifolia (see below), also uppear to be consistent with differ-

ences between the two species. Additional counts from the localised populations

HI Kh. SCMODDE

of M, inopinate and from the eastern Australian populations of MM. myosotidifulia

are needed, however, to determine how far these numbers are characteristic at

the species,

It is noteworthy that a number of the characters distinguishing M. inopinata

from M, myosotidifolia resemble those of the eremaean M. greevesii F, Muell.,

in purticular the habit, shape of leat and Hower head, depauperate pappus, and

corolla colour, This may be a case of parallelism in so far as M, inopinata occurs

towards the north-west and arid margins of the geographic range of M. myosotidi-

folia, There appear to he similar convergences in the characters distinguishing

M, macrocarpa Schodde from its nearest relative, M, tenuifolia Cass,

There are also similarities between the new species and M. depauperata

Stapf, which was reduced to M, myosotidifolia (Schodde, 1963), and to evaluate

them Professor B. L. Turner (pers. comm,) has examined the type of M. ¢de-

paliperate. This specimen, though like M. inopinata rather than M. myosotidifolia

in its few bristled pappus, apparently obtuse-tipped cypsela papillac, and broad-

Hpped involucral bracts, is apparently too young and depanperate in its various

characters to permit certain identification. Turner considers that in gencral habit

it resembles M, myosotidifolia and that its cypsela papillae and involueral bracts.

thingl slightly different from much of the material of M. myosotidifolia lw

examined, do mately certain determined collections of that species well enough,

ww Pritzel 545. In the present circumstances, there seems to he Jittle point in

treating, M. depenperata as anything else than a synonym of M. myoasoticdifolia-

2, Millotia myosotidifolia (Benth,) Steetz

Chromosome nuinber: w= 10 or 11, reported by B. L. Turner. Vouchers:

Turner 5283, 13 miles north of Norseman (1— ca lO); MEL; Turner 5442, 7

miles south of Three Springs (n = 10): MEL; Turner 5482, 30 miles south of

Miawlirah (= 11); MEL, All localities are in sonthern Western Australia,

Glissett (1966) has recorded tolerance to und accumulation of copper in

plants of this species found growing exclusively and abundantly on strips of open

ground directly below old copper-bearing dumps from the Ukaparinga coppe1

mine near Williamstown, South Australia. Spectrographie analysis of sample

collections showed copper contents of approximately 40,000 p.pan. in the ash

of gree plants and approximately 4% in the ash of dead plants, Voucher:

|. Moisseeff su, Ukaparinga Coppermine, L miles cast of Williamstown:

AD96647104.

It is of Interest to mention that the species has also been observed growing

abundantly on the rocky dumps from the old Blinman copper mine, at Blinman,

South Australia. Vouchers: D, N. Kraehenbuehl 18, rocky hillslopes of the old

Blinman Copper Mine at the north end of Blinman; AD 95909049; R. Schodede

997, hillslopes of the old Blinman Mine at the sorth end of Blinman: ADSS9080R4,

3, Millotia macrocarpa Schadde

The shape of the corolla is more accurately described as yery narrow infiruli-

buliform ef, the original description of the species in which the term “narrow

cyathiform™ was intended to apply to the throat only, not the whole corolla.

4. Millotia fenuifolia Cass.

Chromosome ntunber; 1 = 13, reported hy B. L. Turner. Vouchers: Turner

5271.5 miles north of Norseman: MEL, Turner 5315, 14 miles west of Southern

Cross: MEL: Turner 5516, 53 miles north of Albany; MEL. The counts ure all

from yr. tenvifolia and the localities are all in southern Western Australia,

TAXONOMIC NOTES MILLOTIA 31

My attention has also been drawn to a recently collected specimen of this

species from near Collie, in southern Western Australia (leg. P. G. Wilson 3750:

CANB 161828) which is unusual in possessing predominantly 5-lobed corollas,

and short laevigate cypsclae 3-4 mm long at maturity, only 3-2 as long as the

involucre, and with short beaks 4-1 mm long. It extends the known variation of

the species in these characters.

ACKNOWLEDGEMENTS

Thanks are duc to Professor B. L, Turner, University of Texas, Austin, Texas,

U.S.A., and Mr. P. G. Wilson for their collaboration, particularly to Professor

Turner for making available the chromosome counts and checking details of

Millotia inopinata against the type of M. depauperata at Kew. Professor Turner’s

investigations were assisted by a National Science Foundation Grant GBI1216.

REFERENCES

Burssetr, A. H., 1966. Copper Tolerant Plants from the Ukavaringa Copper Mine, Williams-

town. Quart. Geol. Notes no. 18, pp, 1-3 inelL£., publ. Geol, Surv, S. Austral., Dept. of

Mines, South Australia,

ScHopve, R., 1963. A Taxonomic Revision of the genus Millotia Cassini (Compositae), ‘Trans.

Roy. Soc. §, Austral. 87, 209-241,

A NEW SPECIES OF PTILOTUS FROM SOUTH AUSTRALIA

(Amaranthaceae )

By G. Bent, F.L.S.*

(Communicated by Hj. Eichler)

[Read 11 April, 1968]

SUMMARY

A. description and an illustration are given of a new species of Ptilotus,

Pt. symonii from South Australia. The type specimens are cited and some critical

notes are made on some characters of the new taxon which is compared with

Pt. seminudus (J. M. Black) J. M. Black and other species.

A recent examination of specimens of Ptilotus sent us from both the State

Herbarium of South Australia (AD) and Mr. D. E. Symon (Herbarium ADW)

revealed the existence of a hitherto unknown species which is here described as

follows:—

Ptilotus symonii Benl, sp, nov. (Fig. 1. a-c).

Planta perennis, pluricaulis, in statu iuvenili leviter tomentosa, demum

subglabra. Rhizoma lignosum adscendens, plusminusve tortuosum, fusiforme, in

speciminibus examinatis usque ad 25 cm longum, superne 16 mm crassum.

Caules 50 cm et ultra longi, 1-3 mm diametro (in basi lignosa 3,5 mm crassi),

virgati, saepius curyato-adscendentes, visu cinerei dein pallido-virides; iuveniles

teretiusculi per totam longitudinem pilis crispis crassiusculis nodulosis, ad 1,5 mm

longis dense, adulti sulcati sparse induti. Rami valde ramulosi, ramuli tenerrimi

subdensi, adscendentes, divaricati ve] patulo-erecti, ad 20 cm longi; basales 7-22

mm distantes, summi approximati, usque ad inflorescentiam dense foliati,

Folia permulta minora quin etiam minima, alterna interdum specie secunda

(Fig. a), 2-6 mm, raro ad 13 mm distantia, primo modice puberula denique

glabra et laete viridia, plusminusve coriacea, integerrima, siccitate marginibus

raro subsinuatis, nervo medio subtus vix prominente; omnia inferne attenuata,

breviter vel brevissime petiolata (petiolo indistincto ad 2 mm longo), in apice

mucronata (mucrone 0,1-0,2 mm longo); maiora 1-2 cm longa et 2-3 mm lata

oblongo-lanceolata, minora saepe acicularia.

Spicae haud amplae numerosae, hemisphaericae (1,5-2,5 cm diametro) vel

subovoideae (2,5 cm longae et 1,8 cm latae), pedunculatae, solitariae ramulos

terminantes, raro subsessiles laterales, rhachide breviusculo (0,7-1,3 cm longo)

dense villoso, pilis plusminusve flexuosis nodosis, 1-1,5 mm longis. Flores (15-25,

raro ultra) haud dense congesti, demum stramineo-flavescentes, albido-pilosi.

Bractea bracteolaeque acutae integerrimae, pilosae, extus pilis articulatis,

apicem attingentibus quin etiam paullulo excedentibus obsessae, uninervae, post

lapsum perianthii superstites, inaequales: Bractea inferior rigida, subcordato-

lanceolata 2,5-3 mm longa et 1-1,2 mm lata, fuscescens, nervum medium versus

obscura, in dorso omnino pilosa, pilis articulatis, circiter 1 mm longis densius

vestita. Bracteolae 2 distincte maiores subcarinatae, membranaceae, tenues,

ovato-lanceolatae 4-6 mm longae et 2 mm latae, tantum nervum medium fusces-

centem versus pilos circiter 1,5 mm longos gerentes, lateribus glabris generaliter

incoloratis, hyalinis, nitentibus, perianthio adpressae.

* Botanische Staatssammlung Miinchen, Germany.

Trans. Roy. Soc. S.A. (1968), Vol. 92.

34 G. BENL

Fig. 1. Ptilotus symonii Benl: Pedicelled spike (from AD 96131068) with the tiny upper

leaves (Fig. a); perianth (from AD 968020399) with stamens and staminal cup spread open,

inner view (Fig. b); pistil (Fig. c).

ae a

A NEW SPECIES OF PTILOTUS 35

Perianthium pentaphyllum elongato-erectum dein subcampanulato-patens,

basim constrictam cartilagineam versus sensim indurescens, tubum angustum

cylindraceum 0,8-1,2 mm longum, extus pubescentia plusminusve absconditum

formans. Tepala elongato-linearia vel lineari-oblonga, anguste hyalino-marginata,

trinervia—nervis lateralibus (superne saepius indistinctis) areolam medianam

impellucidam, incrassatam, coloratam includentibus—, ecarinata, integerrima,

truncata, apicibus inconspicuis, fere nudis 3-4 mm longis, rufescentia dein viridi-

flavescentia, pilis dorsalibus strictis, rectis, articulatis, in articulis breviter verti-

cillatis ad 9 mm (in ima basi circiter 1 mm) longis, imprimis dimidio inferiori

areolae medianae tepalorum orientibus, sed apicem superantibus adpresse

denseque obtecta, pilis marginalibus tenuioribus, brevioribus sparse ciliata, intus

demum albido-laevigata, inaequalia: 2 exteriora 9-11 mm longa et 0,6-0,9 mm

lata, marginibus (ca. 0,15 mm) in apicem paulo contractum, eroso-denticulatum,

pilis dorsalibus longe (ad 2 mm) superatum transeuntibus (Fig. b); 3 interiora

paullum breviora, sed angustiora (0,4-0,6 mm lata) et acutiora, marginibus

superne involutis, inferne pilis crispis, nodulosis, uni sive duobus Jateribus tepali,

praecipue autem margini tubi perianthii orientibus et introflexis, plusminusve

copiosis munita.

Stamina staminodiaque 5, in floribus examinatis semper 3 fertilia et 2

abortiva, basi modice (ad 0,2 mm) dilatata flamentorum applanatorum cupulam

membranaceam, hyalinam, tubo perianthii adnatam formantia, anulo minimo

(0,15-0,3 mm) libero integro, pseudostaminodiis nullis, filamentis ligulatis superne

subulatis, interdum brunnescentibus, circiter 2 mm longis, abortivis rudimentis

antherarum coronatis vel anantheris brevioribus (0,5-1,5 mm); antherae bilo-

culares, flavae, lato-ellipsoideae 0,3-0,4 mm longae et 0,2-0,25 mm latae, dorso

affixiae.

Ovarium subclavatum, conspicue stipitatum 2,5-3 mm longum (stipite circiter

1 mm longo incluso) et 0,7-1 mm latum; stylus sicut ovarium regulariter glaber-

rimum 1,3 mm longum et circiter 0,1 mm diametro, plusminusve excentricum;

stigma inconspicuum haud distincte capitellatum, papillosum.

Holotype of species—5 miles south of Koonalda Homestead (east of Eucla),

south-western region of South Australia; D, E. Symon, No. 4684, 21.11.1967, AD

No. 968020399.

Isotypes—Idem, A, ADW_. CANB, K, LE, M, TI, UC, W.

Further collection—Other material of this taxon had already been collected

by P. G. Wilson (No, 1635) in North West Plains, ca. 40 km East of the Western

Australian border, off Eyre Highway, 13.1X.1960 (AD 96131068), Our

description is based on Wilson’s specimens, too, which, therefore, may be regarded

as Paratypes.

Habitat—Symon’s plants were growing “in open Mallee scrub” and “mostly

found in the twiggy remains of dead or dying plants of Westringia rigida’.

Wilson’s material had been gathered in Acacia woodland.

Characteristics—The new taxon superficially approaches the South Australian

Pt, seminudus (J. M. Black) J. M. Black as regards the general form and colour

of the spikes. In this species, too, the outer perianth segments bear a truncate

and denticulate apex, and the inner tepals are distinctly exceeded by dorsal hairs.

The stems arising from a strong rhizome (see “Australian Plants” 4: 117, 1967)

are pubescent in about the same way, when young.

In Pt. seminudus, however, the stems and branches are constantly shorter and

thicker, the leaves (especially the radical ones) considerably larger, the spikes

richer, the bract and bracteoles nearly equally long and more acute; the bract

being usually less hairy. Moreover, the tepals are longer, the inner ones more

36 G, BENL

narrowed towards the apex, and the points of the outer segments are not so

markedly overtipped by the erect dorsal hairs.

In both species these hairs are articulate and of nearly the same length, but

in Pt. symonii they primarily rise from the lower half of the tepals thus covering

the perianth tube, while in Pt. seminudus the basal part of the segments looks

naked, revealing the hirsute subglobular tube. In this plant we find inflexed hairs

inside the perianth arising from the margins of the inner tepals above the tube,

whereas in Pt. symonii a woolly pubescence of the inner segments takes its origin

from the edge of the perianth tube, too.

In addition to these characters, numerous details of the reproductive organs

diverge: e.g. two stamens only are fertile in Pt. seminudus, its staminal cup shows

a comparatively higher free ring, and between two filaments much broadened at

their base you may find a small lobe, at times; the ovary being pilose in summit.

Pt. seminudus, therefore, differs markedly from our taxon.

As a striking particular feature of diagnostic importance are to be regarded

the numerous and uncommonly small narrow leaves densely borne along the

branches: and branchlets of Pt. symonii. Except for the bushy and extremely

branched Pt. parvifolius (F. v. Muell.) F. v. Muell., no other representative of

the genus is characterised by such leaflets. But these two plants are quite unlike

one another.

An additional trait of diagnostic interest is given by the relatively long and

strict dorsal hairs of the intense pubescence in the perianth, distinctly exceeding

and concealing the outer as well as the inner tepals, which look tapering because

of this. To a lesser extent, we find a similar appearance in Pt, arthrolasius F.

v, Muell., a subshrub with a yellowish pubescence, and still more in Pt. eriotrichus

(W. V. Fitzg. ex Ewart & White) W. V. Fitzg., another frutescent species with a

dense greyish tomentum. The shorter hairs in the small flowers of Pt. forrestii

F. v. Muell. and the longer ones in Pt. villosiflorus F. v. Muell. are less distinctly

articulate, and mostly lack the verticillate toothlets at the nodules, as is the case

in Pt. arthrolasius, too. The tepals of the narrow-spiked Pt. lanatus Cunn. ex

Mog. (including var. glabrobracteatus Benl) are surpassed by short bristly and

thickish hairs, those of the long-spiked P#, leptotrichus Ben] by a tuft of relatively

few articulate hairs. In Pt. albidus (C. A. Gardn.) Benl, Pt. brachyanthus (F.

vy. Muell. ex Benth.) F. v. Muell. and Pt. petiolatus L. Farmar the perianth is more

or less completely hidden among dorsal hairs forming an intricate wool. Each of

the cited species has a dissimilar appearance of its habit, of its leaves, spikes or

floral organs, and there is no doubt left as to the specific nature of our well

distinguishable taxon: Pt, symonii does not at all agree with any of the species

hitherto described.

Name—The plant is named in honour of Mr. David E. Symon, B.Ag.Sc.,

Botanist at the Waite Agricultural Research Institute. Mr. Symon is one of the

collectors of the new species, drew my attention to it, and supplied us with

sufficient material.

THE YELLOW-EYE MULLET

AGE STRUCTURE, GROWTH RATE AND SPAWNING CYCLE OF A

POPULATION OF YELLOW-EYE MULLET ALDRICHETTA FORSTERI

(CUV. AND VAL.) FROM THE COORONG LAGOON, SOUTH AUSTRALIA

BY J. A. HARRIS*

Summary

The "Coorong mullet" spawns once per year, from January to early April. Males begin maturing a

few months before the females are running ripe. Seven stages can be recognized in the gonads of

the female and measurements of the diameter of the ova in each stage are given. The smallest fish

with mature gonads were found to be 21 cm (males) and 23 cm (females), measuring from the tip of

the snout to the caudal fork.

The ages of the fish were determined from a study of otoliths and this method was supported by the

Petersen method of analysis of frequencies of lengths. The mean lengths attained by the "Coorong

mullet" during their first four years are 14, 21, 26 and 31 cm respectively.

Characteristics of the "Coorong mullet" are compared with both the eastern and western races of the

yellow-eye mullet (See Thomson, 1957a; 1957b). It is concluded that the "Coorong mullet” has the

characteristics of the eastern race.

THE YELLOW-EYE MULLET

AGE STRUCTURE, GROWTH RATE AND SPAWNING CYCLE OF A

POPULATION OF YELLOW-EYE MULLET ALDRICHETTA FORSTERI

(CUV. AND VAL.) FROM THE COORONG LAGOON, SOUTH AUSTRALIA

by J, A. Hanats*

[Read 9th May, 1968]

SUMMARY

The “Coorong mullet™ spawns onee per year, from January to early April.

Males begin maturing a few months before the females are running ripe, Seven

stages can he recognized in the gonads of the female and measurements of the

diameter of the oya in each stave gre given, The smallest fish with mature

gonads were found to he 21 em. (males) and 23 em (females), measuring from

the tip of the snout to the caudal fork,

The ages of the fish were determined frow a study of otoliths and this

micthocd was supported hy the Petersen method of analysis of frequencies of

lerigths, The mean lengths attained by the “Coorong mullet” during their first

four years are 14, 21, 26 and 31 em respectively.

Characteristics of the “Coorong mullet” are conypared with both the eastern

and western races of the yellow-eve inullet (See Thomson, 19574; L957b). It is

concluded that the “Coorong mullet” has the characteristics of the eastern race.

INTRODUCTION

The yellow-eye or fresh-water mullet Aldrichelta forsteri (Cuvier and Valen-

ciennes) occurs in coastal waters of all Australian States except Queensland.

Thomson (1957a) studied the yellow-cye mullet of Western Australia and com-

pared it with those of Victoria and Tasmania. However, very little is known about

the rate of growth and spawning cycle of the yellow-cye mullet of South Aus-

tralia, especially from the Coorong lagoon (see Fig, 1).

The yellow-eye mullet is very common in South Australia, particularly in

coastal brackish waters, It is the principal species of mullet sold commercially in

this State. The only records of the commercial fishery available during this study

were very rough estimates of the total weight of mullet sold in South Australian

markets from 1951-62:

1951-52 770,000 Yh 1957-58 560,000 Ib

1952-53 500,000. lb 1958-59 900,000 Ib

1953-54 500,000 Ib 1959-40 649,405 Ib

1954-53 431,220) Ib 1960-61 612,000 Tb

1955-56 550,000. Ib 1961-62 675,000 Ib

1956-57 650,000 Ib

The total weight of rnullet handled by the Sonth Australian Fisherman's

Co-operative Limited (S.A,F.C.O.L.) from July Ist, 1961 to June 30th, 1962, was

584,984 lb. Approximately 75 per cent of this came from the Covrong Iagoon, the

other 25 per cent from the shallow coastal waters of South. Australia. The fish

* Formerly Department of Zoology, University of Adelaide. Present address, Department

of Zoology, University of Queensland,

Trans, Roy, Soc. $.A, (1968), Vol. 92.

38 J. A. HARRIS

| SOUTH

! AUSTRALIA

lograths Flat

Fig. 1, Location map of the

Coorong lagoon.

from the Coorong lagoon and those from the coastal waters are sold under

different names at $.A.F.C.O.L. as “Coorong mullet” and “sea mullet” respectively.

This investigation only deals with the “Coorong mullet”.

The legal minimum length for yellow-eye mullet in South Australia is 7

inches (17-8 cm), total length. As so little is known about the commercial fishery

in this state, the effectiveness of the present minimum legal length is open to

uestion. The small amount of data pertaining to fish below 17-8 cm is a

eficiency in the present investigation; however, the commercial catch (above

17-8 cm) has been covered as thoroughly as possible in the time available.

THE “COORONG MULLET” FISITERY

The name “Coorong” applies to a long, narrow lagoon and associated shallow

lakes, paralleling most of the upper south-east of South Australia. This area is

divisible into two sections, the zone of permanent lagoonal water to the north and

the shallow, non-perennial lake to the south, In Fig. 1 the lagoon is enclosed by

the rectangle. The “Coorong mullet” fishery is only concerned with the lagoon.

YELLOW EYE MULLEW PROM Tif COORONG 39

The Coorong lagoon extends southwards from the mouth of the River Murray

a distance of 68 miles and reaches a maximum width of 2% miles. The maximum

depth reached in winter near Salt Creek is about 4 feet, but the average depth

is only 6 feet. The mouth of the Murray provides the only connection between

the lagoon and the Southern Ocean. The mouth is about 300 yards in width and

benaiits ob a dal channel through the unconsolidated dunes of Younghusband

Peninsula.

Mesh and gill nets are the principal means of catching the “Coorong mullet”

and regulations restrict the size of the mesh and the lengths of these nets, The

influence that regulations on nets have on the yellow-eye mullet fishery in Western

Australia has been diseussed by Thomson (1957a). Regulations during the period

of this investigation restricted the length of any net to 60 yards, the length of

rohit nets to 35 yards and the size of the mesh used in all nets to not less than 2

Aches.

The “set net” is by far the most common method used for catching nvullet.

Lengths of mesh or gill nets are anchored to stakes across the current. They are

checked at regular intervals and the “gilled” fish collected. “Ring netting” is often

used with great success on calm, still nights. The fisherman lays a ret, one end of

which is attached to a buoy. around 2 school of fish. A few fishermen use beach

seine nets from sandy beaches or sand bars. Unfortunately, these beaches are rare

due to the recky nature of the bottom. During the summer months sand banks are

exposed jn the centre of the lagoon and from these banks beach seining is very

successfully carried out. Most of the mullet from the lagoon is sent te Meningie to

he packed in ice prior to transport to $.A.F.C.O.L.

SAMPLING METHODS

Weekly samplex of “Coorong mullet” were collected from $.A.F.C.O.L.

Adelaide market for one year from February, 1962 to January, 1963. Over 100 fish

were selected at random each week: and their lengths were measured by punctur-

ing, holes in & celluloid strip, mounted on a centimetre rule, with a hootmaker's

awl (see Scott, 1954), From this random sample approximately 12 fish were

chosen, representing the complete ranve of sizes of fish available, for further

detailed examination in the laboratary.

This detailed examination consisted of: (a) collccting scales and otoliths and

stonng them in labelled envelopes, and (b) macroscopic examination of the

gonads, after which they were fixed in 5 per cent formalin for microscopic exami-

nation.

On one occasion on October 7th, material was obtained from the Coorong

lagoon by seine netting in shallow water with a net of 1 inch mesh. Only 31 fish

tunging from 5-5 cm to 15-8 em were caught.

During the year the Jengths of 6,054 fish were measured and 631 otoliths

were examined in order to determine their age and rate of growth, In addition,

the macroscopic and microscopic conditions of about 600 gonads were noted. All

measurements OF lengths were taken from the tip of the snout to the eandal fork

(I..C.F,) onless otherwise stated.

DETERMINATION OF AGE BY MEANS OF OTOLITHS

Otoliths were found to be a good index of age and were tasy to handle anil

examine. Both otoliths in any one fish were found to be identical. No preparation

prior t) the reading of them was required (see Dakin, 1939), Regular opaque

bands and translucent bands are cbyious even in the largest otoliths. The two eto-

liths collected from cach fish were read together using a Jow power binocular

microscope.

40) J. A. HARRIS

As the otolith bands are formed by regular coneretions (Hickling, 1931), the

alternate opaque and translucent zones must at some stage actually constitute the

margin of the otolith. All the otoliths collected from each weckly sample of fish

were placed into two groups, those in which an opaque band constituted the

margin and those in which a translucent band constituted the margin. Transitional

or doubtful ones were discarded and these accounted for about 20 per cent of the

otoliths. The occurrence of opaque margins was plotted as a percentage of all oto-

liths examined during each month of the year from February, 1962 to January,

1963. The results are shown in Fig. 2.

Fig. 2 shows that the lowest percentage of otoliths with opaque margins

aceurs during August and September while the highest percentage occurs in

January. Translucent bands occur during the winter months and opaque bands

form during the summer months. A similar condition was observed for garfish in

South Australia by Ling (1958), for plaice (Molander, 1947) and for the tiger

flathead (Dakin, 1939; Fairbridge, 1951). Thomson (1957b), has shown that the

annuli in the scales of the yellow-eye mullet from both Western and Eastern Aus-

tralia are formed when growth recommences after the winter cessation.

The first translucent band is Jaid down during the first winter, approximately

6 months after spawning. The first opaque band is laid down during the first

summer after spawning. Hence, the number of opaque bands represents the actual

iC)

=60

re]

reg

o50

E40

rs)

fa)

630

na Fig. 2. Monthly percentaye

=20 incidence of otoliths with

rs) opague margins for the 12-

hi month period February, 1962

& to January, 1963.

FEB

MAR

APR

MAY

JUNE

JULY

AUG

SEPT

OcT

NOV

DEC

JAM

YELLOW EYE MULLET FROM THE COORONG Al

age of the fish in years. For this reason the opaque zones were counted and the

age groups denoted as 0+, 1+, 24+, etc., the numbers representing the age of the

fish in years and the + — sign signifying an additional unknown number of

months (but less than 12) above the preceding numbers. The results of the

otolith readings are given in Table 1, the number of fish in each age group are

arranged with their corresponding lengths.

Ling (1958) discusses in great detail the ways of confirming the otolith

method as a means of determining the age of fish. He discusses Graham’s original

five tests to validate the scale method (Graham, 1929). In this investigation Peter-

sen’s method of interpreting the modes of a graph of length distribution as the

model lengths of successive age groups is teed te support the data obtained from

otoliths.

Monthly histograms of the frequencies of lengths of fish measured each week

by the celluloid strip method (see Scott, 1954) are shown in Fig. 3. In the major-

ity of histograms a definite mode occurs about the lengths of 20-22 cm, a less

TABLE 1

Length frequencies of males and females with their ages, as Jetvermined from otoliths, for all fish

examined in the laboratory from February, 1962 to January, 1963.

Age groups Male Female

Length (erm) ot | 14 | 2+ 34 oy | 14 | ot, pe | at | oe

4 1 i 1

1 2

25 . 1 | 10 36

ce)

Total 2; 10 125 | 36 6 14 143 141 32 2

“| AUG

> -

i] Le)

- Z

Z & Zoo

2 2

o g

au uw

m0 JuLy = 0

ue iw

49 J. A. HARRIS

FIG 4

JAN ©

here)

ig SEPT jag b

a eee

LENGTH CCM)

Fig. 3. Monthly frequency distribution of

lengths of fish sampled each week for the

12-month period February, 1962 to January,

1963.

DEC

SEPT

JULY

JUNE

30 450

OVA DIAMETERS (ju)

Fig. 5. Monthly frequency distribution of

diameters of ova examined from fish

selected at random over the 12-month

period February, 1962 to January, 1963.

(Note: Hickling’s “reserve fund eggs” were

only measured in February ),

YELLOW EYE MULLET FROM THE COORONG A

definite one about 24-26 cm, and the trace of a third mode about 30-32 em. By

comparison with the different age groups obtained using otoliths, these modes cor-

respond to the age groups 2+, 3+ and 4+ respectively. Theretore, agreement

between the Petersen method and the determination of age by means of otoliths

is quite good. It can be seen that the majority of fish are in the age group 24-

with a smaller number in the age group 3-+ and very few in the age group 4+.

FREQUENCY

Fig. 4. Frequency dis-

tribution ot lengths of

fish caught October

7th with a net of 1 in,

mesh, a

°o i] ’

LENGTH (CM)

Fig. 4 shows the length frequency histogram of 31 fish caught on October 7th

with a net of 1 inch mesh, Only one mode at 15 cm is obvious, but probably there

is one at 8 em. These modes correspond to the age groups 0+ and 1+ respee-

tively. In general, it may be stated that the determination of age by means of

otoliths is supported by the results of the Petersen method.

RATE OF GROWTII

(a) Using otoliths and measurements of length

Table 1 shows the frequencies of lengths of males and fernales with their

ages, us determined from otoliths, for all fish examined in the laboratory from

February, 1962 to January, 1963, An analysis of these results was carried out to

test the two null hypotheses implicit in these results; (1) that there is no differ-

ence in the lengths of the fish in the age groups 0-+ and 1-++, 1+ and 2+, 2+ and

3+, 3+ and 4+, and (2) that there is no difference in the lengths of a male and

female in the age groups 0+, 1+-, 2-+ and 3+. These null hypotheses were tested

by the “t” test (Snedecor, 1956). The first null hypothesis was disproved for all

cases, but the second null hypothesis was not disproved for all cases. The differ-

ences between the lengths of taale and female fish in groups 0-+- and 1+ were not

significant, but those for groups 2+ and 3+- were highly significant.

The mean lengths attained by the female “Coorong mullet” during their first

four years are 14-2, 21-7, 26-1 and 30-9 cm respectively, Rapid growth occurs

during their first lwo years, but then falls off during their third and fourth years.

The mean lengths attained by the males during their first three years are 13-7,

21-1 and 24-5 cm respectively. Growth is again rapid during their first two years,

but during their third ycar the mean length attained is 1:6 em less than that of

the females. After the first two years, the rate of growth of males (using only

length as the measure of growth) is much slower than that of females.

44 J. 4. HARRIS

The two largest fish examined during this study were both females, approxi-

mately 35 em Jong. Only one male fish exarnined was. found to be longer than 26

em and it was 30 em in length. These data indicate that the females grow tu a

larger size than the males.

(ly) Using the Petersen method

As mentioned earlier, the histograms of frequencies of lengths given in Fig. 3

show two definite modes occurring about the lengths of 20-22 em and 24-26 cin

with perhaps the trace of a third one about 30-32 cm. These modes are considered

to correspond to the age groups 24-, 3+ and 4+ respectively, The evidence far

mouul progression when following the progress of one particular mode in the

ovurse of time is very good,

If March is taken as the tine that spawning activity is at its peak, the histe-

gram for March inay be said to represent the Frequencies of Jengths at the begin-

ning (or end) of one or more complete yeurs in the life-cycle of the fish. Definite

modes occur about the lengths 20-5, 24°5-26 and 30:3 um respectively in the

March histogram. It is easy to trace the mode about 20-5 em throughout the

year. In April and May, just prior to winter, a $mall amount of growth is apparent.

In May the mode is obvious about 21°35 em. Over the winter months June to Sup-

tember this mode about 21°35 em is maiutained, indicating no growth, or very little

(as also indicated by the study of otoliths). From October to January the mode

shifts rapidly from about 22 em in October, to 23-5 cm in November, 24-5 cm iv

December and 24 -5-26 em in January, The histogram for February of the previous

year shows a very good mode aboul 26 cm and supports the evidence for mudal

progression over one year being approximately 20+5 to 26 om. This corresponds. ta

a growth of 5-5 em which is very close to the annual growth of a 2-year-old “Cou-

rong mullet” as sugeested by the study of their otoliths, Over the same time period

possibly the mode about 24:5-26 em gives way to modes about 28:5 em in Qcto-

ber and November, 29-5 em in December and 30°5 um in January. This corres-

ponds toa growth of 4-5-6 em which is roughly equivalent to the rate of growth of

ad-year-old fish as suggested by the study of otoliths.

THE SPAWNING CYCLE

‘The methods by whicli the spawning cycle may be studied inelude a survey

ol the seasonal variations in the conditions of the gonads. This involves a macra-

scopiy examination of the appearance of the gonads and a microscopic examinn-

lion of the ova.

In most investigations (Clark, 1925; Fairbridge, 1951; Scott, 1954) the female

eycle has heen studied in more detail than the male cycle. In the present investi-

gation the male cycle was examined in less detail because it was found hard to

distinguish the stages of the testes and because the condition of the ovaries

changed much more noticeably than that of the testes, A month by month com-

parson of the macroscopic canditions of the avaries was noted and this was later

verrelated with a microscopic examination of the ova, including measurements of

their size.

(a) Macroscopic examination of gonads

Hjort’s classification of the gonads of herring (Hjort, 1910) has heen

adopted with cvertain modifications to suit this particular specivs. Seven stages in

the ovaries of the “Coorong mullet” were distinguished while the testes were dis-

tinguished only us immuture or riature. The classification of each of these seven

staves of the ovaries is given in Table 2,

YELLOW EYE MULLET FROM THE, COORONG 45

TABLE 2

Lengths and stage of maturity of 348 females sampled during January, 1963-

Stages of ovaries

Length (em) | Per cent

Immature Adolescent Maturing Spent maturing

| and spent,

19 2 | | 0

2) 2 2 | 0

2 2 2 | 0

22 3 2 i ()

23 12 ls 2 1

z4 14 55 25 J 27

25 Li 43 a0 47

26 3 21 26 3 49

27 2 17 89

28 4 4 i 70

2a. 9 100

30) 5 10

al 3 100

Total 52 148 | idl 7 43

Stage I. Immature virgins. Ovaries small, thread-like and translucent.

Stage ll. Adolescent, Ovaries larger, walls taut, extending half-way into the

body cavity,

Stage IIT. (a) Maturing virgins or (b) recovering or resting (spent) fish,

Colour ranges from pink at early pigmentation to dark red at a later stage, In (a)

walls of ovaries are taut but in (h) walls are ilaccid, Ovaries extend further into

body cavity.

Stage IV. Maturation continuing. Ovaries larger, red-yellow in colour.

Walls taut and not quite transparent so that eggs can only be seen with difficulty

afler cutting the wall.

Stage V. Mature, Ovaries greatly enlarged from stage IV, occupying the

whole of the body cavity, Yellow-orange in colonr, walls taut and transparent with

eggs clearly visible,

Stage VI, Running-ripe. Yellow-orange in colour with large eggs being

extruded when slight pressure is applied to the abdomen,

Stage VII. Spent. Oyaries dark red aud Haccid. This stage eventually passes

into stage IIL from which it is hard to distinguish.

The seasonal changes in the gonads may be summarized as follows, The

testes of the immature fish are white, thin bodies, triangular in section. During

maturation they become thicker and rounder, protruding further into the body

cavity and milt being exuded if the walls are ruptured. During the months of

May, June and July no mature testes were noticed. For the rest of the year there

were always a few mature testes present and during the spawning season January

to March, over 50 per cent of all testes examined were mature, (See Table 3 for

the percentage of maturing males during January),

When the inyestigation started in February, 80 per cent of the ovaries

exumined were mature (stages IV, V and VI) with 25 per cent of these being

in the “ripe” condition (stage VI). During March a similar set of conditions

46 }, A. HIARAIS

continued with a few spent ovaries (stage VIL) becoming apparent. However.

during April and May spent ovaries became more evident and the last mature

ovary (stage VI) was found on April Eith, except for one ovary at stage V found

on May 2nd which appeared to be abnormal, only one branch having developed,

During the months of June and july, except for one fish on July 25th which

was at stage IV, all ovaries were classified as belonging to stages II and III.

Approximately 85 per cent of the ovaries examined during August, September and

October were at stages IL and IlI. Only 15 per cent were more advanced than

stage IJI and it is possible that they were carly spawners. During November and

December, however, ovaries at stage FV became more evident with an occasional

stage V until in Jannary ovaries at stage V were common with several at stage V1.

(Sce Table 2 for the percentage of maturing females during January. )

(h) Microscopic examination of ovaries

Several ovaries chosen at random, were set aside each week, preserved in 5

per cent formalin and stared in labelled vials. These ovaries were examine:

under the microscope al a later dite.

To measure the diameter of the ova a small piece of ovary was teased out

onto a microscope slide, mounted in 5 per cent formalin and covered with a cover

slip. The diameters were measured using a micrometer eye-piece and a micro-

scope,

; All eggs were measured along whichever axis of the egg lay parallel to the

cross-hair of the micrometer eye-piece. Clark (1925) has shown that this methoul

gives a reliable estimate of the frequencies of diameters of the ova. Throughout

the investigation group A eggs were not measured. except in February, as they

were far too numerous in all of the oyarian stages.

Consistent with the classification of Hickling (1930) and other workers, the

ava of the “Coorong mullet” were classified into four groups.

Group A — Immature ova. Very small, angular and colourless with nucleus

elearly visible. They are Hickling’s “reserve fund eggs” and measure up to 126) »,

Group B — Maturing ova. The smallest eggs in this group are becoming

round and opaque due to accumulation of yolk granules, while the largest ova are

quite round and opaque. Size varies from 120 p to 255 py,

Group C — Mature ova. Large round opaque ova, pale yellow. Size varies

from 255 pe to 380 p.

Group D — Ripe ova. Very large round opaque ova, yellow. Size varies from

380 » to 600 ». This stage merges with group C, but is distinct from group C by

size and coloration,

The microscopic characters of each ovarian stage were established as follows:

Stage I. The ova are all typically group A, with none larger than 60 p.

Stuge II. As for stage I, but an occasional ova of group B may be present

No ova larger than 135 p,

Stuze DT. Ova of groups A and B are very mimerous with an occasional

group C. Usually ne eggs larger than 255 yu.

Stage TV, Ova of groups A, B and © are mimerous but group B is usually

nore numerous than group C.

Stave V. Ova of groups A, B and C are numerous with a few of group 1).

Group C is usually more numerous than groups B and D,

Stage VI. As for stage V, but the ovaries are much larger with ova of croup

D most numerous and lying free in the lumen. Groups A, B and C ere confined to

a layer inside the oyary wall.

YELLOW EYE MULLET FROM THE COOKONG AT

Stage VII. Ova of groups A and B are both numerous. An occasional group

D ovum is present undergoing breakdown.

The frequencies of diameters of the ova examined were plotted as monthly

histograms and are shown in Fig. 5. Throughout the year there is a residue of

Hickling’s “reserve fund eggs” which were too numerous to measure every month

and are shown only in the February histogram. From Fig. 5 it can be seen that

numerous ova of group B also are present throughout the year. From May to

October (except in August when the ovary examined may have been that of an

early spawner) ova-of groups A and B are by far the most numerous. During

November, however, an increase in the diameter of the ova present is obvious. As

maturation continues through December, the diameters of the oya continue to

increase until “ripeness” (stage D) is obtained. Ripe eggs are present in greatest

numbers during March, with large numbers also present in February and April.

Spawning is thus at its greatest intensity during late summer. A sharp drop in the

overall diameter of the ova marks the spent and recovering stages, during April

and May. It is followed by a period of quiescence from June to October.

SIZE AND AGE AT FIRST MATURITY

The gonads of 122 male and 348 female fish were examined; and their lengths

taken, during visits to §,A.F.C.O.L. in January, 1963, The females were classified

arbitrarily as immature (stages I and IL), adolescent (stage III), maturing (stages

{V, V and VI) and spent (stage VIL). The males were classified under two divi-

sions, immature and maturing. Tables 2 and 3 show the lengths and stages of

maturity of these males and females.

Table 2 shows that 15 fish with lengths of less than 22 em were examined

and none showed signs of maturing, Sixteen per cent of the fish 23 cm in length

were maturing and spent, The percentage rose to 100 per cent for all fish 29 cm

and above. Females, therefore, do not reach maturity before 23 cm.

Table 3 illustrates that only 3 males 21 cm long were examined and that one

of these was maturing. Because so few fish of this length were examined it cannot

be proved that males mature at 21 cm. However, 19 fish of length 22 cm were

TABLE 3

Lengths and stage of muturity of 122 males sampled during Jamnary, 1963.

Stages of testes

Length (em) Immature Maturing Per cent raaturiag

20) | 4 0

21 ' 2 i 33

22 1 A 42

24 22 28 57

24 Il a4 6a

25 I 7 87

26 2 100

Total 51 71 61

examined and uf these fish 42 per cent were maturing, Males, consequently, are

definitely mature by 22 em.

Correlating the lengths at which the females and males first mature with the

data obtained from the otoliths, it is concluded that the “Coorong mullet” attain

maturity during their third year.

4h J, Av HARRIS

DISCUSSION

The “Coorong mullet” grows to a relatively small size of 35 cm. At first the

growth, using only length as a measure of growth, is rapid. The mean lengths

attained during their first and second years are 14 and 21 cm respectively, After

this period, however, there is a marked slowing down in growth reaching a mean

length of 26 and 31 cm during their third and fourth ycars respectiyely. This slow-

ing down is more prouounced in ‘the males which do not grow as large as the

females.

TABLE 4 2

Lengths (em) attained cach winter average by the Western and Mastorn races of theYeollow eye

Mullet (‘CThomson, 1957b) and the mean lengths attained by the “Cusrong mullet’? over aa

equivalent time poring,

l 1

Western i !

and Easter I i IL Lit TV ¥ VL Vil

vares | |

i ——— = = nl, ee ne

“Coorong | 4 '

roullet™ | Of Af BOB

Western fish | a 18.19 | 24.25 | 29-32 | 32-25 | 38 49

Eastern fish 5 | wus | i921 | 2427 9 a0

“Oourung rules” | T(t) 14 21 26006¢,~—C 3]

Table 4 compares the growth of the western and eastern races of the yellow-

eye mullet with that of the “Coorong mullet”, The data for the western and eastern

races is taken from Thomson (1957b), The year-classes designated in Thom-

son's paper refer to the length attained each winter average (the western stock

being winter spawners). Over an equivalent time interval the “Coorong mullet”

rows to a mean length of 7(?), 14-2, 26 and 31 cm by their first, second, third,

ourth and fifth winters respectively. Tt can be seen that their growth rate agrees

closely with that of the Victorian and Tasmanian yellow-eye mullet which com-

prise the majority of fish referred to as “eastern” by Thomson (1957b), The “Coo-

rong mullet” grow at a slower rate than the Western Australian yellow-eye mullet

and do not become as large.

The spawning period Jasts for 3-4 months from January to early April. This

proves that the “Coorong mullet” have the characteristics of the eastern race

rather than the western rave of yellaw-eye mullet (see Thomson 1957b, p, 12),

Spawning probably takes place in the Coorong lugoon which would provide a

very productive area in which the newly hatched fish can grow rapidly with little

chance of a shortage of food occurring. The Jargest ova of the “ripe” fish were

upproximately 600 , in diameter which is relatively small compared with whiting

980 » (Scott, 1954) and garfish 3,500 » (Ling, 1958).

The smallest fish recorded with mature gonads were 23.cm for females and 21

em for males, They mature during their third year or by their third winter. The

“Coorong mullet”, therefore mature a little earlier and at a slightly smaller size

than the Western Australian yellow-eye mullet studied by Thomson (1957a). In

addition, due to the spawning periods of the two raccs of fish occurring at differ-

ent times of the year, the summer spawning “Coorong mullet” reach maturity

faster than the winter spawning western racc,

Comparing the commercial catch of “Coorong mullet” with the catch of

yellow-eye mullet in Victoria, some interesting facts emerge. In Victoria the fish

YELLOW EYE MULLET FROM TITE COORONG 49

reach the minimum legal length, 9% inches total length (21:3 em L:C.F.), from 2h

to 32 months prior to spawning (Thomson, 1957a). In Sauth Australia the

tinimum Jegal length is 7 inches total [length (15 em L.C.F.) and the fish attain

this size approximately 12 months before they have spawned for the first time.

South Australia’s minimum legal Jength is obviously too small and at first glance

the “Coorong mullet” fishery appears to be in jeopardy. However, Fig. 3 shows

that no fish below LT om were sampled at S.A-F.C.O.L. The majority of fish, in

fact. were well above 20 cm. This biased sample was due to the size of the mesh

of the nets used by the commercial fishermen. In South Australia the minimum

legal mesh size during the study was 2 ches, compared with 2M inches in

Victoria, Thomson (1957a) showed that, as far as vellow-eye mullet are con-

cerned, any mesh from 2 inches to 34 inches allows the majority of the catch to be

above the length of 20 crn.

Thomson (19571) discusses the management of the yellow-eye mullet fishery

heth in Eastern and Western Australia and concludes that the existing manage-

ment policies are elective in maintaining stocks. Whetber this ix alsa tre fur the

“Coorong mullet” fishery cannot be determined yet due to lack of information,

both biological ancf stulistical. A study of the length/weight relationships 4s

required together with estimations of fecundity and observations of the survival of

fry and small immature fish. Statistics of the catch and fishing effort are also

required hefore a more complete management programme can be suggested,

ACKNOWLEDGMENTS

The project was suggested and supervised by Mr, I. M, Thomas, Zoology

Department of the University of Adelaide, prior to going overseas on sabbatical

leave. I am greatly indeb(ed to Dr. S.J. Edmonds for supervising me during Mr,

Thomas’ absence, for reading the original draft and offering much helpful advice.

Thanks are due to Mr. T, D, Scott, formerly Curator of Fisheries of the South

Australian Museurn for his guidance and to Prof. J. M. Thomson, Zoology Depart-

ment of the University af Queensland, for critically reading this manuscript.

Vinancial support for this investigation came from the Research Grants Committee

uf the University of Adelaide. Finally, thanks are due to S.A.F.C.O.L. for enahling

the sampling programme to be carried out on their premises.

REFERENCES

Crank, Frances N. (1923).—The life history of Leuresthey tennis, an atherine fish with tide

controlled spawning habits, Fish. Bull., Sacramento No. 10: 1-51,

Darin, W, J. (1939), The age determination of the tiger flathead. Neoplatycephalis (Calefrxia)

mactradaw (Ogilbv), by means of otoliths. Rec, Aus, Mus. 20; 282-93,

Famnmripce, W. 5, (£951). The New Sovth Wales tiger fathead Neoplatycephalusa macrodan

(Ogilby). T. Biology and age determination. Aust. J. Mar, Freshw. Res, 2; 117-78,

Granam, M. (1929). Studies of age determination in fish. 1, A survey of the literature, Fivh,

Incest,, Lond, (2)11(3); 1-50,

Thexiing, C, F. (1980). The natural history of the hake. LT, Seasonal changes in the condition

of the hake. ish. Invest, Lond. (2)12(1); 1-78,

Hiexiane, C, F, (1931) The stracture of the otolith of the hake, Quart. J. Mier, Sel. 74: 547-61

your, J. (1910), Report an herring investigations until January, L910. Publ, Cire. Cons. Explor.

Mer, No. 53; 1-173.

Linc, J. K. (1958),. The Sea Garfish, Reporhamphus melunachir (Cuvier & Valenciennes )

(Hemiramphidue). in South Australia; Breecling, age determination and growth rate.

Aust. J. Mar. Freslaw, Res, 9; 60-105,

Morannen. A. R, (1947), Observations om the growth of the plaice and on the formation nf

annual rings in the otolith. Seenska Hydrogr—biol, Komm, Sk, 2: 1-9.

Scatr, T, D. (1954). M.Sc. Thesis. University of Adelaide.

Ssvepecon, G. W. (1956), “Statistical methods applied to experiments in agricultyre and

hiology.” Fifth edition, (The Towa State College Press: Ames, Towa,)

50 J. A. HARRIS

THomson, J. M. (1957a). Biological studies of economic significance of the Yellow-Eye Mullet,

Aldrichetta forsteri (Cuvier & Valenciennes) (Mugilidae). Aust. J. Mar. Freshw. Res. 8;

1-138,

Tnromson, J. M. (1957b). Interpretation of the scales of the Yellow-Eye Mullet Aldrichetta

forstert (Cuvier & Valenciennes) (Mugilidae). Aust. J. Mar. Freshw. Res. 8; 14-28.

THE DISTRIBUTION AND LIFE HISTORY OF THE SKINK

HEMIERGIS PERONII (FITZINGER)

BY MICHAEL SMYTH*

Summary

Hemiergis peronii occurs from south-western Western Australia to southeastern South Australia,

but rarely inside the 12-inch isohyet and not in the Flinders or Mt. Lofty Ranges. Its northern limit

is probably determined by aridity; its southern limit might be determined by the length of the

winter.

H. peronii bears two to five living young in February. Females are inseminated in the autumn, when

they are two years old. They ovulate in spring and do not bear their first young until they are three

years old. Males first come into breeding condition when two years old.

H. peronii eats mostly arthropods and snails.

THE DISTRIBUTION AND LIFE HISTORY OF THE SKINK

HEMIERGIS PERONI (FITZINGER)

hy MicuaAri, SmytTn®

[Read 11 July, 1968]

SUMMARY

Hemiergis peronii ocours from sonth-western Western Australia to south-

eastern South Australia, but rarely inside the 13-inch isohyet and. not in the Flin-

ders or Mt. Lofty Ranges, Its northern limit is probably determined by aridity;

its sonthern limit might be determined by the length of the winter.

A. peronii bears two to five living young in February. Females are insemi-

nated in the autumn, when they are two years ald. They ovulate in spring and

do not bear their first-young until they are three years old, Males first come into

breeding conclition when two years old.

IL. peronii wats mostly arthropods and snails.

INTRODUCTION

Hemiergis peronii is a small, weak-limbed skink, very abundant in coastal

dunes near Adelaide. Its reproductive cycle is unusual in that the females are

inseminated in fhe autumn But do not ovulate until spring Gsesivth and Smith,

1968). I now report some other details of its life history. They will provide a

background against which the adaptive significance of the unusual reproductive

cycle will perhaps become clearer, and they will slightly diminish our great ignor-

ance about our native reptiles,

Some authors use Hemiergis as a sub-genus in the genus Lygosoma, H. per-

onii is called L. (H.) quadridigitatum Werner by Glauert (1961), probably for

reasons which are explained and dismissed by Loveridge (1934). Worrell (1963)

calls it Lygosoma ( Leiolopisma) peronii.

METIIODS

The distribution of H. peronit was mapped from the records of the South

Australian, Western Australian, and Australian Museums and the Department of

Zoalogy, University of Melbourne, from the published records of Wermer (1910),

Waite (1929), Loveridge (1934) and Mitchell and Behrndt (1949), and from my

own collections and those made for me by a group of students of Naracoorte High

School directed by Mr, D, Von Behrens.

The natural history was described from samples of from four to 16 lizards

taken at two to four-weekly intervals for a year at Port Gawler and Middle Beach,

about 30 miles north of Adelaide. Most of them were taken from under dead

clumps of the lily Dianella revoluta R, Br. on shell-grit dunes behind the beach.

The lizards were brought hack to the laboratory and kept at 10°C until they were

* Department of Zoology, University of Adelaide:

Trans, Roy. Soc, S.A. (1968), Vol, 92,

52 MICHAEL SMYTH

dissected. They were then measured with dividers, their reproductive organs

removed and examined, their guts removed, the food taken out and identified, and

the guts examined for parasites, he bodies were then preserved in buffered neu-

tral formalin, The gut parasites are described hy Angel and Mawson (1968),

DISTRIBUTION

H. peronit is confined to southern Australia, from south-western Western

Australia to south-eastern South Australia ( Fig. 1), It probably occurs in western

Victoria as well, for according to Rawlinson (1966) there is a specimen in the

National Muscum, Mclhourne, listed in the catalogue as from “Mallee district,

Victoria’. But it does not occur in southern Victoria, Tasmania, or the Bass Strait

islands ( Rawlinson, 1967). Lucas and Frost's (1894) claim that it has becn taken

in the Dandenong Ranges near Melbourne is probably mistaken, and Weekes’s

4 SOUTH AUSTRALIA |

WESTERN AUSTRALIA

VICTORIA

TASMAN No

Fig. 1

(1930) claim that she took Lygosoma (H,) quadridigitatum at Tanara and

Jenolan, New South Wales, is probably based on a wrong identification.

In Western Australia, I. péronii is found into the 50-60 inch rainfall belt near

Northcliffe, but as far as we know at present its range does not extend right to the

west coast. Loveridge (1934) does record a specimen from Perth, but this might

have been the address of the collector rather than of the lizard, It has been taken

as far inland as Fraser Range, in the 11-12 inch rainfall belt.

In South Australia, H. peronii is very abundant around much of the coastline

(Fig. 2), It also occurs on many of the offshore islands, including St, Francis,

Franklin, Flinders, Pearson, Greenly, Price, Black Rock, the South Neptune, and

Wedge Islands. It occurs inland on sandy or skeletal calcareous soils on Eyre and

Yorke Peninsulas, Kangaroo Island, and in the south-east of the State. Its distribu-

tion extends slightly inside the 12 inch isohyet only on the West Coast and near

the head of St. Vincent Gulf (Fig. 2). It seems not to occur in either the Flinders

or Mt. Lofty Ranges.

Further collecting will no doubt extend the known range of H. peronii. For

instance, it might be found all the way around the Bight, and further into the

Murray Mallee of South Australia.

HEMIERGIS PERONII 53

santana

REPRODUCTION

Male H. peronii come into breeding condition and inseminate the females in

the later summer and autumn (Smyth and Smith, 1968). The sperm is stored in

the female genital tract over winter; the females ovulate and their eggs are fertil-

ized in the spring, between late October and the end of November. Two, three or

four young are born at the end of February, larger mothers bearing on the aver-

age large litters.

3 4

> A

v 3 3 Fig. 3. The frequency distributions of

2 snout-to-vent lengths of female H, peronit

z bearing two, three and four young. The

wi | number of young was counted from either

5 large ovarian eggs (white) or eggs and

O embryos in the oviduct (black),

ur 3 2

]

5 6 7

LENGTH (cM)

a4 MICHAEL SMYTII

TABLE 1

Breeding condition of femulu Hemiergis peroni,

Number of females with*

= = , Total

Date no sign of number

developing ovarian oviducal = | reproductive

follicles; verges eggs activity

16 November 1966 1 1 2

29 November é 1 4

14 December 6 2 8

§ January 1967 8 3

25 January A ui ‘BI

16 February 4 3 7

1 March 5 5

14 March 2 2

28 March a 9

12 April 2 2

25 April 8 8

10 May ij 2 8

24 May + | sy

13 June 6 1 7

18 June 4 4

18 July 1 l 2

18 August <4 1 3

19 September 2 1 3

3. (ctober 4 | 6

17 October j 5

31 October 2 3 4

21 Nouyermber 2 2 4

*Femaics with no sign of reproductive activity included juveniles, ono-year-olds, and, in

March and April, older ammals. Their ovarian follicles were small and transparent. Females were

said to have developing follicles when the follicles berarne an apace white, though thoy had

enlarged very little if at all, Later, after the follicles had obviously begun Lo enlarge, the females

wore sail to have ovarian eggs.

In H. peronii the right ovary is well anterior to the left in the body cavity,

and the right oviduct is much longer than the left. About equal numbers of eggs

are shed from each ovary but a high proportion migrates from the left ovary to

the right oviduct. This is deduced from a comparison of the distribution of the

corpora lutea between the wo ovaries with the distribution of eggs and embryos

between the two oviducts. Thus of 90 eggs shed from the ovaries of 29 females,

48 came from right ovaries, but 61 embryos developed in right oviducts and

only 29 in the left. In only one of the 29 females was there more than onc embryo

in the left oviduct; in this case there were two in each. The most posterior egg

was always in the left oviduct,

GROWTII AND AGE STRUCTURE

Figs. 4 and 5 show the distribution of snout-vent lengths of all individuals

taken during the vear. It can be shown from these figures that males first come

into breeding condition, and females are first inseminated, when they are two

years old. Females therefore bear their first young when they are about three

years old, Consider the snout-vent lengths of females (Fig. 4); in April and May

there are clearly two year-classes in which there is no follicular development.

These have been delineated in Fig. 4; the lower group is of young of the year, the

HEMIERGIS PERONII 5D

SV LENGTH Cem)

N DJ F MA MJ JS A S ON

Fig. 4. The snout-to-vent lengths of all female H, peronii caught from

November, 1965 to December, 1966. @ pregnant; ® not pregnant,

follicles undeveloped; © follicles visibly developing; + juvenile, sexes

not distinguished.

upper group is of onc-year-olds. Now consider the distributions for Noyember,

1966 to February, 1967, when the females were pregnant. It is apparent that those

females not pregnant must have belonged to these two youngest year classes. A

somewhat similar argument can be applied to the males (Fig. 5), From February

to May, when the testes of most malcs in the samples were enlarged, there were

always some males with quite undeveloped testes; these were young of the year

and, as well, larger animals which must have been one year old.

Neither the males nor the females, once they have reached sexual maturity,

can he confidently separated into further year-classes, but it is likely that there are

at least two year-classes of pregnant females in the summer, which by February

will be three and four ycars old respectively. The larger females, probably

approaching four years old, bear the largest litters (Vig. 3), Females grow to be

longer than males.

SV LENGTH Com

N O JS FM A MJ J A S O N

Fig. 5. The snout-to-vent lengths of all male H. peronii caught from

November, 1965 to December, 1966, @ testis and/or epididymis con-

tains sperm; © no sperm; + juvenile, sexes not distinguished.

36 MICHAEL SMYTH

FOOD

Tt was fairly easy to classify most arthropod foods to Order because at least

the heads and usually other parts as well passed through H. peronii intact. But

some other foods were probably missed. Land snails, for instance, were somctimes

found without shells or with only a féw fragments of shell adhering, which made

them hard to notice. Also, the shells of land snails were probably sometimes con-

fused with the small marine shells, mostly of gastropods and foraminifera, which

made up much of the ground on which the lizards were living, and which often

appeared in the lizards’ guts. So the proportion of land snails in the diet was

probably under-estimated.

The abundance of various items of food taken by male, female and juvenile

H, y:eronii is shown in Table 2, Obviously H. peroniié is almost exclusively insecti-

yorous. The only plant food found more than once was the seeds of the lily D.

revolita under which most of the lizards were caught. These seeds are small,

black and shiny, and might have been mistaken for insects. They were largely

undigested.

TABLE 2

The dict, of Hemieripis peronit.

Numbers of each item 9% wf botel

Ttem . ; es i

Males Females | Juveniles Males Females | Juveniles

Beatles 100 133 6 28-3 299 7-1

Ants anc other

Hymenoptera. 24 iG 2 6-8 12.6 24

Cockroaches 30 23 3 8-5 5:2 3-6

Jockroach oothecus 15 25 4-2 516

Moths | 25 22 7-1 49

Bugs (Hemiptera) 17 10 4-8 22

Unidentified insect

larvae i 1} Bel 29

Collembola 2 1 0-6 02

Grasshoppers 2 0-6

Flies L ()-3

Earwigs 1 0-3

Ant-lions : 1 Og

Mites 41 11 60 11-6 27:2 82-1

Spiders 13 1h 3-7 22

Psendoscovpinos 4 2. 0-8 24

Sluturs (Isapoda) 4 7 T-t 16

Centipedes 1 Q.2

Snails 20 6 1 8-2 13 1-2

Lizards 2 1 O+¢ Od

Secda 21 ' 14 5-4 a+4

Unidentified 1 | 1 3-4 1-3

Total numbor of |

individual items a3 445 | 84

The range of size of the foods taken was large, from small mites, mostly Ori-

batids, up to adult skinks Ablepharus greyii (Gray), which are about 4 cm long.

Juveniles took only small items: there is very little ditterence between the foods of

males and females. Most of the animals caten were feeders in litter or on the

surface.

VEMIERGIS PERONI oT

INJURIES TO LIMBS

Several A, peronii had lost digits or parts of limbs. This would he no great

hanclicap to them, since their movement is mainly by lateral undulations. Some of

these members would have been lost in accidents: { found one H. peronid with an

aut's head still firmly clamped onto its foreleg, though the limb distal to the ant’s

mandibles had withered and would soon have dropped off. In other cases, the

members might have been lost by discase or fights.

Males and females apparently lost large portions of front and hind limbs with

about equal frequencies, but in both sexes toes were missing much more often

than fingers (Table 3), This suggests that digits and large portions of limbs are

lost from different causes.

TABLE 3

Limh injuries in Hemieryéx povewit.

——— rrr

| Male | Fernale

Na, feet from which one ue more digits Were lost | front 5 i

| hinrl pm | | 20

No. linbs wholly ov partly last | front Ni 4

hind | 4 0

SO, animals with wll members intart 6 73

No. animals exumined te Ys

One I. peronii was found to have five fingers on cach front foot, though the

usual number is four, This anomaly is worth mentioning only because several spe-

cies of Hemiergis are most easily distinguished on the basis of the number of their

fingers and toes. Werner (1910) claimed that this was the only way he could

separate them at all. Glauert (1961) mentions some other freak numbers of digits.

and gives the impression that the mmmber of digits is not a good specific character.

DISCUSSION

Tt is likely that the northern limit to the distribution of H. peronii is deter-

mined by aridity, for its powers to resist desiccation are relatively poor. Warbur

(1966) has shown that its rate of evaporative water loss increases rapidly wit

increasing temperature, and that it quickly dies in a dry atmosphere at tempera-

tures as low as 35°C,

The southern limits to ils distribution might be delennined by winter temper-

atures. Where winters are cold, lizards usually become torpid, and will hecome

active again only in warm, sunny weather, Even then, they probably do not feed,

for torpid cold reptiles with food in their guts are in mortal danger because they

cannot digest the food, which then putrefies (Regal. 1966), I found food in the

gut of H, peronii throughout the winter, so, though it certainly becomes less

active, it probably does not become torpid for long. Also, abdominal fat bodies,

which in some lizards are known to be important in over-winter metabolism (Des-

sauer, 1955), are absent in H. peronii, though they are prominent in some other

local skinks.

So H. peronii might need to feed over winter in order to survive. If this is 80,

its pteferred body temperature should be low. There is no information available

on this, but Licht, Dawson, Shoemaker and Main (1966) haye shown that the

preferred body temperature of I. quadrilineatum is relatively low, and

38 MICHAEL SMYTH

If. quadrilineatum, like II. peranii, has a very high rate of evaporative water loss

(Licht, Dawson and Shoemaker, 1966).

Why HH. peronii does not occur in the Flinders or Mt. Lofty Ranges is hard to

say. Over much of its range it occurs on sand; in the south-east, for instance, its

range seems to coincide with the relict coastal dunes there (Sprigg, 1952). On

parts of Eyre Peninsula it occurs on soils which, though not yery sandy, are

underlain by kunkar derived from old sand dunes. Perhaps H. peronii avoids soils

prone to watcr-logging. But obviously a full analysis of its distribution reqnires

much more detailed study.

ACKNOWLEDGMENTS

1 wish to thank John Mitchell, Glen Storr, Harold Cogger, Peter Rawlinson,

Dierk Von Behrens and students from Naracoorte High School, Michael] Braysher,

Meredith Smith, Vaal Nielsen, Beverley Jones, Stuart Harris and Lewis Chinner

for help and advice.

REFERENCES

ANGEL, J., M.-and Mawson, P. M. (1968). Helminths of some lizards from South Australia.

Proe. R. Soc. §, Aust. 92; 59-72.

Drssaurr, H, C, (1935), Seasonal changes in the gross organ composition of the lizard, Anolis

carolinensis. J, Exp. Zool. 128: 1-12,

Guaven?, L. (1961). “A Handbook of the Lizards of Western Australia”, (Western Australian

Naturalists’ Club: Perth.)

Lacirr, P., Dawson, W. BR. and Suommaxen, V. H, (1966), [leat resistance of soit: Australian

lizards, Copéiga 1966: 162-169.

Licut. P,, Dawson, W. R.. SHoemAxeR, V. H. and Mat, A. R. (1966). Observations ou the

thermal relations of Wexteru Australian lizards, Copeia 1966: 97-110.

Lovenipce, A. (1984). Australian reptiles in the Museum of Comparative Zoblogy, Cambridge,

Massachusetts. Bull, Mus. Comp. Zool, Harvard 77; 243-383.

Lucas, B Li. S. and Frost, C, (1894). The lizards indigenous to Victoria. Proc. R. Soc, Vict.

6; 24-92.

MrrceEct, I. J. and Brarnnr, A. ©. (1949), Fauna and flora of the Greenly Islinds, Part 1,

Introductory narrative and vertebrate fauna, Rec, South Aust, Mus, 9: 167-179.

Rawniixson, P. A. (1966). Reptiles of the Victorian Mallec. Proc, R, Soc. Vict. 79: 605-619.

Rawrrsson, P. A. (1967). The vertebrate fauna of the Bass Strait Islands; 2. ‘he Reptilia of

Flinders and King Islands. Proc, BR. Soe. Vict. 80; 211-223.

REGAL, e J. (1966). Thermophilie response following feeding in certain reptiles. Copcia 196A:

88-390.

Syeytat, M. and Saar, M, J. (1968). Obligatory sperm storage in the skink Memiergis peronti.

Science 161: 575.

Spence, R, C. (1952). The geology of the Soxth-East Province, South Australia, with special

reference to Quaternary coast-line migrations and modern beach developments, Bull, Ceol,

Surv. §, Aust. no, 29: 1-20.

Warez, % R. {1020}. “The Reptiles and Amphibia of South Australia”, (Government Printer:

Adelaide.

Wanbunc, M. R. (1966). On the water economy of several Australian geckos, agamids and

skinks. Copeia 1966: 230-235.

Werxes, H. C. (1980), On placentation in reptiles. UW. Proc, Linn. Soc. N.S.W. 55: 550-5764.

Wenner, F. (1910). Reptilia (Geckonidue und Scincidae). In “Die Fauna Sidwest-Australiens,

Ergebnisse der Hamburger siidwest-australischen Forschungsreise 1905". Ed. by Michacl-

sen, W. and Hartmeyer, R. Bd. 2, pp, 431-93, (Fischer: Jena.)

Worsert, &. (1963). “Reptiles of Australia’. (Angus and Robertson; Sydney.)

HELMINTHS FROM SOME LIZARDS MOSTLY FROM

SOUTH AUSTRALIA

BY L. MADELINE ANGEL AND PATRICIA M. MAWSon*

Summary

An account is given of helminths from lizards from an area near Port Gawler in South Australia,

including a table of their incidence. Records are also given of trematodes from two Queensland

lizards. Species recorded (all from the Pt. Gawler region unless otherwise stated) are:

Paradistomum. crucifer (Nicoll) (syns. Eurytrema crucifer Nicoll, Paragonimus trachysauri

MacCallum, Cephalogonimus trachysauri MacCallum, Paradistomum muccallumi Johnston) from

Hemiergis peronii, Trachydosaurus rugosus (Pt. Gawler area and from Murray Bridge S.A.),

Tiliqua scincoides (Queensland) , and Varanus varius (Queensland) Oochoristica trachysauri

(MacCallum) (syn. Oochoristica australiensis Spassky (from Trachydosaurus rugosus; Thelandros

kartana Johns ton and Mawson from H. peronii, Phyllodactylus marmoratus; T. trachysauri

Johnston and Mawson from Trachydosaurus rugosus; Skrjabinodon smythi n. sp. From

P. marmoratus. Other helminths recorded are Microphallus sp. from T. rugosus; Trematoda, ? sp..,

from Rhodona bougainvillii; Oochoristica sp. and Baerietta sp. from R. bougainvillii, H. peronii

and P. marmoratus; Skrjabinelazia sp. From P. marmoratus; acanthocephalan cysts from H.

peronii.

HELMINTHS FROM SOME LIZARDS MOSTLY FROM

SOUTH AUSTRALIA

by L. Mapenrse ANGEL anp Patricia M. Mawson*

[Read 8 August 1968]

SUMMARY

An aeeount is given of helminths from lizards froin an urea near Port Gawler

in South Australia, including u table of their incidence. Records are also given of

lrematodes from two Queensland Ligurds. Species recorded (all from the Pt,

Gawler region unless otherwise stated) are; Paradistamum crucifer (Nicoll)

(syns. Eurytrema crucifer Nicoll, Paragonimus trachysuuri MacCallum, Cephala-

conimus trachysanri MacCallum, Paradistomum maccallumi Johnston) from

Hemiergis peronii, Trachydosaurtis rugosus (Pt. Gawler area and from Murray

Bridge, S.A.), Tiliqua scincoides (Queensland), and Varanus varius (Queens-

land); Oochoristica trachysayri (MacQallum) (syn. Oochoristica australiensis

Spassky (from. Trachydesaurus rugosus; Thelandros kartana Johnston and

Mawson from MT. peronii, Phylloductyjlus marmoratus; T. trachysaurt Johnston

and Mawson from Trachydosaurus rugasus; Skriabinodan smythi n. sp. from

P. marmoratis. Other helminths recorded are Microphallits sp. fron T, rygosus;

Trematoda, ? sp., from Rhedona bougainvillit; Oochoristica sp. and Baerietta sp.

from RA. bougainvillit, H, peronii aud P. marmoratus; Skrjabineluzia sp. from

P. marmoratus; acanthocephalan cysts from H, peranii.

INTRODUCTION

During 1967, Dr. Michael Smyth, of this department, undertook an investiga-

tion of aspects of the ecology of lizards inhabiting a coastal strip north of

Adelaide. The parasites of these lizards, and some from the same host species

from different localities, are discussed in this paper. The occurrence of a trema-

tode from two other species of lizards, both from Queensland, is also recorded.

We are very grateful to Dr. Smyth, not only for giving us the viscera for

examination, but also for the regularity and precision of his collection and

records. An aceount of his work is in press.

The study area is a short distance north of Adclaide and the two collecting

centres aré Port Gawler and Middle Beach. The two areas are separated by two

permanent salt water channels. The lizards from each area are listed in Table 1,

with records of parasitism. It will be noted that trematodes were found (except

in oné case) only at Middle Beach; cestodes and nematodes occurred in both

areas. Juvenile lizards seem to be free from helminths. Records were kept of the

sex of each lizard dissected, but this appeared to have no significance in relation

to the infestation by helminths, and has not been indicated in the Table.

Differences in the incidence of parasitism, as well as of the different groups

of helminths, in the different species of lizards, are quite marked, and are

discussed below.

We are grateful to Dr. S. J, Edmonds, of this deparlment, for examining the

acanthocephalan cysts, Our thanks are also due to Mr. John Mitchell, of the

South Australian Museum, for the correct names of the lizards concerned.

* Department of Zaology, University af Adelaide.

Trans, Roy. Soe, S.A. (1968), Vol. 92,

60 L. MADELINE ANGEL ayn PATRICIA M. MAWSON

TABLE 1

Incidence of helminths in the study ures.

The figures refer to the number of lizards dissected or found parasitised. An asterisk mdicates

that the alimentary canal only, not the gall bladder, of these specimens was examined

a UE aE EEE

Number yielding

Lizard Locality Number | Number - - po

dissected parasitiged | ‘Trema- | Cestoda | Nema- jAcantho-

toda toda | cophala

Trachydoxsauras | Pt. Gawler 3 3 a 3 =

rugosa (Gray) Middle Beach! 6 i 1 2 ; | —

Ablepharus greytt Pt. Gawler 5* 0

(Gray) { | 0

Middle Beach) 2 | 9

Hemierqia peronit Pt. Gawler au* 16 os 7 12

(Fitzinger) Li) 26 — 7 eal] 2

~ 10 juv. 0

Middle Beach 13* + — 4

il 37 15 2 84 3

§ juv.

A, devrestensis Pt. Gawler I 0

(Fitzinger) i

Rhadona Pt. Gawler oth 14 iT Ii -—

bougeinvéllii (Gray) :

Phyllodactylus Pk. Gawler .P 1) a 2 1

nearmoratus (Gray ) 1 juv, 0

Middle Beach l [ -- i

PARASITES RECORDED, ARRANGED UNDER THEIR HOSTS

Trachydosaurus rugosus Gray. Microphallus sp.; Paradistomum crucifer

(Nicoll); Thelandros trachysauri Johnston and Mawson.

Hemiergis peronii (Fitzinger). Paradistomum crucifer (Nicoll);

Oochoristica sp. Baerietta sp.; Pharyngodon kartana Johnston and Mawson.

Tiliqua scincoides (Shaw). Paradistomum erucifer (Nicoll).

Rhodona bougainvillii (Gray). Trematoda, 2 sp.; Oochoristica sp;

Baerietta sp.

Phyllodactylus marmotatus (Gray), — Skrjabinodon smythi n. sp., Thelan-

dios kartana Johnston and Mawson; Skrjabinelazia sp.

Varanus varius (Shaw). Paradistomum crucifer (Nicoll).

TREMATODA

Paradistomum crucifer (Nicoll)

(Figs. 1-6)

Eurytrema crucifer Nicoll, 1914, 338, in gall-bladder, Dela fraseri.

Paradistomum crucifer (Nicoll) Travassos, L919, 12; 1944, 256,

Patagonimus trachysauri MacCallum, 1921. 173. im gall-bladder, Trachydosaturus rugosus (syn.

Trachysauris rugosus ).

Cephalogonimus trachysauri MacCallum, 1921, 176, in gall-bladder, Travhydosauras rugasus,

Paradistoma trachysauri (MacCalltmm) Dollfus, 1922, 329.

Paradistoma truchysauri (MacCallum) Dollfus; Johnston, 1932, 64,

Paradistomum maccullumi Johnston, 1932, 64 (nom. nov. for Cephalogonimus. truchysauri

(MacCalhm ).

Paradistumum. trachysaurt (MacCallum) Dollfus; (syns. Paragonimus Wwachysauri MacCallum

and Paradistomum maceallumt Johnston). Malan 1959, 87.

Paradistomum trachysauri (MacCallum) Dollfus; Travassos, 1944, 262.

TIELMINTHS FROM SOME LIZARDS &1

Hosts and Localities: Hemiergis peronii, Middle Beach, South Australia, from

gall-bladder, occasionally in sma!l intestine; Trachydosaurus +tigosus, Murray

Bridge district, South Australia, from gall-bladder and liver; Tiliqua scincoides,

Iyueing Island, near Gladstone, Queensland, apparently collected by Professor

T. Harvey Johnston, in 1918, from gall-bladder (ten specimens from one host);

Varanus varius, Townsville, Queensland, collected by Dr, G, A. M, Heydon in

1927, from gall-bladder (two specimens from one host).

The description is based on 24 stained and mounted specimens from

Hemiergis peronii, with details from living specimens, Notes on the trematode

from Trachydosaurus rugosus are given in the discussion.

TAINS

ANN

Fins. 1-6 Paradistomum ecrucifer, Drawings were made with the aid of a camera lucida, Scales

in millimetres, Figs. 1 and 6 to same scale. Evws not drawn exactly to scale.

Figs. 1-3. 5, 6, from Memiergiy pereniiz 1, 2, 6, adults, showing variation in form, and in

extent of uterus, (2. flattened); 3. cirrus sae, fram specimen shown in fig, 2, enlarged;

5, youny adult, excretory system from living specimen. Fig. 4, type specimen redrawn

(fronr Delma frasert). ¢, cirrus; eb, excretory bladder; mg, Mehlis’ gland; 0, ovary;

bie, prostiie gland cells; rs, receptacuhim seminis; sv, sernimal vesicle: vd, vitelline

cluct.

62 lL. MADELINE ANGEL any FATRICIA M. MAWSON

The numbers of trematodes in the infected H. peronii ranged from I-15, and

generally included juveniles as well as adults. One gall-bladder contained 65

specimens, but all were very young, Nearly all specimens were found in the gall-

bladder, but a few were in the liver, and four were taken from the intestine. OF

these last, three were in two hosts whose galJ-bladders were uninfected, and the

fourth was in a lizard whose gall-bladder contained twelve trematodes. In one

lizard only, the trematodes were found not only in the gall-bladder (thirteen).

but alse in ducts which appear to be pancreatic, running through the clongated

panereas from the gall-bladder to the intestine (fifteen).

Specimens were fixed, under a coverslip with only slight pressure, in formol

acetic aleohel, stained in Van Cleave’s combination haematoxylin stain, and

mounted in Depex.

DESCRIPTION

Shape: Elongated ov leaf-like when living, fixed specimens with bluntly

rounded posterior and often somewhat elongated anterior end. Colour pale pink,

with caeca yellow.

Rody 0748-2890 mm long by 0:306-1 -496 mm wide. Cuticle not spined,

Oral sucker rounded or slightly oval, 223-494 » (average 342 4) Jong by 129-

493 yp, (330 p) wide. Acetabulum rather inconspicuous, rounded, 170-353 4

(235 «) long hy 141-306 » (239 ») wide; anterior border near anterior third of

body, Ratio of width of oral to width of ventral sucker 1:0:7 to 1:0:9. Pre-

pharynx short; pharynx 50 by 50 «—1L0 by 100 », dorsal to oral sucker or partly

posterior to it; oesephagus short; caeca wide, sinuous, extending nearly to pos-

terior end of body.

Testes symmetrical, at sides of acctabulum, rounded in living, may he

irregulae in Gxed specimens, equal or sub-equal, 71-165 ~ (110 4) long by 5Y-

176 p. (106 ») wide. Cirrus sac 92-184 p» long by 45-100 » wide, between suckers,

at an angle to left of mid-line, Seminal vesicle internal, much coiled. Cirrus

inconspicuous, slightly coiled, surrounded by diffuse prostatic cells, Cenital pore

median, near posterior border of oral sucker.

Ovary post-ncetabular, sinistral, irregular, §2-223 » long by 71-188 ,» wide.

Receptaculum seminis mostly dorsal ta ovary, may be posterior or to cither side

of it, 47-141 » (96 ») by 59-212 » (107 »). Mehlis’ gland irregular, generally

posterior to ovary and slightly to right. Laurer’s canal not seen. Uterus varying in

extent from condition shown by Nicoll, with relutively few eggs (Fig. 6), to one

in which it fills all of hind-body and an area in front of acctabulum; passing to

right of cirrus sac and opening by museulur nctraterm at genital pore.

Eygs variable, larecst 40 » by 22 p.

Vitellaria extracaecal, restricted to middle of body, reaching level of posterior

third of acetabulum anteriorly; 270-764 » (476 «) in extent; Jobules variable,

some moderately large. Transverse yolk ducts widening at junction in mid-line,

but forming no obvious yolk reservoir.

Lxeretory bladder elongate, main arms leaving anterior end, Exerctory pore

terminal.

DISCUSSION

The muture trematodes found in Hemiergis peéronii in this study show a

considerable variation in size, ranging from 0-748 by 0-374 mm to 2-890 hy

1258 mm, and in appearance. The trematodes found in the pancreatic ducts

were greatly elangated, while the worms from the gall-bladder tended to be

TELMINTHS FROM SOME LIZARDS 63

foliate, As immature specimens are often present with the adults, some of the

differeices among egg-beuring adults ire probably due to differences in age. For

example, the testes and ovary may be as large in small worms as in much larger

ones (Figs, 2, 6) giving a disproportionate emphasis to these organs in the small

specimens, The specimen shown in Fig, 1 shows the acetabulum very near the

oral sucker; & specimen of similar size from the same host showed a similar

arrangement of eggs, but the body was more elongated and the acetabulum was

in the more Gypical position, Even among specimens of the same size there is

great variation in the number of eggs, and this leads to differences of form andl

general appearance, In the original description of Paradistomum crucifer, Nicoll

stated that the uterus, especially in the less mature specimens, had a charavteristic

cruciform course, but that in more mature specimens this arrangement was, to

some extent, obliterated. Among the sinall trematodes of my collection. some

(Fig. 6) have a uterus showing the crucitorm course, while in others (Fig. 2) the

eges form a inore or less solid mass in the hind-body and sometimes anteriorly

to the acctabulum, so that most of the organs are obscnred, Even among the

larger specimets there is alsoa good deal of variation in the number of eggs.

By courtesy of the Director of the School of Public Health and Tropical

Medicine, Sydney, one of us (L.M.A.) has been able to examine the type of

P. crucifer. Some of our smaller specimens are very similar to it in appearance,

aml one of them is almost identical with it, not only in overall size, but in the

sive and arrangement of the organs. The trematodes from Hemdergis peronii

must be regarded as Paradistomum ertcifer.

Although the range of measurements given by Nicoll shows that his speci-

mens were umformly smaller than the trematodes found in H. peronit, Nicoll's

description was based only on “a few" worms. In addition, all measurements

given by Nicoll are consistently smaller than those now made on the type (Table

2), It is possihle that this is due to some fattening of the specimen over the

years; it 1s also possible that Nicoll had made a mistake in his scale.

MacCallum (1921) found about twenty trematodes in the gall-bladder of a

stump-tailed lizard, Trachydoseurus rugesus, which died in the New York Zoo,

He stated: “there were three different sizes among them, which, with the decided

differences in form, would almost make it necessary to divide them into three

species, but as they are in many particulars alike, and for brevity’s sake, we shall

describe them as one species”. This he named Paragonimus trachysaurt, Later in

the same paper MacCallum stated that he had found two different trematodes in

the lizard, one being P. trachysauri; the second he described as Cephalogonimus

frachysouri. Malin (1939) regarded the two as identical, and thus synonymous

with Paradis!omum trachysauri, and this has been accepted by later workers.

Although the largest specinens frotn H. peronii are smaller than the measure-

ments given by MacCallum for Paradistomum trachysauri, the measurements of

ovarics and testes are comparable, the ratios of the suckers appear similar, and

the specimens resemble the one figured hy MacCallum. Differences in fixation

could gecount for some differenves in body dimensions, as also could the relative

sives of the hosts*. There seems no reason ta regard the specimens from Hemieryis

perenié as distinct from those deserihed by MacCallum trom Trachydosaurus

gugosus, so that Paradistomum trachysauri must become synonymous with

I’. crucifer, (It should be noted, however, that P. crucifer as now interpreted

* It is of interest to note that Delma frasert (the type host of Paradistomum orucifey), a

yeapodid lizard, and Temicrais peronil are much smaller than Trachydosaurus ttigosus. 1

ave Tut Uisseted D. freseri, ut if can be expected that its gall-bladder is very lithe Larger

than that at If neronit, which is much smaller than that ef T. rugasts,

EL anp PATRICIA M. MAWSON

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HELMINTHS FROM SOME LIZARDS 6n

shows considerable variation in size and general appearance. Variation has

already been reported for another species of Parailistomum by Dollfus (1922.

328, footnote), who referred to the great morphological variations he had found

in P. miutabile (Molin).

This is the first Australian record of 4 trematode from « lizard since

MavCallum’s report of P. trachysauri. In 1932, Harvey Johnston reported that

“a mimber” of 7, rugosus had been searched for parasites from time to time, but

thut no trematodes had been found, After P. crucifer was found in H. peronii in

the present study, as many T. rugosus as possible were examined. Jt was not until

this paper was completed that P. crucifer was found in this host, in two lizards

from the Murray Bridge district. (Of the forty-five T. rugasus dissected only nine

were from the study area north of Adelaide. The remainder came from a number

at different localities.) In the first lizard there were 34 trematodes in the gull-

bladder smd 12 in the liver. In the second, there were 56 living and a number of

disintegrating worms (which were mere collections of eggs) in the gall-bladuer,

and 13 living worms in the liver, The lizards had been in captivity for almost two

months when they were dissected, All the trematodes were mature, but great

ditterences in size were found. The specimens were fixed in formol acctic alcohol

under a coverslip. Virtually no pressure was required to fix them Rat, so that the

measurements are not truly comparable with those of the specimens from

A. peronii (which were fixed with slight pressure) but would probably have been

greater (certainly in the width of the body) had the worms heen fixed with the

same pressure. Measurements of the largest and the smallest specimens fram the

first infected lizard aré given in Table 2.

The following observations were made on the living trematodes fram

T. mugosus. The caeca appear hright yellow (due to bile), and contain many

crystals, as recorded by MacCallum. These are tetragonal in shape, and vary in

size from fine slender crystals to forms up to 223 by 35 by 35 w. (Crystals were

not present in P. erucifer from H. peronii; this is presumably due to a difference

Hi composition of the hile of the two lizards.) The part of the hody which is not

coloured by bile or obscured by eggs is pale pink in colour, Twenty mature eggs

tukem at random from the liver washings measured 34-39 « (37 «) hy 21-24 p»

(22 4).

P. crucifer fram both hosts was very sensitive to changes in the medium in

which it was kept. In 0-65% suline it very quickly hecame swollen and <¢lied. It

vould be kept alive in bile at 6°C for several days.

LIFE HISTORY

The only species of Parudistomum for which investigations on the life-history

have been recorded is P. mutabile (Molin). Timon-David and Timon-David

(1967) infected Helicella arenosa experimentally and obtained brevicercous

xiphidiocercariae, (HH. arenasa was not the normal host since it did not occur on

the islands on which Paradistomum mutabile was commonly found in livards).

The cyst stage was not found, but the authors thought it probable that a second

intermediate host is necessary, and suggested an isopod or an insect.

In the present study no information could be gathered on the life-history of

P. erteifer, The only land snails found in the study area are Avstrosuccinee

australis Ferrusac, Omegapilla australis Anvas, Australbinula margaretae Cox and

Paralaoma stabilis Iredale, The last three of these ave very small snails and quite

diffieult te find in the litter in which they occur, (P. stabilis was not. in fact, found

66 J,, MADELINE ANGEL ann PATRICIA M. MAWSON

in the open at all, but was recovered from the stomachs of lizards, in which it

occurred quite often). It was therefore not possible to conduct any trematode

life-history studies with these molluscs, Attempts were made to infect Austro-

succinica autralis (collected from another area) but it proved impossible to keep

the snails alive Jong enough to obtain any results,

It will be seen from Table 1 that whereas fiftcen of seventy-one Hemiergis

peronii from Middle Beach were infected with Paradistomum crucifer, none of

ninely-nme of these lizards from Port Gawler harboured the parasite. We can

suggest no reason for this.

If, as seems likely, a second intermediate host is necessary, it is to be sought

among the animals listed by Smyth (1968) as found in the stomachs of H. peronii.

Of these, the most common are weevils, free-living mites (very small species),

ants, cockroaches, moths and snails.

Microphallus sp,

Host and Locality. Trachydosaurus rizgosus, Middle Beach.

One specimen, in upper small intestine.

Trematoda, ? sp.

Host and Locality, Rhodona bougainvillii, Port Gawler,

One specimen, in Intestine.

CESTODA

The authors, neither of whom is a cestodologist, are greatly indebted to

Dr, John Hickman, of the Zoology Department, University of Tasmania, for

identification of Buerietta sp, and verification of Oochoristica spp. Further identi-

fication of the material will be made by Dr. Hickman.

Oochoristica trachysauri (MacCallum)

(Fig. 7)

Tuenia trachysuuri MacCallum, 1921, 229.

Oochovistica trachysauri (MacCallum), Johnston, 1932, 65.

Oochoristica australiensis Spassky, 1951, 547.

Host and Locality. Trachydosaurus rugosus, Middle Beach, South Australia,

This species was fully described by Johnston (1932). Spassky (1951) con-

sidered Johnston’s specimens different from those of MacCallum and proposed

for therm a new species, O. australiensis. The material now examined, all from

one host, shows similarity to all earlier collections—some specimens with an

obvious rostellum, some with rounded anterior end; the mature segments vary

(sometimes abruptly, fig. 7) from more or less square to elongate. It is considered

that all helong to the same species.

Oochoristica sp,

Hosts and Locality, Rhodona bougainvillii, Hemiergis peronii, Phyllodactylus

marmoratus, Port Gawler.

More than one species may be present.

Baerietta sp.

Hosts and Locality, Rhodona bougainvillii, Hemiergis peronii, Port Gawler.

The cestodes from these hosts arc similar and probably belong to the same

specics.

HELMINTHS FROM SOME LIZARDS 67

a../

Soa

300

Fig. T. Ovochoristica trechysauri, part of strobila showing variation in shape of segments.

Figs, 8-10, Thelancros kartana, 8, anterior end; 9, vontral yiew of male tail; 10, female

tail, Figs. 11-14, T. trachysauri, 11, anterior end; 12 and 13, yentral and lateral views

of male tail: 14, tail of female. Figs, 9, 12, 13, to scale beside 13, figs. 8, 10, 11 and

14 to seale beside 10.

NEMATODA

The nematodes taken from the lizards at Port Gawler show an interesting

host distribution. None were found in Ablepharus greyii or Rhodona bougain-

villii. Oxyurids were found in Hemiergis peronii, Trachydosaurus rugosus and

Phyllodactylus marmoratus, and in almost all cases each of these hosts carried

only its own species. The exceptions were two P. marmoratus, in which were

Thelandros kartana, typically present in H, peronii, Although H. bougainvillit is

apparently free of nematodes in this area, it carries the same cestode species as

does #7, peronii and these species are in turn different from that from T. rugosus.

This difference in parasites may be duc to a high degree of host specificity among

the oxyurids, and perhaps of resistance among the lizards, or it may be explained

in part by the different niches occupied by the lizards. Dr, Smyth has informed

us that R. bougainwillii and H, peronti live mainly under certain bushes, A. greyii

and P. marmoratus mainly under wood, stones, etc., and T. rugosus (a much

larger lizard) may be found in either habitat. Most of the lizards are diurnal and

insectivorous; T. rugosus is diurnal and predominantly vegetarian; P. marmoratus

is nocturnal and insectivorous. Oxyurid eggs might be expected to be quite

common over the surface in the area, especially those of Thelandros trachysauri

which occurs in hundreds in each host animal,

68 L. MADELINE ANGEL sxp PATRICIA M. MAWSON

Thelandros kartana Johnston and Mawson

(Figs. 8-10)

Thelandtag kartana Jolinston and Mawson, 1941, 145; from Iemiergis peronii, Kangaroo

Slana,

Hosts and Localities. Hemiergis peronii, Port Gawler, Middle Beach.

Phyllodactylus marmoratus, Port Gawler.

Thelandros kartana was taken from 22 specimens of I. peronii from Middle

Beach and 27 from Port Gawler. Female worms with eggs were taken from two

P. marmoratus, No more than four worms were present in any one lizard, and

usually only one or two. They occurred almost always in the short caecum at

the junction of small and large intestine. The new specimens have been com-

pared with paratypes of T. kartana and found to agree closely. The original

description can now be augmented by Figures 8-10 and measurements in Table 3.

The measurements of the specimens from P, marmorattus are within the range of

those from H. peronii,

Figs. 15-21, Skrjahinodon smythi, 15 and 16, anterior end of specimens in relaxed and con-

tracted states respectively; 17, posterior end of male; 18, ventral view of cloacal

region, male; 19, tail of female; 20, part of spike of female tail showing ‘spines’; 21,

egg, Figs. 17 and 18 to same scale,

HELMINTTIS FROM SOME LIZARDS 42

Thelandros teachysauri Johnston and Mawson

(Vigs. 11-14)

T Relandyye 5 frachysairt Jobeston and Mawson, 1947. 24; fron Traehytlosanris rupovun,

Jselehnue,

Host and Localities. Trachydosaurus rugosus, Port Gawler, Middle Beach.

Thelandros trachysauri has been found in nearly all T, rugosus dissected in

this Department—more than thirty-five from various places. It is present in very

large numbers in the middle and posterior parts of the large intestine. The col-

lections [rom the Port Gawler area agree with the original description, which ean

now he amplified by measurements of more specimens (Table §) and

by Figures 11-14,

The lateral alae, present in both sexes, are not very wide; in the mule they

extend from the posterior oesophageal region to near the cloaca; in the female

they are restricted. commencing just posterior to the oesophagus and reaching to,

not past, the vulva. The nerve ring is 190-210 4 from the head in females, rather

more anterior than described earlier. The eggs are oval in shape, contain a bent

larva, and measure L0G « by 59-60 p.

Skrjabinodon smythi n.sp.

(Figs, 15-21)

Type Host and Locality. Phyllodactylus nurmoratus, Port Gawler. Other

Jovalities: Middle Beach, Chowilla, Loxton, Lock, all in South Australia,

Type 6 and allotype @ will be deposited in the South Australian Museum.

This species appears to be common in P. marmeratus throughout southern

Australia, although another species (unpublished) apparently takes its place in

northern parts of the state, tt has been taken from fourtcen of seventeen host

animals examined, In most cases there are about 6-8 worms in cach host, but in

some there are more, the greatest number being sixty-seven of which 35 were

males, 7 females with eggs, and 22 females without eggs, All ovcur in a mass in

the small caecum at the origin of the large intestine. Where few worms are

present, the gravid lemales are stuffed with eggs, but where there are many

worms, the eggs are few,

There is an apparent variation in the position of the vulva and excretory

pore in the females. These appear to be oesophageal structures in some collec-

tions and well behind the oesophagus in others. This however is largely dependent

on the degrec of contraction of the body. It has been noted that those worns in

which the excretory pore (and vulva) are oesophageal are stiff and barreltike

in appearance, with strongly marked ringed or ruched cuticle, whereas in flaccid

specimens with smooth cuticle these two pores are further back, This is shown

hy the measurements, in Table 8, for $. smyth! as, although the total length

measurements show a great variation, the length of the ocsophagus and the [gil

spike do not. In this Table, measurements are also given of some very flaccid

specimens from Chowilla.

Lateral alae are present in both sexes, from about the midlength of the

oesophageal region to the level of the anus, The amphids are large. mere promi-

nent, and slightly further back, than the four large cephalic papillae. The three

lips are bilobed in the female and single in the male. At the anterior end of the

oesophagus of the male are three small teeth, not present in the female. The

position of the nerve ring is not clear in mast specimens, The excretory pare is a

transverse slit with cuticularised lips and it lies posterior ta the wcsuphagus dn

relaxed specimens, It is more posterior in the male than in the female,

The male is without caudal alae; the male tail spike has a few very minute

spines. There is no spicule, but cuticularised projection of the cloacal wall is

70 L.. MADELINE ANGEL ann PATRICIA M. MAWSON

TABLE 3

Measurements of Thelandros kartina, T. trachysaurié and Skrjubinadon wmylhi-

All nvusurements ara in jo unless otherwise indicated. The tail of the male S. vnylht fram Part

Gawler is broken.

Hpuciex T. kartana T. trneh yaar. | Si smytht

Locality ' Port Gawler | Middle Bouwh | Port Gawler | Port Gawler Chowilla.

ae of wonlraatior (eontracted) (rotaxedl) (flaccid)

Males:

myth (111) 163-0 | 1628-5 1-40-1-78 2-4-2-5 1) 75-23-10

vosophapus 320-480 =| 9 5A0-6H0) 220-300 | 250-275 200-31)

antr. end-cxer, pore | 50-1000 Oo0-OtN) SAU-8 70) 600 650 400-490

tail spike 50-74) \ 100-120 ahU-410 4110420) 450. FAO

apicule 65-75 100 150 — —

Yemales: | |

length (rn) 2-1-6-5 3°3-3-5 31-4 Fi 4-0-7°-0 | 6-9 Bed

beso phage 400-L200 820 980 ANO-AfiN) 470-500 440-600

antr. end-exer, pore 520-1400 00-1000 200 40 5HU-GH) 550-700

-yvulva 3-0-4°8 1-2: 350-45) 650-TH0 GON-7580

tuil 100-150 340-450 1390-1500 1350. 1800 15-1600

taal spike —_ 1900-1050 800 -1050 HO0=LO00.

present, The preanal, adanal, and postcloacal papillac are almost evenly spaced

on the ventral surface, Dorsulaterally to each adanal papilla is a papilja-like

extension into, but not lifting, the cuticle, and terminating at a tiny pore, These:

are probably the phasmids.

The tail of the female bears about 7-9 irregularly spaced projections which

are more digitiform than spinous (Fig. 20). The vulva is a transverse slit, without

thickened lips. The eggs are 150-165 » long, with a plug at cach end and slightly

flattened on one side, The most mature eggs contain a larva 120 p long.

The genus Skrjabinodon was proposed by Inglis (1968, 179) for some species

which had been placed in Parathelandras Baylis but which differed from the

type species of this genus, and from other species which he described at that

time. The species attributed to Parathelundros are all from Australian frogs, and

those to Skrjubinedon are from lizards, mostly from places other than Australia,

but one, §. oedurae (Johnston and Mawson) from an Australian lizard. The

present specimens agree generally with other species of Skrjabinodon and with

the gencric diagnosis proposed by Inglis, except in the two characters which he

marks as doubtful, namely the absence of onchia at the anterior end of the

oesophagus {present in the male of $. smythi) and the lip shape, which does not

uppear to be bilobed in the male of S. smythi.

The species is differentiated from S. anolis (Chitwood), the only other

species of the genus in-which the spicule is absent and the female tail bears large

“harbs”, by the shape of the barbs, the larger size of the eggs, the longer oeso-

pligus,. the more anterior position of the anus in the female, and by the rather

different spacing of the male caudal papillae.

Skrjabinelazia sp.

Host and Localities; Phyllodactylus marmoratus, Port Gawler, Middle Beach,

The material available consists of several female worms from the intestine

of five geckos. No males have heen found.

The worms are large, up to 18°3 mm in length, and the cuticle at each end

is markedly inflated. The anterior end bears four large papillae and two amphids,

There are no lips, The mouth, more or Jess circular, Ieads into a short buccal

HELMINTITS FROM SOME LIZARDS Tl

cavity from the walls of which project 2 ring of tiny teeth, like an internal leaf

erown. The oesophagus, 1:2 mm long, widens towards its posterior end, but is

not obviously divided into muscular and glandular parts, The nerve ring is at

about one third its length from the head, and the excretory pore at the same

level. Cervical papillae were not secn, The posterior end narrows suddenly 300 p

behind the anus, and the bady proper ends in a short spike about 110 j long.

The inflated cuticle extends behind this for about 50 p.

The vulva lies shortly behind the nerve ring, about 490 » from the head,

The eggs are large, sub-spherical, 90 » in diamcter, and contain a coiled embryo,

The egy shell is thin and apparently not rigid, as some change their shape under

pressure.

These worms are very similar in appearance to those of the genera

Skyjabinelazia Sypliakova und Salobrella Freitas. The species of these genera

differ, as far as the female is concerned, in the absence or presence, respectively.

of lips, so the present specimens are identified as Skrjabinelazia sp. In the absence

af inales, no further identification is attempted. No species of Skrjabinelazia, or af

Salabrella, has so far been recorded from Australian hosts,

ACANTHOCEPHALA

Host and Locality: Wemiergis peronii, Port Gawler (2) and Middle Beach

(3),

Five acanthocephalan cysts were taken from the mesenteries of these hosts.

They have been examined by Dr. S. J. Edmonds, who has kindly given the

following information.

All the cysts appear to belong to the same genus, In only one is: the introvert

extended enough to permit an estimate of its measurements, The. ellipsoidal cyst

is approximately 800 » long by 300 » wide, The introvert, which is cylindrical

and bears numerous hooks, is about four-sevenths extended, and is estimated 45

1100 » long and 260 » wide, The number of hooks is hard to estimate, possibly

about 80 rings cach of 26 hooks.

The only acanthocephalan so far recorded from Australian reptiles is

Sphaerechinorhynchus rotundocapitatus (Johnston, 1912), from the black snake

Psewdechis porphyriacus Shaw, The cysts from H. peronii da not belong to this

species,

REFERENCES

Curiwoon, B. G., 1934, Reports on the collections ohtained by the first Johnson-Smithsyonian

sleep-sea expedition to the Puerta-Ricun Deep. Two new nematodes, Smithsonian Mise,

Coll. (Publ, 3243) 91 pp.

Duturvus, R, P.. 1922. Variations dans: la forme dw corps, la position et Ia forme des tostionles

chez Dicracoelinm lanceolatum (Rudolphi), Bull, Sue. zool, France, 47, pp. 312-342.

Faenmas, J. F. Teixeira de, 1941. Sobre um intercssante nematodes parasita de reptil (Spiru-

roidea) Mem. Inst. Osw, Crnz., 28, po, 239-245.

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72 L. MADELINE ANGE! ann PATRICIA M, MAWSON

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TWO SPECIES OF SACCOGLOSSUS (ENTEROPNEUSTA)

FROM SOUTH AUSTRALIA

BY I. M. THOMAS*

Summary

A new species of the Enteropneusta, Saccoglossus aulakoeis is described. It possesses a deep,

dorsal, longitudinal groove on its proboscis. It is compared with three previously described

members of the genus, which have similar grooves. One of these species, Saccoglossus otagoensis

(Benham) has been found in the same vicinity. This is a new record for Australia. The two species

however, occupy different habitats, S. aulakoeis being found in coarse sand and shellgrit in amongst

the roots of Zostera while S. otagoensis, in this locality, has been found only under stones at or

below the level of low water spring tides.

TWO SPECIES OF SACCOGLOSSUS (ENTEROPNEUSTA)

FROM SOUTH AUSTRALIA

by I. M. Troxtas*

[Read 8 August 1968]

A new speeies of the Enteropneusta, Saccoglossus aulakoeis is deseribed. LU

possesses a deep, dorsal, longitudinal groave on its probosvis. It is campared with

three previously described members of the genus which have similar grooves.

One of these species, Saccoglossus otagoensis (Benhani) has been found in the

same vicinity. This is a new record tor Australia, The two species however,

occupy different habitats, $, aulakocis being found in coarse sand and shellgrit in

amongst. the routs of Zostere while 8. ofagoensis, in this locality, has been found

only under stones ut or below the leyel of low water spring tides.

During an investigation to determine the distribution of Saccoglossus apan-

tesis. (Thomas, 1956) in South Australia, two other members of the genus have

been found. One of these is a new species and the other a new record for Aus-

tralia..

Material of both species was fixed in Bouin’s-in-seawater and scctioned at

10 4. Sections were stained either with Ehrlich’s haematoxylin and eosin or with

Mallory’s triple stain. The latter was very eflective for showing the basement

membrane and the skeletal parts derived from it. Proboscis skeletons were

extracted by maceration of unfixed material in 4% borax for several days. After

carelul cleaning by brushing, they were stained in aniline blue and mounted in

cedarwood oil.

Saccoglossus aulakoeis n.sp.

The trivial name is descriptive of the deep, median grooye on the dorsal side

of the proboscis (avAaé = a groove or furrow),

Specimens have been found at Port Willunga, South Australia (lat. 35° 16°

50” S, leng. 138° 27’ 20” E) in a shallow tidal pool about half an acre (0-2 hec-

tares) in extent about a quarter of a mile (400 m) south of the rnins of the Port

Willunga jetty. The pool is covered to a depth of five or six feet (about 1-6 m) at

high spring tides and about half its floor is exposed at low spring tides.

Zostera tasmanica G, V. Martens grows sparsely in small patches at about the

level of normal spring lows and in amongst the roots of some of these patches, the

enteropneust has been found, Il is by no means common, The soil over the under-

lying rock is not more than an inch or two-deep and consists of sand and shell grit

with little or no mud, The animal has not been found in sand without Zostera and

many patches of Zostera do not contain it. One specimen has been found in

amongst the roots of Cymodocea antarctica (Labill.) which grows abundantly in

slightly deeper water in the pool. The pool is partly protected by a low, rocky

elevation (covered at mid-tide) on its scawards side, but it is open to the sea at

all states of the tide.

At Brighton, (lat. 35° 02’ S, long, 138° 31’ E), about thirty miles north of

Port Willunga, after a heavy storm, some masses of Zoster roots were washed

* Zoology Departnient, Uniyersity of Adelaide,

Trans. Roy. Soc. S.A. (1968), Vol. 92.

T4 T. M. TITOMAS

ashore on a sandy beach, In amongst the roots were found four fragments of

enleropneusts similur in size to the Port Willunga specimens, One of these frag-

meyts included a portion of a pruboscix with a deep dorsal groove, Sectioning of

this material has confirmed that it is $8. aulakoeis.

The species agrees with the diagnosis of the genus Saccoglossus Schimke-

witsch, 1892 (syn, Dolighoglossus Spengel, 1893) iu the possession of the follow-

ing characters; (a) proboscis very long, (b) collar about as long as broad, (¢)

lateral genital folds present but dorsal gonads absent, (d) gill pores small lait dis-

tinct, (e) perihacmal spaces present, (£) posterior oesophageal pores present,

anterior ones absent,

EXTERNAL FEATURES

S. aulakoeis (Fig, 1) is a species of small to moderate size, Full grown, intact

specimens are diffeult to obtain but a specimen of reasonable size which las heen

relaxed by the method recommended by Ledingham und Wells (1942) (isotonic

magnesinm chloride). had the following dimensions, Proboscis 16 mm; collar 2+5

mm; branchial region 5 mm; oesophageal region 2 im; hepatic region 13 mm;

intestinal region 24 mm: genital region (which overlaps from the branchial to the

hepatic region) 10 mm. The gonads were not folly mature.

The proboscis is orange-red and its base, usually hidden by the collar, slightly

darker, On this the preoral ciliary organ (Fig. 1) shows up as a U-shaped yellow

line. The anterior three-fifths of the collar is similar in shade to the proboscis, The

remaining two-fifths bears two broad, slightly elevated bands, paley and yellower

and separated by a narrow groove, the posterior wall of which shows wp as a still

paler band. This latter band corresponds to zone four in the histological divisions

of the collar epidermis (Fig. 6).

Dorsally and laterally, the branchial region is slightly browner than the pro-

boscis and is beset with irregular flecks of yellow-orange. These are groups of

eland cells (epidermal glandular eminences, Figs, 1, 7, 8 and 10), The ventral

musculature of this region is again similar in shade to the proboscis. but with

narrow and slightly paler transverse striations. (Fig, 1), The oesophageal region

is similar in colour to the dorsal branchial region hut the flecking, dorsally and

laterally, is denser. In the hepatic region, the body wall is somewhat translucent

and the surface flecks on dhe epidermis are smaller and sparser. Tn livmg soeci-

mens, the deep brown colour ef the lateral saccules of the liver region of the

alimentary canal show through clearly. The intestinal region, too, is transhicent

and the sandy gut contents show through, The flecks on the epidermis are still

evident but ure more widely spaced and are smaller than they are on the

hepatic repion.

In preserved specimens, the proboscis tapers slightly throughout its length

but in ving aud fully relaxed material, it has the form of « very elongate pear.

heing appreciably thicker at its base. It has a deep mid-dorsal groove which, at its

deepest, is a third to a half of the diameter of the organ and it extends almust to

the anterior extremity (Figs. 1 and 4). Posteriorly, in the vicinity of the proboscis

complex, it is shallower so that on the basal face of the proboscis, its depth is

about one-fifth of the diameter. The preoral ciliary organ has a pattern similar to

that in other members of the genus in which it has been described (Brambell and

Cole, 1939; Brambell and Goodhart, 1941; Thomas, 1956). It is U-shaped and lies

an the base of the proboseis closer to the stalk than to the outer border of the

hese. The dorsal tins of its arms, however, are detected inwards and herein it dif-

fers slightly from the organ as it is pictured by Brambell and Cole (1939b) in 5,

suber (syn. 8. cambrensiv Burdon-Joncs and Patil, 1960) in which the arms are

Fig, 1. Saccoglossus aulakoeis, anterior end, lateral view.

Fig. 2, Oesophageal region, dorsal view.

Fig. 3. Transverse section, dorsal proboseis yroove.

Fig. 4. Transverse section, proboseis.

Fig. 5. Proboscis skeleton, yentral view,

Fig. 6. Zones of collar epidermis, b., body of proboscis skeleton; b.I., basement layer: ¢., collar;

ce, 1 to 5, zones of collar epidermis numbered from anterior end; d.pr.. dorsal groove. of pro-~

hoseis; e.gLe,, epidermal glandular eminence; g.p,, gill pore; g.r, genital ridge; h., horn of

proboscis skeleton; i:c.m.pr., inner circular muscle layer af proboscis; k,. keel: Lb)., lateral blood

vessel; lm.pr., longitudinal muscles of probosvis; m.d.bl., mid-dorsal blood vessel; n.J., nerve

layer, O.c,m.pr,, outer circular muscle layer of proboscis: o.p., sesaphayeal pore; p.c.or., preoral

ciliary organ; pr.proboscis; pric., proboscis coclom; pr, sk,, proboscis skeleton; s., sole, formed

hy ventral longitudinal muscles; th.n.l., dorsal thickening of nerve layer: v.gle., vacuole of

epidermal gland cell,

v6 1. M, THOMAS

straighter, There is a single, slit-like proboscis pore on the stalk, close to its junc-

tium with the collar and slightly to the Jeft of the mid-te.

The collar is longer than its diameter in the proportion nf about 1/0 to 0-77,

It is also longer dorsally than it is veatrally im the proportion of about 1:0 to 0-95.

This is dne partly toa slight retraction of the ventral part of the anterior Hange

under the mouth and partly to the dorsal part of the posterior lange extending

farther over the branchial region than does the baistrale part (Fig. 1). The pos-

terior two-fifths, approximately, of the collar bears two broad, slightly elevated

bands of about equal width and separated hy the narrow groove already

mentioned,

Anteriorly, the branchial region is somewhat quadrangular in transverse sec-

tion as most of the main musculature is veutral in position, The ventral surface of

the body thns toms a broad sole on which the animal creeps (Fig, 1). Farther

back in the branchial region, the sole has a shallow median ventral groove { Figs,

7 and 8).

There are from twelve to twenty-five pairs of small gill pores set in shallow

dorse-lateral depressions. The anterior two or three pairs are coyered by the pos-

terior flange of the collar. The Just five or six diminish in size and get successively

closer to the mid-dersal line (Fig, 2). In small decinatte) specimens the first

gonads are seen wbout half-way alony the branchial region but in filly mature

specimens, they hegin closely behind the collar, Depending on the maturity of the

specimen, the gonads form more or less conspicuous genital ridges (Fig, 7). In

immature specimens (Fig. 8) the ridges are relatively inconspicuous, These

extend to the beginning of the hepatic region in mature specimens and end rather

abruptly. In immature specimens they do not extend as far back, In the branchial

region, the gonads are rather lateral in position but behind the gill pores they

hhecume more dorso-lateral oe 2).

About 1 mm behind the last gill pore lie the ocsophageal pores (Fig. 2).

These are arranged in two rows one on each side of the mid-dorsal line. The:

pores number from two to seven or eight pairs but the more anterior ones are not

patent. The specimen shown in Fig. 10 hud two patent (1 and 2) and three rucdti-

mentary (3, 4.and 5). The disposition of the rows is variable. Seven specimens

were examined in this regard, In three of these the rows were almost parallel to

each other and ta the mid-dorsal line, in three they were slightly divergent. the

more posterior being the farthest apart and in one they diverged at an angle of

about 30°. This specimen is shown in Fig. 2. The larger and patent pores are the

most posterior. In the anterior oesophageal region the sole (Figs. 7 and 8) is as

broad as it is in the posterior branchial region but begins to narrow towards the

posterior cnd of the oesophagus. The hepatic region can he recognized hy the

more or less regular lateral dilatations of the alimentary canal which can he seen

through the somewhat transparent body wall, The sole here narrows further and

becomes less in width than the dorsal part of the body though its lateral bulges

aré still apparent, In the intestinal region the hody tapers slightly to the terminal

anus and the ventral musculature diminishes in amount so that the sole disappears

sind the hody is rounded in transverse section.

INTERNAL ANATOMY

Praboscis

The glandular and ciliated epidermis of the praboseis extends to the bottom

of the dorsal groove and beneath it Hes the nerve layer (Fig. 3 and 4), Over most

af the proboscis, this is about 12 to 14 « thick, At the bottom of the proboscis

groave there is. a marked ridge in the nerve layer so that at this puint it is 30 to 32

TWO SPECIES OF SACCOCLOSSUS 77

Fig. 7. Succoylossus aulukodis, transverse section of nearly oature fermale in first region of

oesophugns.

Vig. §, As Fig. 7 but of immature female.

Fig. 9. Longitudinal sagittal section of proboscis conyplex.

Fix. 10. Longitudinal section showing oesophageal pores. hic,, buccal cavity; c.. collar: c.co..

collar cord; c.cl., collar coelom; ¢.s, central sinus (blood sinus); d.bly.. dorsal bload vessel;

dium. dorsal longitadinal muscle; dn.c. dorsal nerve cord; e.gle., epidermal glandular emi-

nence; gl,. glomerulus; hev., heart vesicle (pericardium ): ic.m.pr., inner layer of vircular muscles

of proboscis: |m.pr., longitudinal muscles of proboscis; Lin.p.c.. longitudinal muscles of peri-

huemal cavity; Lw.d.gr., vertical section through wall of dorsal groove; m., mouth; n.ljpr., nerve

Jnyer of proboscis; o.c.mpr,, outer layer of circular muscles of proboscis; oes, 1, first region of

oesophagus: o.p. 1 and. 2, first and second oesophageal pores (patent); o.p. 3, 4, and 5, third,

fourth and fifth acsephageal pores (rudimentary ); o.p.sk., supporting skeleton of cesophageal

pores; ov., ovary; pr., proboscis; pr.c., praboscis cotlom; pr, ca., proboseis canal (endsav); p.sk..

proboscis skeleton: sid., distal portion of buceal diverticulum; st.p., proximal portion of buccal

diverticulum; tr.c., trunk coelom: y.bl., ventral blood vessel; v.lam., ventral longitudinal mus-

cles: v.n.e., ventral nérve cord.

Ts LM. THOMAS

zw thick (Fig, 3). There fs also i general thickening nt the nerve layer at the bise

of the proboscis. particnlarly under the preoral ciliary organi where it may be

three times its normal thickness (Fig. 9) The outer layer of circular iuscles

immediutely under the basement membrane is a half to two-thirds of the thickuess

of the nerve layer. The longitudinal muscles form nine or ten concentric layers

separated by loose connective tissue. These are compressed and barely distin

cuishable immediatcly under the dorsal grouve (Fig, 4) and they are reduced to

five or six in number posteriorly in the vicinity of the proboscis complex (Fig. 9),

An inner, thin Jayer of circular muscles limes the coelomic cavity, The latter

extends almost to the tip of the proboscis. Anteriorly it is narrow, being only

about one tenth of the diameter of the prohoseis, but it expands considerably pos-

terlorly to accommodate the proboscis complex.

Two blood vessels are present immediately outside fhe outer circular musele

layer on the erests of the ridges formed as the result of the presence of the dorsal

wroboseis groove (Fig. 4). A small, subneural vessel (median dorsal vessel) has

heen seen in some specimens in the mid-dorsal line but a median ventral blood

vessel lias not heen observed in specimens so far examined.

The buecal diverticulum (stomochord) (Fig. 9) has a wide, ventral pocket

the posterior wall of which is indented by the blunt, anterior tip of the proboscis

skeleton. The diverticulum is bent slightly backwards at its tip under the end of

the skeleton. The lumen of the pocket is wide and broadly in contact with the

main lumen of the buccal diverticulum within the neck of the proboscis but it

is not confluent with the huceal cavity. It is occluded at the level! of the peint

of migin of the horns of the proboscis skeleton. Anterior to the ventral packet,

the buccal diverticulum has no continuous lumen, but only a series of uncon-

nected cavities which: diminish in size anterforly, This anterior part of the

buecal diverticulum differs from the form usual in the genus in that it is sharply

dividect into two regions of more or less equal length, The proximal part is thiek

and conical, while the distal part is very thin being only about one sixth of the

diameter of the proximal part at its widest, The distal part is attached antero-

ventrally and curves dorsally to end near the anterior extremity of the glomerulus.

Its cells are much smaller and less vacuolate than those of the proximal part but

thy separate portions of its lumen are apparent almost to its anterior end.

The glomerulus (Fig. 9) surrounds the distal part of the bueeal diverticulum

bul al the level of the proximal part it is almost entirely lateral and ventral. [+

ceases posteriorly at the level of the ventral pocket of the huceal diverticulum.

The curdiac vesicle (pericardium) and ventral sinus (Fig. 9) call for no speciul

voninent, The dorsal mesentery of the probuseis extends forwards nearly to the

level of the constriction of the buccal diverticulim, The ventral musentery is shor-

ter, extending forwards only. to the level uf the ventral pocket of the buccal diver-

Hivulum, The Teft coelomic pouch so formed, communicates as is usual in the

genus, throngh the proboscis canal (endsac) with the prolioseis pore which is dor-

solateral on the left side of the proboscis stalk,

The proboscis skeleton (Figs. 5 and 9) is slender, terminating anteriorly in a

rounded tip which partly penetrates the posterior wall of the ventral pocket of the

huceal diverticulum. The ventral keel is well formed and posteriorly it bifureates

to become continuous with the homs. These extend aliout halfway along the

collar and embrace about half the circumference of the buccal cavity.

Collar

The five transverse, epidermal zones of the collar distinguished by Spengel

(1893) are present (Fig. 6). The first, which overlaps the anterior Range of (he

collar, contains cells which have large vacunles distally. This zone meres into the

TWO SPECIES OF SACCOGLOSSUS 7

second. This forms an epithelium, which, at its thickest, is more than three times

as thick as the first zone, and occupies nearly three-fifths of the length of the

colliv. Its cells contain large numbers of small basophil granules in the imyer

three-quarters Of (heir lengths. The third zone and the fourth and fifth zones com-

bined, form two elevated bands which are readily visible externally. The third and

fourth zones are little more than half as thick as the second, The third is histalogi-

cally similar to the second except in that the basophil granules are concentrated in

the inner half to two-thirds of its cells. The fourth zone is by far the narrmayest

and forms part of the posterior wall of the groove. Its granules are more densely

packed and are fairly evenly distribnted throughout the length of its cells. In the

fifth zone the granules are more sparsely and evenly distributed and towards its

posterior end there are peripheral vacuales which are characteristic of the epithe-

lium of the brinchial region with which this zone merges.

The nerve cord of the collar is solid throughout its length, The dorsal anvseu-

lery is Incomplete and ends anteriorly a little behind the point where the pro-

koscis skeletan divides to form the two horns, The ventral mesentery is mare

variable in extent, In some specimens it extends as far forwards as the level of

the posterior tips of the hors. ie. about halfway wong the collar while in others

it js apparent only near the posterior end.

Trienk

The gill pores imerease in size over the first two or three and decrease in size

uver the last four or six. The posterior gill pores are very small and may lack gill

youches. Behind these there may be two or more rudimentary gills. The tongucs

project farther into the pharyny than do the septa. The gills extend a little more

than half-way around the circumference of the pharynx.

In the oesophagus there are three regions. In the first, the epithelium is

similar to that of the ventral part of the pharynx but its walls are thicker and have

a higher proportion of gland cells, Its lateral walls are irregularly sacculated (Fig,

10). In the second region the walls are much thicker and the lumen correspond-

ingly narrower, At its untero-dorsal end there are deep grooves in the dorsolateral

walls into which the oesophageal pores open. Of these, two lo five are patent and

two to four do not open to the surface but are represented hy outpushings of the

dorsolateral grooves. There may be cormesponding indentations of the epidermis

wbove them, All ure supported by skeletal eloments which usually fuse to form

an almost continuous plate which is perforated in the positions of the patent and

the nou-patent pores. These clements are, like the other skeletal structures of the

animal, thickenings of the basement laver which underlies the nerve layer over the

whole of the body. In the third region of the oesophagus the walls are thinner and

the lumen wider, Laterally there is an irregular sequence of shallow pouches. This

region merges into the hepatic region which differs from it maimly in that the

lateral pouches are Jarger and more regularly arranged. In the intestinal region,

the alimentary canal is simple. Its wall is thin and the lumen wide. There are

hawever, two ventral thickenings separited by a narrow, median, longitudinal

groove which extend nearly to the posterior end.

COMPARISON WITH OTHER SPECIES

Three other species af Saccoglossus have been described as possessing dee

dorsal grooves an the proboscis. These are 8. mereschkowskil (Nic, Wagner) 188) 5

S. ofayvoensis (Benham) 1809, and S, sudcates (Spengel) 1893. While the first two

have been adequately described (van der Horst. 1939), the third was described

from three anterior ends only which became dried up so that detailed examination

was impossible (Spengel, 1893),

ait 1. M. THOMAS

The main differences between these three species and S. aulakoels are listed

in Table 1. The relevant data on S. mereschkowskii, 5. otagoensis and S, sulcatus,

have heen taken, in the main, from van der Horst, 1930 and 1939.

§, aulakoeis is established on its possession of the following combination of

characters.

1. A deep dorsal proboscis groove.

4. The ventral musculature forms a promiticut “sole” in the posterior bran-

chial and ocsophageal regions,

3. The collar is slightly longer than broad.

4, The fourth epidermal zone of the collar epidermis is very narrow.

5. There are two to five pairs of patent oesophageal pores preceded by twa

to four rudimentary ones.

6. The longitudinal muscles of the proboscis ure arranged in nine or ten

concentric layers.

Epidermal gland cells extend to the bottom of the proboscis groove.

8. The glomerulus covers the anterior end of the buccal diverticuhim.

4, The buccal diverticulum has a very narrow distal portion which is not

conspicuously bent and in which the lumen is incomplete.

10. The proboscis skeleton is bluntly rounded anteriorly,

1l. There are no cavities in the dorsal nerve cord of the collar,

DISCUSSION

8. aulakoeis has been found only in the restricted regions indicated in the

introduction. Even here it is not common. It is rarc to find as many as two or

three specimens in a spadeful of soil, It is interesting however, that frequently

specimens varying in size from 1-5 cm to 5:5 cm may he found at the same time.

This suggests that the breeding season for the species is an extended one or that

there is a considerable variation in growth rate, However, animals with mature or

maturing gonads are generally seen in late winter and early spring (Fig, 7), while

in the summer months the gonads are invariably small (Fig. 8). Mature specimens

do not coil markedly in the post-lranchial and oesophageal regions as do those of

S. apantesis (Thomas, 1956), This is, no doubt, to be associated with the lessee

degree of development of the genital ridges,

In all specimens examined, with the exception of those fixed without

adequute narcosis, the collar is slightly longer than it is broad, The definition of

the genus states that the collar is “about as Jong as it is broad”. The proportion ot

the length to breadth of 1-0 to 0:77 is considered to fall within the limits of the

definition but the extent of the ratio is noteworthy.

Non-patent oesophageal pores have heen described for $, ruber (syn. 8. cam-

brensis Brambell and Cole, 1939) and they also occur in the present species, If

will be necessary to determine their presence or absence in several other forms

before reasons for their existence can be discussed,

‘There is a possible association between the dorsal thickening of the uerve

layer and the dorsal groove of the proboscis. Both S. ofagoensis and S. aulakoeis

have deep dorsal grooves and a thickening of their nerve layers. Similar thicken-

ings of the nerve lavers have been described in S. ruber and in $. horsti (Brambell

and Goodhart, 1941) and in these too there is a dorsal groove in the proboscis

TWO SPECIES OF SACCOGLOSSUS $1

though it is net nearly as deep as it is in the first two species named. In S. apani-

tesis however, the dorsa! groove is only slightly developed and the thickening of

the nerve layer is also slight.

While the description of this species was being prepared, some specimens

which clearly belonged to the same genus were found under stones on Snapper

Point, This hes about half a mile (about 800 m) south of the pool in which S.

aulakoeis had been found. These specimens were at first thought to be the same

species as they also had u deep dorsal groove on the proboscis, Closer examination

and Sater, a study of longitudinal and transverse sections showed that this was not

the case but that they were Saccoglossus ctagoensis (Benham).

Balanoglossus ofagoensis, Benham, 1899

Dolichoglossus otagoensis, (Benham) v:n der Horst, 1930

Succoglossus otagoensis, (Benham) Van der Lorst, 1939 (p. 399)

This is the first record of the species outside New Zealand, Three fairly com-

plete specimens and two fragments were found under stones in about one foot of

water at low tide on the northern (more sheltered) side of Snapper Point Reef

(Jat. 35° 17° S, long. 138° 26’ 30” E), This is an extensive wave-cut platform

of almost horizontally bedded sandy limestone of Pliocene age. It ix relatively

hard in parts but on its northern border and in some other regions, it is softer

and somewhat friable. Thus, along its northern margin, a secondary, narrow:

reef flat has been formed about two feet below the general level of the main rect.

This is covered by about au foot of water st normal spring lows. One Specimen

was found under a stone in a permanent rock peal about six inches deep on the

main reef surface.

It is a crawling rather than a burrowing species, as is 5S. aulakocis and it

agrees closely with the description of 8. etagoensis as given by Benham ( 1899).

and van der Horst (1930 and 1939). The points of undeniable resemblance are

asterisked in Table 1. In regard to the remaining points listed, the collar is rather

shorter than it is broad in the proportion of abont 1-0 to 0-70. This may be due to

contraction in fixation. The specimens from Snapper Point are relatively immature

so that the irregularity in the lateral genital bulges is not very apparent. In regard

to the presence of gland cells in the proboscis groove, van der Horst (1930) states

that they are absent from the base of the groove and his Fig. 2 (p. 137) shows

them to be present in the lateral walls, This is the case in the Snapper Point speci-

mens too. However in his diagnosis of the species (1939) he says “Keine

Driisenzellen in der Epidermis in der dorsalen Furche’, which implies that they

are absent from the groove altogether. This seems to be an oversight as they are

present in the walls of the groove (though not at its base) in specimens in the

author's possession which were collected at Portobello. New Zealand. which is

close to the type locality of the species.

Van der Horst (1930 p. 139 and 1939 p, —61 ! describes the buceal diverticu-

lum as haying two marked flexures in front of the ventral diverticulum. This is

figured in his 1930 description (Fig. 4, p. 139), The Snapper Point specimens do

not have these marked flexures. It is telt that these may well be fixation artifacts

in van der Horst’s specimens,

In New Zealand, the species is found in amongst coralline algxe at Wellington

and amongst the holdfasts of Macrocystis at Portobello. This shows it to be a

crawling rather than a burrowing species and indeed, Benham in his original

account of the specics writes of it as crawling on ua stem of seaweed,

1, M. THOMAS

pepunor Ayunyq aoz

-ajays sfasoqoid jo dr} Joraywy ¢

“Kraarp yeoonq Jo dy puosaq |

Ajroliayue = spuayxa snyniauta[g |

sonxay payrwist

ynoyyM nq ApIOUe}UR LOL

-Inu AOA WNpUIAearp Peony

yayood [ey

-UaA OU} JoLia}ue snonuyuos you

WH[NOWaAD PRONG JO WeluNT

AYAYD YINOUL OJUT Nodo JOU seop

woynoaniaarp ywoong jo AyAvy

SBULI OLIUaOUOS {)T

10 § WL sapsnut [eUTpPNySuoT

9AQOIS slosoqoid [esiop jo gseq

ur jquasaid spars ypeuuepidy |

gjeutay pue opeul yyoq |

ut qvpnBar Apey sprog TeMuaD

peor

wey] dasuop Apysys sBoD |

proaq sv suo] sv ynoge AvTJOD |

juiod dreys “auoj yya uo}

-ajays stosoquad yo dy stopeyay,

wnNITyIIAIp

Jeaong Jo joy Ul payun jou

SnMsawo9p3s ayy JO saaTey OML,

APIOMA}UR MOLTB fsarnxay

jowayue way} peIyUaA poyrelw

quan winqMaLAaAtp yeoong

snonuiyu09

Un MonAeATp [Been jo SULIT

AYACI TQNOU OUT uedo Avot

lunMoHaArp [e90ngG JO AWARD,

SSULL OLNQUSAIOD P

JO © UL saposnul [BUIpUpPSWOT,

aAooIs stosogoid [esuop

yo aseq Ul spur[s [euLspide ON

ayewoay Ul saspnq ey]

-NSaIM FO WIOF UL SPlOF TeWUOS)

proiq sv Buoy se qnoqr seyjOD

qurod divys “suo, yyy U0}

-g[ays stasoqoid jo diy Joey

umypnoyiaarp Teoonq

jo [Je Apuau sano snpniewlo[s

qysreys

APPAQU[aT WinynoAwaAip Rong

snonurywo09

WINnpWarHeATp yeaonq fo wsuM’y

slosoqgold

SBULL S1}UB009

OL OF L Ur epost Teurpnysu0T

A[ao apeuiay uy Surmeadde 34y

-ngal ssay Jo 3100r Sp[OF pequsy

pvorq se Suaoy sv ynoqe AOD

saingoa y [DULay

: slaoynynn *s

Ssnjoopns °S

SISUBOTDIO 'S

MYSMOYYISAAIUL *§

“sla0yD]ND *g pu snynopns. *g “HYSMOYYIsasauL -g ‘s|sUaORDJO snssojzog9vg Jo samyvay aysouderp Jo wosriedurop

T HTaViL

TWO SPECIES OF SACCOGLOSSUS

‘sualujoads puvjeaz MeN pue UeyeYsny NOG UsaAjaq AjETLUTS yo sjaiod sayeorpuy ,

Bypeasny wos AK IOVFO

‘NY Ys FIND qsoura 4g

AJVAG QUO UL BAO Ad aroUr IO g

soxod AIVUaTUIT pI

jo sired % 0} g pue sarod wad

-vydosao yuayed jo sued ¢ 0}

suoyoeas ¢ YYWAX susdeydosag

Ss yo sued og 04 Z]

saspii [v4912] stonordsuos WUOF

Sapsuu = RUIPNySuey yeruaA

piod aarauU [sIOp UL SaVIAvD ON

ajo[dmoour yng yasaid

Solayuaselt [LIJUVA PUR TesI0g

sauoz [Butapida 3Aly

SSoUYIIY] [LUIOU Jo syutseprd|

AYIABI

tqnour Fey yNoqe aoviquia pur

Ipyjoa Buope Ava-Jey joqe pua)

“x9 UO}IfeXs stIasoqoid Fo su10}F{

uvdef |

SITES Jo sured [] 0} OT

FOOOULA "19 pug PuRlfeoZ Mon

AIBAO OUO UT

BAO adir 9 uel} azo TUOPT9S gy

sarod yeaseydosso jo amd aug,

suolpes g YIM snsrydosag

Sys JO sed cy 07 OTs

saspi

[Blaze] WIoF Jou Op Yyuns jo

Sappsnur [RUIPNysuoc], [eyooA,

p40. vAtou JPSIOp UT SaT}TAvr),

yuasqv

SaLlo}Uasaul [BQUaA PUL [BSIOC

sauoz Teuoapida dear], ,

ssauyaty] [eUuOU jo stuaprd'yz,

yoo et)

pus Jopuny yeal pure yeyuozoy

udzJoOAS stosoqoid jo SUIOY ,.

JO Rag OY} O} LIssNY UIsIION

Uu01}D90°T

saiod

[vaseydosao jo sued y ynoqy

suotpoas § Ym suseydosag

S]T4 jo sed gg Jnoqy

yun,

pi02 OAraU [esIOp UT saryTatry

aqyayduioaur ynq yuosaid

SASPUSsAU [RIYUDA puUe Tesi0dy

(war co‘9)

IGT AIA stuwopidy

Ingjog

Slaoyojnp *¢

snyp2 ns “S

HSUZOBDIJO “S

inysmoyyosasau ‘5

_-e_,———— Eee

84 J, M, THOMAS

REFERENCES

Bennam, W. B. (1899). Balanoglossus otugoensis n.sp. Quart. J. Micr. Sci. 42; 497-504.

BramMBE.L., F. W. R. and Core, H. A, (1939a). The preoral ciliary organ of the Enteropneusta;

its occurrence, structure and possible phylogenetic significance. Proc. Zool. Soc. London.

B., 109, 181-193.

BRAMBELL, F, W. R. and Coreg, H. A. (1939b). Saccoglossus cumbrensis sp.n., an Enterapneust

ovcurring in Wales. Proc. Zool. Soc. London. B., 109, 211-236.

Braneert., F. W. R. and Gocpmart, C. B. (1941). Saccoglossus horsti sp.n., an Enteropneust

cecurring in the Solent. J. Mar. Biol. Ass. U.K. 25, 283-301.

Bunpex-Jones, C. and Patiz, A. M. (1960), A revision of the genus Saccoglossus (Entero-

pneusta) in British waters. Proc. Zool. Soc. Lond., 134, 635-645.

Horst, C. J. van der (1930). Observations on some Enterapneusta. (Papers from Dr. Th.

Mortensen’s Pacific Expedition 1914-16). Vidensk. Medd. naturh. Foren. Kobenhavn.

87, 135-200.

Hors, C. J. van der (1939). Hemichordata, Bronns Klassen u, Ordnungen des Tierreichs

Bd. 4, Abt. 4, Buch 2, Tiel 2.

LepincuaM, Isaper. C. and Wexxs, G. P. (1942). Narcotic for marine invertebrates. Nature,

London, 150, 121-22.

SpENcEL, J. W. (1893), Die Enteropneusten des Golfes von Neapel. Fauna u, Flora des Golfes

von Neapel. Monogr. 18.

Tuomas, I. M. (1956). Saccoglossus apantesis a new species of Enteropneust from South

Australia, Trans, Roy. Soc. S. Aust. 79, 167-176.

LUNETTES IN SOUTHERN SOUTH AUSTRALIA

BY ELIZABETH M. CAMPBELL*

Summary

Lunettes are low crescentic ridges, which commonly occur on the eastern side of ephemeral lakes.

They are of two types-those composed predominantly of sand, and those composed of sand, silt and

clay. The mineral composition of lunettes varies, though lunettes rich in quartz or in gypsum are

common.

The morphology, composition and distribution of lunettes in southern South Australia are examined

in the light of suggested theories of origin. The deflation hypothesis of Stephens and Crocker and

many other workers can account for most, though not all, of the field evidence. Lunettes develop as

a result of wind action, but wave transport is an important factor in concentrating debris on the Ice

sides of lakes.

The available evidence suggests that although some lJunettes are apparently still forming, they are

essentially relict features dating from the recent past.

LUNETTES IN SOUTHERN SOUTH AUSTRALIA

by ExizapetH M. Campseii®

[Read 12 September 1965]

SUMMARY

Tameltes: are low éréscentic ridges which ecmmonly occur on the castern

side of ephemeral Jakes. They are of two types—those composed predosuinantly

of sand, and those composed of sand, silt and clay, The mineral womposition

of Juncttes varies, though lunettus rich im quartz or in gypsum are common,

The morphology, composition and distribution of Juncttes in southern South

Australia are examined in the light of suggested theories of origin. The deflation

hypothesis of Stephens and Cravker and many other workers can account for

most. thongh not all, of the Held evidence. Lamettes develop as a result of wine

action, but wave trausport is an important factor in concentratng debris on the

lee sides of lakes.

The available evidence suggests that although some Iuncttes are apparently

still forming. they are essentially relict features datiny from the recent past.

INTRODUCTION

The purpose: of this paper is to describe the shape, cornposition and internal

structures of lunettes in southern South Australia, and to discuss their possible

age and origin. The term lunette as here used includes not only the crescentic

dunes located on the lee side of lake and swamp depressions and composed

variously of silt-clay, clay-loam and sand, but also the so-called gypsum dunes

of Jack (1921) and others. |

Features of similar morphology and situation have been described from

several parts of the world, including North Africa (Tricart, 1954a; Boulaine,

1954) and North America (Coffey, 1909, Huffman and Price, 1949), They have

also been recorded from several parts of the Australian continent. In general,

they are best and most commonly developed in the semi-arid regions of Victoria,

South Australia and Westem Anstralia, thongh they occur also in humid zones,

for example Tasmania, and in truly arid regions as, for example, the north of

Sonth Australia,

DESCRIPTION

Morphology and Size

The dune on the eastern side of Lake Bumbunga, Mid North, 6-4 km long,

21 m high, and 1:2 km wide, is an example of the larger lunettes known in South

Australia, At the other end of the scale are the small rises about 0+S km long,

4-5-6-0 m high and 45 m wide, typical of the flat, swampy interdune areas of the

Southeast. There is no consistent and direct relationship between the size of each

Iunette and the extent of its associated lake, Lake Greenly, Eyre Peninsula,

which is about 23 km? in area, is bordered by a Junette which is only just dis-

tinguishable. Lake Baird, Eyre Peninsula, on the ather band, which covers only

1-3 km? is bordered by a lunette 2 km long, 21 m high and 0-8 kin wide (Fig. 2),

Nor is there any consistent distributional pattern; in a given district, some lakes

have associated limettes, athers do not.

~ Geography Department, University of Adelaide.

1 For the locution of all South Australian place naiues mentioned, sec Fig. 1,

Trans. Roy. Soc. S.A. (1968), Vol. 92,

86 ELIZABETH M. CAMPBELL

Yongala *Hutlant Lageon

Bordertown

Moone Flab Ss

* Sere

wy odulla

eXyanculta wHites Lagoon

eSnowlawn

*Loke Gumbunga

én mark

Barri

*ADELAIDE

#Cookes Plains

Lake

mots oe eCumm'nt

Greenly® li

\ ®Port bineoln

ing

Yorketawn

Siske fowler

hearst

White aqean

Enlargemest A “Sarndurce 4

Kongoreo Island PReth

Burden iow

{

Enlargamen! 6B Frances

; i ”

Narorsa:tel loke Cadn

. Cockatao Lake

Lo] 4

loo mj as f “soll tale {

° Penola I

° igo kilometres Mount "Lochaber

5 um bey ;

= Swamp

Unnamed ve Hynom Swarnp

. ae ae

et Hy 6 am

pant

Nayacoerle

Unnamed « Smiles

lose d

—

week Rest ys

— |

a lakilomet es —— Roads

———!

='™Stale

& Skilemerres \ Bel lagoon Bowndory

i Baebes

Fig, 1, Southern South Australia—Location, Enlargement A. Enlargement B.

Lunettes are crescent-shaped in plan and extend between one-quarter and

one-half the way round the perimeter of the lake (Figs, 3, 4, Pl. 1).* They are

most commonly found either on the eastern or southeastern side.

The smooth form of the lunette on its western or lake side contrasts with the

irregular or sinuous shape of the eastern slope of many lunettes. The occurrence

® For reasons not yet clear, an wonamed lake on the Jaterite plateau of Kangaroo Island is

completely surrounded by a dune, whilst several depressions south-east of Renmark are

bordered on their western side by gypsum dunes. In the south-west of Western Australia

between Kindinin and Corrigin where summer easterly winds are significant linettes occur

on the western side of depressions (C. R. I'widale—personal communication).

See opposite page Fig. 2. Contour plan of Lake Baird lunetic. Plane table survey.

Five feet = 3-3 metres, 500 yards = 500 metres,

LAKE

BAIRD

LUNETTE

CONTOUR INTERVAL

to FEET

ARBITRARY CATUM

seater tin yarns}

igh

Height in ¢

o *

o sa

Vertical Exaggeration 3-75

8&8 ELIZABETH M, CAMPBELL

LOCHABER

SWAMP

LUNETTE

CONTOUR PLAN

Contour interval

S feet

Arbitrary datum

Lochaber

Swamp

Scale in feet

ro) Q 500 19000 isoo

7 ‘ct = st

Artificial

Drain

PROFILE A-B

Vertical exaggeration 7:1]

8 a0 4 40

= 3

= 20 ae 20

> 0 A

x 500 1000 «=1SO00 feet

17-18°5-65,E.C+ DC.

Fig. 3. Contour plan of Lochaber Swamp lunette. Plane table survey.

Five fect = 1-7 metres, 1500 feet = 500 metres.

LUNETTES IN SOUTHERN SOUTH AUSTRALIA

LEGEND

Arbitrary datum

Contour interval 10 feet

——— Road

Quarry

2400 feet

Fig. 4. Contour plan of Cooke Plains lunette. Redrawn from King 1949a.

Ten feet = 3.3 metres, 2400 feet = 800 metres.

90 ELIZABETH M. CAMPBELL

of consolidated rock outerops at the Jake edge, as for example at Lake Greenly,

introduces complications and further irregularities in the plan,

Many lakes are bordered not by a single Junette, but by two or more luncttes

arranged concentrically, or nearly so, around the eastern shore. At White Lagoon,

Kangaroo Island (PI. 2), for example, there are no fewer than three distinct and

separate dune ridges bordering the lake on its eastern side. The pair of lunettes

which borders Lake Fowler, Yorke Peninsula, and those close to the eastern shore

of Hiles Lagoon, Mid North, are of similar composition, but at White Lagoon the

outer or easterly dune is sandy, the westerly ridge immediately bordering the

lake is composed predominantly of silt and clay, and the one between these is

—____ fe

er eer ee

—I,

—

500 7000

See an vee 2 ——— ee ae

2nd 490

HORIZONTAL DISTANCE

Fig. 5. Transverse profiles across selected lunettes from west to east (left to right). A—Moona

Flat lunette, B—Lake Cadnite lunette, C—Lochaber Swamp Imette, D—Cockatoo

Lake lunette, E—Hutton’s Lagoon Junette, F—Hiles Lagoon lunette, G--Bool Lagoon

lunette.

LUNETTES IN SOUTHERN SOUTET AUSTRALIA a

composed of sand, silt and clay. More complex patterns of lake and multiple

lunettes are displayed at Bool Layoon and Salt Lake near Naracoorte, Southeast

(Pl. 3). Here complexes of smal! lakes each with their associated lunette or

lunettes, occur within larger lake basins on the caste margins of which are

found individual luncttes.

Most Junettes in southern South Australia are asymmettical in transverse

section, the steeper slope being the western or lake slope (Fig. 5). The gradient

of the western slate of Bool Lagoon linette (Pl. 4) averuges 1 in 3+3, whilst the

eastern slope is 1 in 18-3, Some few, however, are roughly symmetrical and

several examples have been noted in which the castern slope is steeper than the

western. This is so along part of the Lake Baird hmette (Fig. 2), part of the

middle lunette on White Lagoon and along most of the inner lunette at the same

site. The western slopes are tor the most part smooth and rectilinear and, though

varying from one hinctte to another, are essentially uniform on any given feature,

The inclination of the eastern slope, however, varies considerabl , even on the

same Iunette.

In longitudinal section, the crests of lunettes typically rise gradnally to a

high point near the centre of the feature (Fig, 3) but many crests are undulating

(Pigs. 2, 4). On the innermost of the White Lagoon linettes, there are no [ewer

than four distinct crest lines, all of them undulating, the high points of one crest

invariably located opposite depressions in the adjacent ridge (Fig. 6),

Fig G Diagram showing the multiple crestline of the linette mnmediately adjacent tw White

Lagoon. viewer! from the est.

Composition

Samples of lunctte material were collected on the crest where it reached its

maximum development, and from the lake hed adjacent to this position, Borings

were made to below the level of soil development.

On the basis of particle size, lunettes are of two distinct types—those com-

posed of almost pure sand and those composed of sand, silt and clay (Fig. 7),

The predominantly gypscous and predominantly quartzose sandy Junettes are

clearly differentiated. The sorting value developed by Trask (Krambein and Sloss,

1963 p. 101) is a measure of the uniformity of the sample and is derived from

the formula Sy = Q:/Qs where §, is the sorting value and Q; and Qs, represent

the grain size values corresponding to the 25% and 75% marks respectively. of the

sample. The sorting value of the sandy Iunettes is 15 or less, that is. they are well

sorted, whilst the fine-grained lnnettes are poorly sorted, with a value ranging

from 5-6 to 7:6, Field determinations of samples collected at various depths and

from various positions across the linette indicate that the mechanical composition

is generally uniform, although there are Iocal and minor irregularities,

92 ELIZABETH M. CAMPBELL

100

Per { ——$ _—__—__—— a = 4

Fy .

e Lunettes. |

e 1, Cocke Plains Gypteoyve) |

a 1

r. Zlake Fawler Sandy Quariose \

A 3, Loke Cadnite Lunerics Sandy .

. 4, abe Board } Lumens 7

£ 5, White boagoon

2 b Lochaber Swenp

£50 =

> FHynem Swamp |

a 8 Cockulos Luhe

= \

“

= Fineegriined

Lunettes

25 _ -|— eC —_ |

cas

Al 4

3 =

1G 4 i} 4s 4 05 2) 005, “007

Pesticle Size (millimeters }

Fig. 7. Cumulative percentage curves of the size distribution of Imette samples,

The mineralogy of samples from six Junettes and their associated Jake beds

was analysed by Australian Mineral Development Laboratories (A.M.D.L.),

Adelaide. The < 0-002 mm fraction (Table 1) of the lunette samples is com-

posed predominantly of “amorphous” material, which was not positively identified

but which shows physical properties similar to illite and smectite, or “amorphous”

material and illite. Quartz and kaolin ave present in all cases, and illite-chlorite,

illite, calcite and chlorite in some.

The > 0-002 mm fraction was separated into a light and a heavy [raction

(S.C. 2-96), The light minerals (Table 2) include quartz which is abundant in

all and dominant in most, silt- and clay-sized particles too small to be identified

by the methods used, and, in some samples, minor amounts of calcite, layer

silicates, organic remains and opal.

The heavy minerals (‘Table 3) vary from 0-007%-0-31% of the total sample,

They are numerous and vary from onc lunette to the other. Hematite, magnetite,

other opaqnes, tourmaline and zircon are the most common.-

Gypseous lumettes were not included in the selection analysed by A.M.D.L.

Apart from varying amounts of gypsum which is mainly the “seed” variety, calcite,

halite and insoluble materials, including quartz, opaques and many other minerals

are present in some samples (Table 4),

There is generally a close relationship between the mechanical and mineralo-

gical composition of samples taken from the hinette and from the associated

lake bed, though minor variations occur, The mineral composition (‘Tuble 4) is

LUNETTES IN SOUTHERN SOUTH AUSTRALIA 93

TABLE 1

Semiquantitative analysis of clay minerals in samples from Iunettes and associated

lake beds.

Minerals Present.

Weight | Amor- | Illite-

total | Material

oe joes. open or bs: _ a

Moons Flat

lunette < 10

bed <h0

Lake Cadnite |

lunette <10 |

bed 17 bb A A T |

bed (surface) <10

Lochaber Swamp

lunette 32 D A A T

bed lL D A T I

Cockatoo Lake

lunctte 3] Db A T A

bed 63 D F T A A

Hutton’s Lagoon :

lunctte 8 8 A A 8 F T

hed 22 8 A A r 8 F T

Boo] Lagoon |

lunette 28 D A F T T

bed 76 Db A A T T

D — Dominant: >50%

§ == Subdominant 20-50%,

A = Accessory 10-20%

T = Trace <10%,

F = Faint trace just detectable.

TABLE 2

Analysis of the light fraction mineralogy (8.G. <2+96 and diameter ~-(-002 mm) of samples from

lunettes and associated lake beds.

| Mincrals present a3 percentage of light fraction

Sample Weight Silt Layer Organic

oof | Quartz + Opal sili- Calcite | Remains

total clay eatcs.

Moona Flat linctte > 90 100

bed >90 of 5

Lake Cadnite lunette >90 100

bed &3 75 20 5

bed (surface) > 90 100 <2

Lochaber Swamp lhinette 68 BO 65 5

bed so | 85 15

Coekatoo Lake lunette 63 | 75 as) 20 <5

bed 37 30 | 50 20

Hutton’s Lagoon lunetto 92 24 70 5

bed 78 40 5h | 5

Bool Lagoon lunctte 72 20 i 50) 25 5

bed 24 | 15 i 5 60 10 10

| J}

ILIZABETIT M. CAMPBELL

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(uoowdy Aavey Jo % OT >) Tueserg = q

(uoorry AABOY Jo % 07-07) UouuIE)) = 9

(uoyariy AARoy Jo %OF<) yaRpunqy =

| 20-0

LOO +0

s0-0

60-0

S00

01-0

c+)

£00

Té-0

el:

90°+0

9T-0

OT+0

l | 1 5

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; d ral i => i a - od d | V dj}-|da

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=| |

32,

“spaq eYE] poyeLoosse pu soz, 9UN] WoIy saydures ur {"wIW Z0-0 < dayeWeIp put 9g:g < OS) syesaUTO AAvaT, Jo sis(jeue Arey yUrnhruag

£ ATAVL

LUNETTES IN SOUTHERN SOUTH AUSTRALIA 85

TABLE 4

Mineral comnposition of two pypscous lunettes and their associated lake beds.

Minerals present

“ree Gypsum pp Sodinm | Ferric | Insolu- Total

E bonate Chloride! Oxide bles

Cooke Plains lunctte (average of 16

samples) (a) 83°88 2°80 na. na. 11:21 ' 97°89

Cooke Plains bed (trial hole ,

$12:2'107-3'2”) (a) 47-77 | 21-70] ma | na. 18-43 - 87-90

Lake Fowler lunette (average of 33

samples) (b) 92-9 | Mel Q+7 (hel 3:9 98-7

Lake Fowler bed (sample

C4:0-294”) («) 75-70 1-11 1:95 0-34 18-3 97-4

Source:

(a) King (19494 p. 145)

(b) King (1949b p. 66)

(¢) Hiern (1957 p. 40)

n.@, not available.

TABLE 5

Percentages by weight of sand, silt and clay.

(The silt and clay fraction was not separated. in some samples: sill and clay—less than

0-066 mm. sieve size.)

Sample i Sand Silt & Silt Clay

Clay

Cooke Plains lunctte 98:4 16

Lake Fowler lunette 98-7 1:3

Lake Baird lunette ! 99-4 0:6

White Lagoon lunette 93-2 6-8

Moona, Flat lunette : 96-1 3:9

bed 96-4 3:6

Unnamed lake 2 lunette 97-0 3:0

bed , 97-2 2-8

Unnamed lake 1 lunette 94-2 5:8

bed 92°5 7-5

Lake Cadnite lunette 95+2 4-8

bed T4e4 25-6

bed-surface 2” 98°38 1-7

Hynam Swamp lunette | 37-5 B7-d 25:0

Lochaber Swamp lunette 45-0 32-1 21-9

bed 85-0 4-0 1150

Cockatoo Lake lunette 33-0 37-0 40-0

bed 79 92-1

Hutton’s Lagoon lunctte 34-2 65-8

bed 37-7 §2-3

Tliles Lagoon lunette 29-7 70-3

bed 41-1 58-9

TABLE 6

percentage of the total sample.

s from lunettes and lake beds

each fraction expressed us 1

Percentage size distribution (by weight) of sample

size in millimetres:

range of particle

Yas

Pipette Anal

ELIZABETH M, CAMPBELL

‘

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gs AM SOPRANO HOA I GAARAS w

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LUNETTES IN SOUTHERN SOUTH AUSTRALIA 97

TABLE 7

Quartile measures (in mms) and sorting af samples from hinettes ancl associated lake beds,

Sample Q, | Mg Q: | Sa

Cooke Plains hinette 0-85 0-61 0:35 | Ls

Lake Fowler lanette 0-68 0-61 0-37 143

Lake Baird Inneatte O26 ! 0-2 0-16 1:3

White Lagoon Iunette 2] | 0-16 O12 1-3

Monona Flat lunette 0-18 : O15 : 0-12 1-2

bed 0-18 15 OIL 1-3

Sandy Lake. hinette 0-23 | 0-19 (14 1-3

beet 0-25 0:20 0-16 1-4

Northwest. ake lanetic O32 423 O- 16 1-4

bed H Oap 0:20 - 14 I-5

Linke Cadnite hinctte | 0-38 0-26 0-18 bed

bed 0-26 | 0-16 0-06 2-4

bed—surtace 2" +25 0-18 O14 1-3

Wynn Swamp limette 0-086 0+ 0084 00018 7-6

Lochaber Swamp linette 0+096 O15 0- 0026 t-1

bed Q-17 0-098 0-44 1-0

Cockatoo Lake linette Q-Q31 | ()- 0966 (he (01 56

bed | na. H T.w, ra, Lith.

Huttou’s Lagoon lunette 0: 107 | TB, na. ied.

bel (125. i iA. Lh. ni.

Hilegy Lagoon lunetty ()- LL nA na, 11.8.

bored 0-105 | Tha. nea. | 1,8.

Sy O1/ Qs

mem > notoayailable,

the only information available by which to compare the gypseous sandy lunettes

with their associated Jake beds. The same minerals are present in both cases and

the percentages vary only slightly. The percentages by weight of sand, silt and

clay (Table 5), the size distribution (Table 6, Fig. §), the quartile measures and

degrees of sorting (Table 7), and the mineralogy (Tables 2, 3) all indicate that

the composition of the quartzose sandy lunettes is very similar to that of the

associated lake bed. The size distribution of samples from the fine-grained

luncttes varies slightly from that of the lake bed samples, especially in the small

size grades (Table 6, Fig. 8), but the mineralogy of the two is similar (Tables

1, 2, 3). The sample from Lochabcr Swamp lunette, Southeast, shows a lower

percentage of light minerals than might be expected from the percentages

observed in the same lake bed (Table 2). Uowever, the composition of the

Jake bed may have been altered by the man-made drain entering the swamp

(Fig.3). Other dissimilarities, for example, at Cockatoo Lake, can be accounted

for by the difficulties associated with the analysis of the samples.

Structures

Internal structures have not been observed in fine-grained Innettes, bul there

are several exposures in yypseous and quartzose sandy Junettes, The most exten-

sive of these are in Cooke Plains and Lake Cadnite lunettes, Southeast.

Cooke Plains linette is composed almost entirely of seed gypsum of sand-

grain size, In the quarry on the southern end of the lunette, near the swamp edge,

the dip of the beds is about 2° towards the Juke; near the centre of the lunette

the din is in the same direction but varies over short distances from 20° ta almost

flat (Pl. 5); and on the side of the Iunette away from the swamp, the dip is

consistently abeut 30° away from the depression,

100

ELIZABETH M. CAMPBELIL

LAKE

CADNITE

SIZES)

25 -

CERTAIN

{LARGER TRAN

30.

PERCENTAGE

CUMULATIVE

COCKaToOo

PARTICLE

100

PARTICLE

“01 ‘O05 -O01

DIAMETER (MILLIMETERS)

i i

| HUTTONS LAGOON

HILES LAGOON

(MILLIMETERS)

DIAMETER

Fig. 8 Cumulative per

centage curves of the size

distribution of lunette and

lake bed samples. The solid

line represents the Iunette

sample, and the dashed line

the sample from the lake

bed. In the case of Lake

Cadnite, the dotted line re-

presents the sample from

the surface of the lake bed.

LUNE TTES IN SOUTHERN SOUTH AUSTRALIA bY

The structures of Lake Cadnite lunette, composed jlmost entirely of quartz

sund, are only visible in isolated places where the quarry face has been exposed

to weathering. Typical measurements near the centre of the lunette are 2° to 12°

lo Uhe east (PL. 6), but whatever the value near the centre of the feature, the dips

in all cases Hatten to the east. A series of superimposed soil profiles is exposed in

a Quarry face to the cast of the crest for a distance of a few metres, Each «oil

consists of a black humus-rich layer which grades to light brown with depth and

overlies buft-coloured sand. This rests with sharp demareation on the black

honzon of the next lower soil,

Vegetation Cover

The surface of nearly all the Iunettes in southern South Australia is protected

by a dense cover of grasses, Trees are rare, hut red gums about 6 m high are

lound on Lake Cadnite lunette, and the Cooke Plains lunette is covered hy dense

Mallee scrub, The grass cover has apparently been sufficient to protect the

lunettes from crosion; the only occurrence where modifications to the original

Junette form are evident are those in which wave action and slumping have

resulted in a cliff on the lake cdge, or where overgrazing by shecp and rabbits,

or overcropping, have resulted i erosion of the surface.

ORIGIN

Precious Investivations

The two principal hypotheses advanced in explanation of Innettes are those

of Hills (1939, 1940) on the one hand, and Stephens and Crocker (1946) on the

other, Lt should be noted, however, that although Stephens and Crocker first gave

prominence to the deflation theory, similar ideas are either implicit or explicit in

the writings of Coffey (1909), Grayson and Mahoney (1910), Wyherzh (1918),

ane ITarris (1939),

Hills suggested that spray droplets whipped from the surface of the lake by

wind and dew formed preferentially on the dawnwind side of the lake cause the

deposition and retention of atmospheric dust on the lee shore of the lake, The

presence of the water in the lake is thus an essential feature of this mechanism.

Hills did not preclude the possibility of the addition of minor amounts of material

by deflation from the dry lake bed but considered this of minor siznificance.

Stephens and Crocker and many later workers {Huffman and Price, 1949; Gautier,

1953; Boulaine, 1954; Tricart, 1954a, b, 1955; Jennings, 1955) considered, huw-

ever, that this latter process was of prime importance. They considered that

wind scouring of the unconsolidated sediments exposed in the dry Jake bed

provides material which is trapped hy vegetation geawing on the lee shore of

the lakes. Once the lunette has developed, it propagates itself by deflecting the

wind over it whilst the material in transport is deposited on the lunette. The

essential contrast between these two hypotheses is that Hills’ requires water in

the lake while the deflation hypothesis required a desiccated lake hed.

Thus it is that these lwo hypotheses have been construed by different authors

as implying the association of the hinette cither with wel ( glacial) or dry (inter-

glacial) phases of the Quaternary. In view of the marked seasonality of climate

and lake conditions in southern South Australia at present, it seoms immecessary

lo invoke such definite and long term changes of climate. But the age or ape

range of lunettes is certainly significant, for, if the features developed under

2 Various other hypotheses have heen put fonvard io explain ximilag landfornis in other

133; Frouty, 1952), hut none seems ta have any relevance to the situation in southern South

Australin,

101) ELIZABETH M. GCAMPBRU),

conditions widely different from those which obtain at present, modern climatic

parameters have little or no relevance to considerations of their evolution. Thus,

before any assessment of the various lines of evidence, and a discussion of the

possible genesis of these Teulures, it is necessary to examine (heir likely age or

age range.

Age of Lunettes

Soil profile development on lunettes varies considerably but most have leer

sufficiently stable to allow the development of mature soils on the surface, Sisni-

larly, a layer of “flour” gypsum, the weathering product of the “seed” varicty,

occurs at the surface of the gypseous lunettes. This is an indication that weather:

ing has latterly outpaced accumulation, suggesting, that lumette formation has

ceased or has ut least slowed down. However, it is not necessarily evidence af

antiquity; as soils can develop in a relatively short time, as evidenced by horizon

development on sediments no more thar 100 years old at Beefaeres, an Adelaide

suburb (C. R. Twidale, personal communication ),

There are isolated indications that linetles are forming today; the bed of an

unnamed depression 13 km. northwest of Naracoorte is bare, but flat-topped

remnants standing 4 metre above the bed are vegetated. This is taken to mean

that fhe mesa slopes and the bottom of the unsalted depression have heen

recently removed, apparently by deflation; movement of material across the

lunette on ITutton’s Lagoon has heen observed: and there are reports of luneites

forming today in Western Australia (Bettenay, 1962), Victoria (Baldwin, Burvill

and Freedman, 1939) and Texas (Caftey, 1909; Huflman and Prive, 1949) under

conditions similar to those in southern South Australia.

In any case, the evidence points to the luncttes heing geologically youthful

rither than to their antiquity. From a variety of evidence throughout southern

Australia, Stephens and Crocker (1946) concluded {hat “it is apparent that the

halk of the Tnettes belong to the lute Pleistocene to Recent: Period”. King

(419492) concluded that the Cooke Plains gypsum dime postdated the emergence

of fend after the wid-Recent high sea-level when conditions permitted the evapo-

ration of water and the precipitation of salts, In the Riverine Plains of south-

eastern Australia, Pely (1866) noted that lanettes predate the most recent period

uf stream incision into the plain but postdate the lust phase of deposition by the

prior streams which occurred in the Pleistocene (Pols, 1964), Bowler and Ifarford

(1966) in the same area, worked out a geomorphic sequence in which they retog-

nized three phases of lunette formation: the first oceurred about 7,000 B,P.,

indicated by a date from lacustrine silts in the hed of the lake on which the hinette

formed: the second occurred following shrinking of the lake: and the thirel alter

a period of prior stream activity, deposits from which were dated by the carbon

14 method at 4.000 B.P. Bowler and Harford thus showed that linette formation

in this area began at least 7,000 years ago, and continued at least until 4,000

years ago, that is, through the middle Recent."

Whilst lunettes in southern Australia may vary in age, it is likely that their

age range is of the same general order, and it can be assumed that the lumettes in

southern South Australia developed during the Recent. They are, therefore, a

relatively youthfnl feature of the landscape. Sinew this time, the climate of

sonttiert: Australia is thought to heve changed only slightly se Uiat conditions

fayunrable to linettc formation are not very different Gout the range of those

‘Carbon 14 determination of material at the use anel at the snrlice ofa linete offer Te

next satislactory answer to the range af time clupine which formation Wok place, Datahl

fuiterial has so Fur heen found only vear the surface of lunetles in South Austrlia aud poage

donominitions lave heen carried out.

LUNETIES TN SOUTITRRN SOUTLIE AUSTRALIA td

expericnced at present. An examination of present climatic parameters is there-

fare relevant in a sludy of the origin of luncttes.

Discussion of Orisin

The fine-grained hinettes in southern Soufh Australia are composed of 76-90%

mulerial less thay O-L mim diameter (Fig. 7), ie. it can be transported in stes-

pension, All the hinettes deserfhed by Hills (1939, 1940) ia north-western Victoria

wre dominuntly fine-grained. Most of these lunettes could have been formed by

the deposition of atmospheric dust on the lee side of the lake, Deflation is not,

however, excluded, Most of the particles in the sandy lunettes and sore in the

fine-grained luncttes are too large to be transported jn suspension, and thervfore

cannot be explained by the mechanism suggested by ills, Other lines of evidenve

fram scuthern South Australia throw further doubt on this theory.

No measurements for the precipitation of dust over rural areas. are available,

but the amount precipitated in wetter months, as required by the atmospheric

dust hypothesis, must he extremely small (R. Culver—personal communication ).

ly view of the youthfulmess of lunettes, it is considered that there has heen in-

sufficient time tor the large amounts of material required to make up a lunette to

have heen deposited from material suspended in the atmosphere.

Variation in composition of linettes In close proximity and in an area of

uniform soil types is incompatible with the atmospheric dust hwpothesis. Cackutna

Lake and Lochaber Swamp, no more than 10 km apart, are associated with

lunettes of quite different composition (Tables 1, 2, 3), and the composition ot

the three hunettes associated with White Lagoon, as stated previously, varies

considerably,

The position of the hinettes on the castern side of lake depressions indlcates

formetiun by westerly winds, and, in general, winds in southern South Australia

hlow froin this seetor (Fig. 9), In most areas where luncttes are numerous, these

winds are the least dust-bearing as they blow for only short distances across land

(Stephens and Crocker, 1946),

Twe morphological characteristics of lmettes which arc atypical of dunes,

their smooth and regular contours as opposed to # hummocky surface, and their

asymmetry with the steeper slapes on the windward rather than the leo side, were

interpreted by Hills (1940) as evidence against the deflation hypothesis of Innette

formation. There are, however, several factors in the lunette environment different

from that of dunes in general, which can account for these anomalies, The surface

of Junettes is covered and protected hy vegetation, thus preventing major re-

shaping of the material, and the development of a hummocky surface. A slow and

steady (although not necessarily continuous) rate of accumulation which js in-

tlicated by the presence of low angle dips and the absence of marked diseon-

formities in bedding.” would allow such a vegetation cover to develop and be

maintained. This vegetahion cover, together with the facts that hmeltes are

stationary, the rate of supply of material is limited, and most deposition takes

pluce near the Jake shore, can account for the steeper windward slope ot luncttes.

Tt is suggested that « large and continuous supply of material destroys the

vegetation cover and permits the development of a hummocky surface and of

steeper lee slopes. In fact, sume lynettes, for example Cooke Plains lunette

( Fig. 4) and some of those in northern South Australia (C. R. Twidale, persunyl

communication) do liave a hummocky surface due to wind and water erosion,

The 30° dips of the beds in the Cook: Plains limette on the side away fron the luke

presumably represent true foreset bedding, hut the low angle beds in Cooke Plains and Lake

Ciucluite lonettes, although nat characteristic af transverse dunes, are consistent with an aeolian

(rigin,

Servicéton G9OOhrs

| i

Noracoorte OFOChLS Coimas

elle

Calms

Serviceton |5Q00nrs

Yoreetown GFOGhes

Noo cales jeetieded

th SNOWFOWN OF DD hia

Yange a OF Obes Colmes

Vergata 3G hrs Cols

KYAMECUTTA 1500hrs

SNOWTOWN fad brs Cabins

Al,

PARRDANA OF00 hry Cains

PORT LINCOLN O8906 hrs

BERRI OFOO hrs Cayms

ula.

Calms

sills t

PAINDANA [S500 his Calms PORT LINCOLN ‘SDQOh-2 Calms

Calnis

MOUNT GAMBIER @830 his MOUNT GAMBIER 1490 bys

Fig, 9, Wind roses—all winds, The mouthly percentage frequency distribution of wind

directions from ¢ight points af the compass. The sides of the ovtigons face tawards the

cardingl and semicardinal points—north is at the top. Projecting from cach side are twelve

colunms representing the twelve months of the year and the lengths of the columns are propor-

tionul to the percentage frequency of winds from the given direction working round clockwise

from January to December. The solid black part of the column represents the winds of

speeds 13 m.p.h. (21 kilometres per hour) aid greater, and the open part of the colunin

represents the winds of speeds less than 13 xi-p.h. The scale is shown in the ventre of each

wind rose. The percentage frequency of calms in the twelve months is shown to the right

of the wind rose, working from left to right from January tu Devember. The distribution of

winds at 0900 hours is shown for each station and at 1500 hours for some stations, The wind

roses for Mount Gambier show the distribution of winds from sixteen points of the compass.

for 0830 and 1430 hours. ‘The solid black part of the columns represents winds of speeds

15 uup.h. (24 kilometres per hour} and greater, and the open part winds of speeds Jess

than 15 m,p.h.

14 ELIZABETIL M. CAMPBELL

and some, for example parts of Lake Baird lunette (Fig. 2) are asymmetrical with

j steeper lee side, ‘These unusual morphological characteristics of Janettes

represent conditions similar to those of normal dune formation, and can be

expected in the hmette environment as a result of the season! and Jonger tern

yatiations in climalic conditions such as are experienced at preseul.

Similarly, normal variations in the climate of today can explain the multiple

lanettes which developed at various stages in the recession of lake shorelines."

There is little evidence in southern South Australia to support the atrospherie

dust theory and much which contradicts it. On the other hand, there im ample

evidence in support of the deflation hypothesis of lunette formation,

As stated previously, the particle size distribution, the mineralogy, the

quarble mneasures and the sorting values of each Innette sample analysed are

similar to those of the associated lake bed sample. Several components charac-

teristic of, though nat confined to lakes, have been found to luncttes; Hingston

aud Bettenay (1960) reported alunite in lunettes, and gypsum exystals make up

« large proportion of many, such as those at Cooke Plains and Take Fowler

specimens of Coxiella sp. have been found in Lochaber Swamp hinctte and

Jhuraceae oospores im Hiles Lagoon lunctte, The close similarity between Ute

composilon of the lunette and of its assuciated luke bed suggests that the two are

eaisally related, ie, that the material inthe hnictte was derived from the lake

hed. as required by the deflation hypothesis,

Although Hills discounted the likelihood of the formation of agyrcyates: af

particles in the lake beds of north-western Victoria, ageregales have been noted

ihuring analysis of samples from southern South Anstralia, and have bee seen

in transport across dry lake beds in Texas (Coffey, 1909). Although the process

of ageregate formation is incomplelely mmderstood, aggregates are considered to

explain the large proportion of particles in fine-grained Iunettes (76-90% ) which

are ton small to be transported by defation as individual particles, hut whieh,

when bound together, behave as sand grains, The size cornposition of the sandy

Jnmettes is also consistent with the deflation hypothesis.

If the Jake depressions and their associated lnnettes ure not causally related,

the lakes must be explained otherwise than by deflation, In southern Sonth Aus-

tralia eximples can be cited where wave action, solution. blocking of surface

drainage of Huviatile action have contributed at least in part to the origin of the

lake, Lut these proecsses cannot salistuctorily explain all depressions. Owing to

tectonic disturbances and the semi-arid climate of the area, condifiars.are Lavon

able for the creation of basins of interna] drainage. The periodic aceumulation

of water-transported debris in the centres of these basins under conditions of

strong evaporation makes possible the removal of this material by wind, There

are fumerous examples in southern South Australia where depressions have been

formed by this continuing process of deflation. The origin of the lakes is thus

consistent with the deflation theory of limette fornntion.

‘i is gonerally accepted (J, T. Hutton—personal communication) that ery

conditions favour the generation of electric charges on particles, and these

charges murkedly assist in the deflation of material. In southern South Australia

the following air temperature and relative humidity relationships usually apply

(B. Mason—personal communication );

6 Variations in campositian of fanettes associated with the one Jake result front variations

in Che material in the lake bed whieh bas heen transported by strewoss of varied rates of flow or

(iim varivd sources,

LUNETTES IN SOUTHERN SOUTIT AUSTRALIA 105

Air Temperature— FP. Relative Humidity—%

< 60 > 50

60-80 35-55

> 80 <5

The number of days on which the air temperature reaches these categories varies

throughout the arca, bul statistics for Mount Gambier and Yongala represent

vilues near the two extremes:

No. of Days Temp. reached

Air Temperature— PF’. Mt. Gambier Yongalu

< 60 120 LOL

60-50 204 159

> 60 41 105

For i considerable part of the year, dry conditions assist in the process uf

deflation,

However, the deflation hypothesis of lunctte formation, though on hroad

terms consistent with the observed evidence, presents some difficulties.

First, lunettes are common on the edge of salt encrusted depressions from

whieh deflation of the underlying material is impossible. However, the sult erust

may be a more recent development.

Second, although the climate where luncttes are found varies from arid to

subhumid, they are best developed in semi-arid arcas, and not in truly air regions

where deflation is most pronounced.

Third, an examination of the strong winds (greater than 13 miles per hour—

about 21 km, per hour)? for selected stations in southern South Australia shows

that. in the wet season (May to October, based on the uumber of rainy days per

inonth) westerly and northwesterly winds are predominant, whilst in the dry

season (November to April) the stroug winds most commonly blow from a

southerly to westerly direction (Fig, 10). Lunettes occur most commonly on the

eastern or south-eastern sides of lake depressions. That is, they facc the wet

season winds, those which blaw when the lakes are occupied by water, when

deflation from the lake bed is impossible. They are not primarily associated with

the dry season witids which could achieve deflation from the drv lake bed.

These difficulties could he overcome by postulating a change in climate, for

exiunple, a change in temperature-humidity relationships resulting in different

evaporation rates, a decrease in rainfall during the wet season, or a change in

wind direction. However, there is no evidence that such drastic chunpes. ol

ave more commonly required, 13 m.p,b. is also the lower limit of one of the classes on the

Beaufort scale.

Wet

KYANCUTTA

ae; ‘

Wet

PORT LINCOLW

| | N | ‘\

aL.

aes

———— \

a — th |

ai Ca ied

‘Ss a ra ]

AQ fe “a a

a Wer s .

‘ ee Dry i Wet

5 Mes “

tee

PARNDANA SERVICETON

MOUNT GAMBIER

BERRI

Fig, 10, Wind roses—strong winds in the wet and dry halves of the year. These wind roses

show the distribution of strong winds from the cardinal and semicardinal points of the compass

in the wet and dry halves of the year, differentiated on the basis of numbers of rainy days per

month, and averaged over the selected stations, The monthly percentage frequency of the

strong winds (13 m.p.h. (21 kilometres per hour) and greater) at 1500 hours for November

to April and for May to October are expressed as a percentage of strong winds for the year.

LUNETTES IN SOUTHERN SGUTH AUSTRALIA 107

Price and Kornicker (1961) nated the importance of waves aid currents in

transporting shell fragments on to the lower slupes of clay dunes similar to

Irmettes. in Texas. Bowler (1964) suggested that medium io coarse sand is con-

centrated in beaches before being blown by the wind on to the lunctte, allhough

finer particles, le considered, could not be concentrated in this way. However,

although silt- and clay-sized particles are generally carried in suspension in water,

some may be conventrated in a beach by one or more of the following methods,

Claw particles may adhere to sand grains or themselves form aggregates. The

salinity of the water in the lake will determine whether the clay particles wre in

(vue colloidal suspension, or are capable of transport by wives in the form ot

agerceutes, As J, T. Hutton pointed out (personal conwiunication) wave acting

sufficient to cause transport would probably also be capable of breaking up the

aguregates; but many remained unseparated after mixing during the analysis of

samples. Silt and clay in susperision may be filtered out by the sand and debris

ina beach or hy vegetation growing on the edge of a lake, It is suggested, there

force, that wave transpnrt of sand, and possibly of silt- ancl clay-sizecl particles, is

important in lunette formation.

As Stephens and Crocker (1946) pointed ont, the vegetation growing on the

vlge af a lake provides the rongh type of surface required ta cause deposition of

wind-Lransperted material, The line of vegetable matter, floated on the surface

of the Jake and deposited as the water level recedes. has a similur result, as was

recognized lyy Woods (1862, pp, 28-29).

The effect of waves and the presence of vegetation on the edge of the lake

“an overcome each of the difficulties encountered by the deflation hypothesis,

First, in the case of those lunettes occurring on the edge of the salt-encrusted

lakes, the major part of deflation ocenrs from a beach built hy wave action in fhe

wet scason, and not from the luke bed. Second, the reason for lunettes being best

developed in semi-arid rather than arid regions, is that in arid areas there is

insufficient vegetation on the lake edge to Uap the material which has been

scourcd from the lake hed and to prevent later degradation, and the lakes in arid

areas do not contain water lony cnough for significant wave transport* to occur,"

The third diffienlty concerns the distribution of the Junette in relation to the lake.

The position of the Junette on the eastern or south-eastern side of the lake oppasite

the wet season winds is a result of wind generated waves. A beach is built on the

castern and south-eastern side of the lake, and the material in the beach is able to

he transported by the wind. In the dry season, once the water level has fallen

and with the winds blowing from the west and south-west, deflation from the lake

hed takes place. In this way, « cambination of wave and wind action results in the

formation of the lunette on the eastern side, and where waye action and wind

transport in winter are relatively more important on the south-eastern side. ‘hus,

the anulogy of hinettes with primary coastal foredunes is readily apparent.

ACKNOWLEDGEMENTS

The research. reported in this paper was made possible by a grant from the

Uprversitv of Adelaide. A mineral analysis of samples from lunettes and their

associated lake beds was carried ont by Australian Mincral Development Labora-

5 1n suh-humid vegians. the raintall is sufficient to allow the development of co-erdinaled

drainage systeros rather than dry busing of internul druinage,.and the evaporition mite is lower.

Therefore, deflition is limited,

‘In several districts in sootherm South Australia, for example near Snowtown, sume likes

have associated lometles. whilst others do not. Such factors as the amonnt aod nature of material

curried Inte the lake, and the length of time the lake is oceupied by water (enabling wave

transport) or is dry (enabling deflation) must be considered in explaining this distribution.

Ws ELIZABHTH M, CAMPBELL

torics, Officers of the Bureau of Meteorology, Adelaide, kindly supplied informa-

tion [rom which the wind roses were drawn. The thanks of the writer are due to

Dr C, RK. Twidale, Department of Geography, University fo Adelaide, for helpful

advice on many aspects of the research, and for a critical reading of the text, and

to Mn. J. T. Hutton, of the Division of Soils, C,S.1.R,0,, Adelaide, and Mr. B.

Mason of the Department of Geography, University of Adelaide.

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384-97.

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Memoir 11, p. 74

Piate J {LIZABETH M. CAMPBELL

PLATE 1

Hitton’s Lagoon linette—aerial view. The smoothly rounded eastern shore is typical of the

lakes in southern South Australia, The dashed line indicates the position of the crest of the

lunette. (Reproduced by courtesy of South Australian Department of Lands. )

EvimaBetru M, CAMPBELL

Phare 2

White Lagoon multiple lunette—ae view. White Lagoon is an example of the more or less

concentric arrangement of a multiple lunette bordering ore lake. The three ridges are marked

by dished lines. (Reproduced by courtesy of South Australian Department of Lands. )

PLATE 3 Exurzazetru M, CAMPBELL

PLATE 3

Salt Lake multiple lnnette—aerial view. A complex of small lakes, each with its associated

lunette or limettes. occurs within a larger lake basin on the edge of which is a large lunette.

(a)

(b)

(c)

(d)

(Reproduced by courtesy of South Austraiian Department of Lands. )

PLATE 4

(See next page )

Western slope of Bool Lagoon lunette. The smooth and rectilinear western slope of this

lunette is typical.

Bedding in Cooke Plains lunette near the crest. The dip of the beds is variable over short

distances. but nowhere exceeds 20) degrees. View looking north-east.

Bedding in Lake Cadnite lunette. The characteristic dip of the beds in this lunette near

the crest is 2 to 12 degrees to the east. The example shown here indicates a rare cross-

bedded sequence.

Shoreline of Lake Greenly, Gneissic material outcrops on the south-eastern shoreline of

Lake Greenly. A shore platform and a beach with rounded pebbles are evidence of the

effectiveness of waves in transporting debris even in this shallow lake. The white material

near the top of the photograph is “Hour” gypsum on the surface of the lunette. (Photo-

graph by C. R, Twidale. )

PLari 4

CAMPBELL

ELizABETH

ORIGIN OF WAVE ROCK, HYDEN, WESTERN AUSTRALIA

BY C. R. TWIDALE*

Summary

Wave Rock is a long high overhanging natural wall located on the northern side of Hyden Rock, in

Western Australia's Wheat Belt. Though it displays some apparently unusual features, it has much

in common with similar forms described from Eyre Peninsula and other areas both within Australia

and overseas. The available evidence suggests that Wave Rock also evolved in similar fashion: by

strong scarp foot weathering, and subsequent erosion of the weathered debris. This and other

hypotheses advanced in explanation of Wave Rock are discussed in light of the field evidence.

ORIGIN OF WAVE ROCK, HYDEN, WESTERN AUSTRALIA

C. R. Twipate*

[Read October 10 1968]

SUMMARY

Wuve Ruel isu long high overhanging natural wall located on the northern

side of Myden Rock, in Western Australia’s Wheat Belt. Though it displays some

upparertly unusual features, it has much in common with similar fornis deseribed

from Eyre Peninsula and other areas both within Australia aud bverseas. The

available evidence suggests that Wave Rock also evolved in similar fashion: by

strong scarp foot weathering, and subsequent erosion of the weathered debris,

‘Chis and other hypotheses advanced in explanation of Wave Rock are discussed

in ight of the ficld evidence,

INTRODUCTION

Various opinions haye been offered concerning the origin of Wave Rock, a

well-known and spectacular feature located near Hyden, W.A. In Land of the

Southern Cress (1956), for example, a popular picture book designed to attract

immigrants and tourists, it is attributed to the work of “wind and rain” (p, 89). In

Vincent Serventy’s Landforms of Australia (1967, p, 25) it, and similar sharply

sigmoidal or flared forms at Ayers Rock, are tentatively attributed to erosion hy

wind-driven sand. An alternative explanation has heen offered by O'Riley (1967)

who, citing professional opinion, suggests that Wave Rock is due fundamentally to

selective erosion by running water. It is suggested that the upper, and earlier

exposed, surface of Hyden Rock, on which Wave Rock occurs, has been indurated

with silica to 4 greater extent than the more recently revealed flanks, Water

coursing over the bare rock surface therefore crodes the weaker lower slopes more

than the toughened upper surface; the lower slopes are “undercut by the contin-

ued flow of water”; and the stone wall erected around the flattish upper surface of

the rock is intended to “arrest erosion”.

Comparable, though less spectacular, flared forms on Eyre Peninsula (S.A, )

and elsewhere (Twidale, 1962; 1964; 1965, pp. 145-147, 347-350) are evidently

due to strong localised subsurface scarp foot weathering and subsequent erosion.

Because this interpretation is at variance with the opinions cited above concerning

the origin of similar forms, it may be both useful and interesting to describe the

process of scarp foot weathering and its characteristic results; and to match the

several hypotheses with the field evidence at Flyden Rock,

SCARP FOOT WEATHERING AND EROSION

Flared forms developed on granitic rocks have been examined on Eyre Penin-

sula, in the Kosciusko region of southern N.S.W., in the western Mt. Lofty Ranges

(S.A.), and in the Sierra Nevada of California. They occur on sandstone outcrops

in the Flinders Ranges, and at Ayers Rock, as well as on the conglomerate of the

Trans. Roy. Soc. 5.A. (1968), Vol. 92.

Olgas massil, Norther Territory, Although Hares are well developed on the north

tn west facing slopes, and although exceptions have been noted, they are best

and most commonly developed on the southern and eastern, ie. the shady, aspects

af the residual hills or inselbergs on whicl they occur. Double and multiple Hares

nr Convavities are in places displaved. The flarcd slopes are not everywhere found

in horizontally disposed zones of steepening or overbang. At several sites they are

inclined or even irregularly distributed, though in all cases the disposition of the

steepened zone varies in evident sympathy with that of the hill-plain or rock-soil,

junction. Flares are very well developed on the points of spurs.

Incipient flared slopes are developed beneath the land surface, beneath

weathered country rock which is in situ. At the margin of the inselberg, the suv-

face of the fresh rock, continued subsurface as the weathering front (Mabbutt,

1561) or limit af weathering, characteristically phmges steeply for a few fret

heneath the land surface before gradually fattening out. Snch concavities in the

fresh bedrock surface, and formed beneath the natural land surface, have: been

observed in dry reservoirs located on the inselberg margins, and have also heen

detected by augering und excavation,

Finally, small flared slopes oceur in clefts within the inselhergs, around

boulders and on gentle slopes, in association with accumulations of soil or weath-

erect debris,

Although several possible explanations of flared slopes have been entertained

(Twidale, loc. vit. 1962; op. cit. pp. 347-350), all save one are found wanting

when the field evidence is idtched against the deducible consequences of the

hypotheses. The interpretation which has survived rigorons testing and which

acuournts for the field evidence is that the flares evolve in two distinct stages

involving first, strony lovalised scarp foot weathering, and, secon], marked) difter

ential erosion of the scarp foot zone and exposure of the erstwhile weathering

front (Fig. 1).

HYDEN ROCK

Ceneral Setting

Hyden Rock lies about 3 km, east of Hyden township, which is, in turn, 296

km, E.S-E. of Perth (Fig. 2). The Rock is about 2 km. long on an east-west axis

and on a N.N.W-S.S.E. line a litde more than 0:8 km. wide (see Fig. 3), It is

composed of granite, most of which is strongly porphyritic, though pegmatilic,

even and medium grained, aid fine gramed, phases occur in patches and veins,

No horizontal dat-lying zones of petrological differentiation have been detected in

the granite.

The granite is jointed but most of the joints are tight and widely spaced.

Avenues of weathering within the Rock are few, and its greater resistance to

weitherng aud crosion, as evidenced by ils remaining as an upstanding hill, is

probably due to its being built of especially massive joint blocks, This is demrons-

trably the case in places on Eyre Peninsula and elsewhere,

Hyden Rock is a granitic inselherg which rises abruptly to a maximum eleva-

tion of some 55 m, above the surrounding plains, whieh, on the evidence of lores

wd dam excavations, are also underlain by granite, Tere, however, the rock is

deeply weathered: Berliat (1965) reports over 20 m, of weuthered grunite wbuut

400m), from the margin of a residual located 63 km. east of Hyden and well over

AQ nm. of disintegrated rock elsewhere beneath the plains.

The plains are not Aat, for there is a pronounced slope down to the north; the

lew vaguely defined watercourses How toward a complex of ephemeral Jakes

ORIGIN OF WAVE ROCK, ITYDEN 7

| | F

pr --2=-"F71', basistaet during dry season

! F

Fig. 1. Evolution of flared slope

by scarp foot weathering and sub-

sequent erosion, (a) Strong

weathering by water (derived from

run-off) at the lower margin of

the residual hill. Numbered lines

represent stages in advanue of

weathering front. (b) Lowering of

base level, erosion of weathered

debris, and exposure of weathering

front as a flared slope, with

further weathering beneath the

new plain surface, (c) Develop-

ment of double flare by repetition

of the twa-stage process.

c , -}-

located immediately to the north of the Rock. The plain to the south of the Rock

is about 25 m. higher than the plain on its northern side,

The inselberg consists of three distinct hills, the central dome-shaped hill

being both slightly higher and more extensive than its neighbours to cast and

west, All three hills are delineated by prominent vertical or near vertical joints,

the more important of which run E.N.E.-W.S.W., N.W.-S.W., and N.W.-S.E. Many

clefts within the hills follow along such joints. Flat lying or gently dipping joints

are also important, for, although discontinuous, they subdivide the hill into mas-

sive slabs or sheets. The weathering and erosion of several of the latter gives the

Rock its stepped appearance in broad view ( Fig. 4).

Flared Slopes

Flared and basally steepened slopes are well developed on all three hills, not

only around the margins but in clefts (Pl. 1) and amphitheatres at higher levels

also. The flared and steepened slopes are well formed on the long southern wall of

the Rock, for instance, adjacent to the 13th fairway of Hyden Golf Club (A on

Fig. 3) and they are well developed along much of the northern foot wall. In

clefts and depressions the south-facing wall is commonly, though not in all cases,

steeper than that opposite,

118 Cc. R. TWIDALE

PERTH

@Hyden

Sr & ——______ limit of granite outcrop

major joint

watercourse, loke

ming kts road

0 Tmile

——

0 Vim.

Fig. 2 and Fig. 3, Location map and map of

Hyden Rock (drawn from W.A. Lands Dept.

air photographs). Sites A-D are referred to in

the text.

N. sheet >, S.

boulders formed by 25

disintegration of sheet ee SS

gm.

incipient flare —-"S, 3

Bet 25

Fig. 4. Diagrammatic N.-S. section, not to scale, of Hyden Rock, showing stepped form, sheet

structure, flared slopes, probable present weathering fronts, and the disparity in elevation

between the plain north and south of the Rock.

ORIGIN OF WAVE ROCK, HYDEN 119

But what is unquestionably the most majestic and impressive Aared and over-

hanging slope is Wave Nock (PI. IT) which is 4 steepened and overhanging hasul

slope 10-12 m. high located on the north side of the central hill, Some fared

slopes on Pildappa Hill, Eyre Peninsula, are as high or higher than Wave Rock,

some at Ucontitchie Hill (also on Eyre Peninsula) overhang toa greater extent or

are more complex, but as a high continuous overhanging wall, Wave Rock indubi-

tably stands alone.

At the base of the flared slopes on the northern side is a continuous, virtually

level platform up ta 10 m. wide (Pl. IL) and located at, or a litle above, the

present plain level. Similar platforms have been observed marginal to the insel-

bergs of northwestern Eyre Peninsula, hut nowhere are they as extensive and level

as at Hyden. Another notable feature is the angular junction between flared slope

and platform. Double concavities or flares are fairly common, as. for instance, on

Wave Rock itself (PJ. IL), Some flared zones on Hyden Rock are inclined with

respect to the horizontal, as, for mstance, on the northern slope (PI. IIT), in the

enclave hetween the central and eastern hills. Finally, it is very typical that the

weathering front—the surface between the fresh and the weathered bedrock—

plunges steeply beneath the weathered granite which, with a veneer of colluvial

debris, underlies the plain (Pi. IIL). Excavations reveal that this steeply inclined

slope continues beneath the plain surface for rather more than one metre, a

which depth it shows ne sign of levelling out.

Thus, in many respects the flared slopes at Hyden Rock, and particularly

Wave Rock, share characteristics with similar forms in other regions. The great

contrast lies in Wave Rock itself, which is on the northern, and not the southern

side of the inselberg, and which is located not on the point of a spur, but in a

broad, prohably joint-controlled, crbayment (B on Fig. 3). These apparent

anomalies are, however, susceptible of ready explanation in terms of the

subsurface water weathering hypothesis.

Being in a broad cmbayment, to which drains nim-off from a wide area of

the central hill, the plains marginal to the inselberg in that area undouhtedly

reocive a more than average quantity of water, so that especially pronounced and

deep scarp foot weathering is probably developed there. Furthermore, inmedi-

ately to the north of the Rock and virtually at the same clevation as the plain on

the northern side is a complex depression—probably an old drainage line—which

receives water from a wide area and which in winter is full of water. Ground-

waters from this drainage line may penetrate to the Rock itself.

The comparative degree of erosional exposure on the northern and sonthern

aspects of Hyden Rack should also be considered. As already mentioned, the

northem plain, clase to the drainage depression, is about 25 m. lower than the

plain to the south of the Rock (Fig. 4): indeed, the whole plain surrounding the

inselberg slopes down to the north, Any lowering of the plain by strcams must

have been initiated from the old drainage line ‘ Fig. 3) aid. therefore would have

affected the northern slope of Hyden Rock before the southern. Thus, it may be

argued that while greater erosion and lowering of plain level has already exposed

the deep weathering front on the northern face, similar lowering of the surfaer

has not yet extended to the southern side. where an incipient flared slope may be

present beneath the surface (Fig. 4).

Thus, local circumstances may have combined to cause unusually deep and

pronounced subsurface weathering of the granite on the northern side of Hyden

Rovk. The same northern slope may have been exposed by erosion to a greater

extent than has the southern. These are the probable reasons for the wnhusual

degree of steepening of the northern slopes, and for Wave Rock in particular.

120 CB. TWIDALE

The other suggested mechanisms tor the development of Waye Rock either

cannot explain the field evidence, or are inconsistent with it. Por instance, aeolian

sand blasting cannot account for the occurrence of incipient Hures beneath the

sresont land surface, below weathered granite in site, and for the fares at high

Fevels an the Rock (sand blasting is most effective within 1 m. or so of ground

level). The areal distribution of the fares is also at odds with the éxpectable

consequences ot sand blasting. Luncttes duc to deflation of exposed lake deposits

are located on the western side of lakes in the Hyden-Corrigin area, indicating a

prevalence of strong winds from the east, and in these terms the “undercut” slopes

slintilel be best developed on the eastern aspects of the hills, No such consistent

and preferred orientation has been detected. Furthermore, the very fact of the

inselberys being there would disturb the air flow and divert winds (and sand}

around the residuals; the abundant vegetation growing aronnil the hills ar the

better watered scarp foot zone also serves to protect the lower rock walls against

avalian action.

The suggestion concerned with running water (O'Riley, loe. cit.) faces oom-

parable difficulties:

(1) Induration of the snrtace of Hyden Rovk has certainly acewrred, partly as

a result of climatic conditions, partly resulting from the work of the lichens which

grew in profusion on the rock surface. Iron oxides have been concentrated! at and

newr the surface, though much of this indurated layer appears to have heen

eroded, only small Hat-topped nobs remaining in a few places. But the induration

appears equally distributed over most of the surface, It is certainly aol preferen-

tially developed at upper levels of the Rock surface, so that selective crosion

cannot be invoked. urthermore, though the upper area must have been first

exposed, there is a strong suggestion that far from being tougher than the lower

slopes, the upper part of the rock has suffered cousiderable weathering, the upper

sheets of granite having disintegrated iuto groups of blocks or ronnded boulders.

(2) Some form of induration has undoubtedly affected many boulders, and

preferential weathering has given rise to cayernous forms or tortoiseshell rocks.

But in these forms the hardened crust forms thin projecting lips or visors, which

da nat oceur in the flared slopes. At one site on the northern slope of eastern hill

(C on Fig, 3) there is a concentration of cavernous and alveolar weathering

which may be due to petrological factors, or may be a reflection of local concen-

trated weathering at or just below a former plain level (the site is some 6 m.

above the present plain). In this area, quite deep hollows are developed beneath

thin external crusts,

(3) Although rimning water, hecanse of its downslope increase in velocity

and volume, may appear capable of producing slopes of irereasing downslope

inclingtion (see for instance Fenneman, 1908; Cotton, 1952), particularly in an

embayment such as those at Wave Rock and at D (Fig, 3) on the southern side of

Tlyden Rock, where flow is centripetal, there are sound hydrological reasons for

suggesting that it dacs not do so (Twidale, loc, cit., 1962). Moreover, ounce a

cortain critical inclination is atlained, fast Howing water becomes detached fram

the rock surface to form a jct or waterfall. Only the slowest flowing thin films of

water adhere to the steep rock face by surface tension—vyet these are supposed to

aucomplish considerable erosion, and even pronounced undercutting. At Wave

Rock in June 1968, some thin films of water were creeping down the overhanging

slope, but there was a continuous drip of water from the upper lip to the foot

wall. Also, on the upper part of Ilydun Rock, pronounced overhanging Mares are

develoned on spurs of very limited catchment, and.in other places on boulders of

similarly limited potential run-off,

ORIGIN OF WAVE ROCK, HYDEN 121

(4) Selective erosion by running water cannot explain multiple flares, the

development of fares in clefts and amphitheatres (nor, indeed, the amphithcatres

themselves), the inclination of the flared zone, and their apparent occurrence suly-

surface marginal to the Hyden residual.

The searp foot weathering hypothesis can readily account for these and other

relevant details of the flared forms, and it is suggested that it is this process which

is responsible for the development of Waye Rock and other similar features at

Tyden Rock, as well as on other granite residuals in the southwest of Western

Australia.

Angular Footwall

The pronounced angularity of the junction between the fared slope and the

basal platform, us displayed at Wave Rock, and other similar features at Elyden

Rock and other inselbergs in the region, develops as a result of subaerial weather-

ing, Ramwater and seepage waters trickle down the steepened lower slopes and

cause the platform immediately adjacent to the hillslope to be wetted. The rnck

here disintegrates, and is subsequently washed out, leaving a miniature depres-

sium, one flake (1-2 ems) deep between the main platform level and the base of

the fared slope, Such depressions, observed at the base of Wave Rock and on

Gorge Rock, near Corrigin, W.A., carry water after rain, causing further weather-

ing of the granite with which it comes into contact, inclnding that exposed at the

searp foot, As the shallow depression is extended laterally by such weathering, the

basal slope is undermined and steepened, and an angular junction is deyeluped

between hillslope and platform (Fig. 3).

Amphitheatres

Another feature well displayed at Hyden Rock, but not so far described from

the Eastern States. is the amphitheatre, Joint clefts with flared hounding slopes

occur on Hyden Rock (Pl, T) and, for instance, on several of the inselhergs of

northwestern Eyre Peninsula. These two areas have similar clirmates (both with

hot summers, coo! winters, 35-45 cm. average annual rainfall and a marked dry

season), but at Hyden Rock the slopes of the clefts have suffered marked reces-

sion as a result of soil moisture weathering and as a result have been widened to

form large amphitheatres (Fig. 6). It is not known whether this more advanced

stage of development reflects a time factor or the less resistant character of the

porphyritic granite exposed at Wyden Rock, but such developments clearly contri-

brite ta ihe breakdown of the sheets of granite and to the formation of an overall

stepped morphology on the inselbergs.

CONCLUSION

Though Wave Rock displays apparently unusual features, it hag much in

common with similar forms described from Eyre Peninsula and elsewhere. The

available evidence suggests that Wave Rock evolved, as did the other flared slopes,

by strong scarp foot weathering followed by erosional exposure. This interpreta-

tion accounts for the field evidence more readily than any of the other hypotheses

so far advanced.

ACKNOWLEDCEMENTS

The writer wishes to thank Professor M_]. Webb, Department of Geogniphy,

University of Western Australia, and the Director, Geological Survey of Western

Australia, for facilitating in vations ways the investigation of Ilyden Rock and

several other inselbergs in the region. Some travel expenses were met from a

Research Grant from the University of Adelaide,

C. R. TWIDALE

Fig. 5. Development of an angular junction

between flared slope and platform by local

subaerial water and pool weathering. (a)

Water trickling down flared slope soaks the

bedrock at the inner edge of the basal platforin

and. weathers it. (tb) The weathered debris is

washed away, a shallow pool of water accumu-

lates in the hollow so formed, and further

weathering takes place. (e) The base of the

shallow pool flare is steepened and made angular as a result

\ of weathering by pool water, and consequent

b - 7 lateral extension of the pool depression.

grenite debris

*_weathered

granite

Fig. 6. Sequential development of an amphitheatre from a joint-controlled cleft by soil mois-

ture, weathering, undermining of bounding slopes, and progressive widening. Several such

phases of weathering, with intervening spells of debris evacuation, are evidently represented in

these clefts and depressions with complex sides. ‘The phases of weathering and erosion in the

clefts, like those to which the flared stoned are attributed, are probably controlled by climatic

changes or fluctuations or even seasonal and storm effects. a-d represents a sequence, represen-

tatives of which have been observed on Hyden Rock. d1 shows the result of baselevel lowering

in a narrow cleft (cf. Pl, 1). The depth of fill must decrease as the area of the depression floor

increases,

ORIGIN OF WAVE ROCK, HYDEN 123

REFERENCES

Anon., 1956 Land of the Southern Cross. Australian Publicity Council, Melbourne, Australia;

BERLIAT, Xe, 1965 Exploratory drilling for water in the sand-plain country 40 miles east of

Hyden. pp. 8-9 in Annual Report (1964) Geol. Surv. W. Austr., Perth, 72 p.

Corton, C, A., 1952. The erosional grading of convex and concave slopes. Geog. Jour., 118,

pp. 197-204.

FENNEMAN, N. F., 1908. Some features of erosion by unconcentrated wash. Jour. Geol., 16,

pp. 746-754.

Massutt, J. A., 1961. “Basal surface” or “weathering front”. Proc. Geol. Assoc. London, 72,

pp. 857-358.

O’Riey, [., 1967. Wave that turned to stone. Walkabout, 33, (6), pp. 12-15.

ServeNTY, V., 1967. Landforms of Australia. Angus & Robertson, Sydney, 132 p.

Twipace, C. R., 1962. Steepened margins of inselbergs from north-western Eyre Peninsula,

South Australia. Zeitschr. f. Geomorph. (N.S.), 6, pp. 51-69.

Twimate, C. R., 1964. Contribution to the general theory of domed inselbergs. Conclusions

derived from observations in South Australia, Trans. and Papers Inst. Brit. Geogr., 34,

pp. 91-1138.

Twwateg, C. R., 1968. Geomorphology, with special reference to Australia. Nelson, Melbourne,

A406 p.

. BR. Twipate

PLates 1 to 3

(CSTEPIAL “WD Ofoyd) — “yloursnc ute

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‘I

OBITUARY: THOMAS DRAPER CAMPBELL 1894-1967

Summary

OBITUARY

THoMAs Drarer Cantppen. (1894-1967)

Born at Millicent in the South-east of South Australia Campbell received his early

education at Prince Alfred College and undertook tertiary study ut the University of Adelaide

where he graduated in dental surgery in 192! continuing these studies ta receive a doctorate

in 1923, His doctoral thesis was published by the University in 1925 under the title “Dentition

ae rain of the Australian Aboriginal” and has been widely acclaimed as a pioneer work

in this field.

Soou after graduation Campbell entered the Dental Department of the Royal Adelaide

Nospital as the first Dental House Surgeon, and in 1926 was appointed Superintendent. He

wus elected Dean of the Faculty of Dentistry in 1938, and appointed to the full-time University

position of Director of Dental Studies in 1949. He became the first Professor of Dentistry in

1954 antl upon retirenent four years later was made Professor Emeritus,

During his professional career Prof, Campbell received high recaguition for his work, In

1948 he was made F,D.S.B,C.S, (London); in 1959, F.D.S.R.C.S. (Edinburgh): iv 1952 he

wus. clected an Honorary Member of the Qdontological Section of the Royal Society af

Medicine, London, and in 1966 « Fellow af the Australian College af Dental Sargeurs,

At all limes Professor Canipbell combined his dental studies with a keen interest in

anthropology. In December 1926 he was associated with the formation of the Board for

Anthropologic:l Research, University of Adelaide, and has played a leading role in its research

activities. As organiser and leader of many expeditions ta Central Australia in the 1920’s and

30's he was responsible for the collection of a large amount of important data and material on

many aspects of the traditional life of the aboriginal, Storie technology fascinated him and he

spent countless hours searching for and making a detailed study of aboriginal stone implements.

He contributed important papers to, the literature on this subject. Well known. as an aecom-

plished amateur film producer, Professor Canmphell made a series af colour-sound movics on

ge bf aboriginal technology for the Board. He was chairman of the Board at the time of

tis death;

He became a member of this Society in 1922 and spent ten years on the Council, holding

the office of President in 1934-35. Following appointment to the Board of the Public Library.

Musenm and Art Gallery in 1932, he became a foundation member of the Museu Board in

1939, a position which he held until his death. In addition to being a Fellow of the Australian

Research Council, Foundation Member of the Australian Institute of Aboriginal Studies and a

Lite Member of the Anthropological Socicty of South Australia, Professor Campbell was an

panty member of the Anthropology Staff of the South Australian Musewn from 1923 until

us death,

Te was well known as a keen debater and will be remembered for lis participation in

many lively discussions at Royal. Society meetings. A member of this Society with an out-

standing international reputation, he died at his home at Tusmare on 8th December, 1967,

Robert Edwards,

BALANCE SHEETS

Summary

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REPORT ON ACTIVITIES OF THE COUNCIL, 1967-68

Summary

REPORT ON ACTIVITIES OF THE COUNCIL, 1967-68

Meetings

The usual eight ordinary meetings were held in the Society's rooms during

the past year at which the attendance averaged 32, One special meeting was

held on December 7th to ratify the new Rules and By-laws,

A total of 12 papers were read devoted to the following disciplines:— Zoology

8, Botany 3, Geology 1, Lectures were given at each meeting and two exhibits

presented,

Membership

Twenty-two new members were elected during the year and nine resigna-

tions were received. The membership now stands at 266, the highest number in

the history of the Society.

The Council records with great regret the death of Fellows; Emeritus Pro-

fessor T. D. Campbell, a past president of the Society; Sir Tom Bart Smith and

Mr, J. K, Powrie.

Rules and By-laws

During the year the new set of Rules and By-laws approved at the December

1967 meetings Was issued to all members. One amendment has since been passed

at the meeting of September 12th, 1968. Rule VI (2) now reads;—

The nomination furm shall be lodged with the secretary and shall be sub-

mitted to the Council at its next meeting, Upon approval by the Council the

nomination shall be submitted to the next meeting of the Society and an

election shall be held at the next meeting of the Society thereafter,

Sub-Committees of Council

The business of Council has been facilitated by the work of four sub-

committees ;—

Library Committee—management of the Library

Publications Committee—production and maintenance of standards in the

Transactions and any other publications of the Society

Awards Committee—nominations to the Council of names of members pro-

posed for awards by the Society and other bodies

Research and Endowment Fund Committee—manayement of investment

funds.

Library

The library has operated efficiently and profitably during the year, the

Library Account having a credit balance of $1630 as at June 30th 1968. The

Council wishes to express their grateful thanks to the Assistants, Mrs. Dunlop and

Mrs. Dougal for the services they have rendered to the Society both in the library

and in assisting the Treasurer.

Rechecking of the bookstock has resulted in 200 new entries being forwarded

to the C.S.1.R.0, Index,

Fifty volumes have been bound and another 160 prepared for binding.

The exchange list has been completely reviewed and adjustments made

129

where necessary. Some 340 journals are now received on exchange from 46

countries, 10 new exchanges were negotiated during the year. In addition 37

continuing subscriptions are now in operation,

Altogether 245 volumes were borrowed during the year, mainly on inter-

library loan within the state, interstate and with New Zealand.

Author and subject indexes for volumes 45 to 91 have been prepared for

publication.

Publications

Vol. 91 of the Society's Transactions was published in December 1967, It

contained 204 pages compared with 191 pages in Vol. 90. The cost of printing

seems to rise steadily each year but it has been possible so far to include all

papers received before the June Council meeting which is normally taken as

the final date for acceptance of material for the current volume.

Research and Endowment Fund

As a result of moves which were initiated two years ago, to improve the

management of the fund, its finances are now on a reasonably sound basis.

Accordingly this year it was found possible to make available some of the income,

in the form of grants, to aid scientific research. Two grants for the calendar

year 1968 were made, Dr, C, R. Twidale was awarded a grant to study the

geomorphology of the Arcoona Plateau, and Mr. I. M. Thomas and Mr. S.

Shepherd on behalf of a group of workers will receive a sum of money to

finance an expedition to Pearson Islands to study the flora and fauna of the

islands and adjacent waters.

Verco Medal

The Council awarded the Verco Medal for the year 1968 to Mr. R. C. Sprigg.

OFFICERS FOR 1967-68

Summary

130

ROYAL SOCIETY OF SOUTH AUSTRALIA

INCORPORATED

Patron:

IUS EXCELLENCY LIEUTENANT GENERAL SIR EDRIC M. BASTYAN,

K.C.M.G., K.B.E., C.B.

COUNCIL FOR 1967-68

President:

K. R. MILES, D.Sc., F.G.S.

Vice-Presidents:

H. B. S. WOMERSLEY, D.Sc. F. J. MITCHELL

Treasurer:

Secretary:

S, A. SEIEPHERD, B.A., LL.B.

W. K. HARRIS, B.Sc.

Editor: Assistant Editor:

1. M. THOMAS, M.Sc., M.1.Biol.

J. K. TAYLOR, B.A., M.Sc., B.Se.Agr,

Programme Secretary:

Librarian:

Db. E, SYMON, B.Ag.Se.

N. H. LUDROOK, M.A., Ph.D., D.LC., F.G.S,

Members of Council:

J. A, PRESCOTT, C.B.E., D.Se., M, J. TYLER

F.RS., FAA, FRAC.

J. T. HUTTON, B.Sc., A.S.A.S.M.

K. E. LEE, D.Sc.

C. B. WELL, M.Ag.Se.

H. E. WOPFNER, Ph.D.

Auditors:

Messrs. MILNE, STEVENS, SEARCY & CO.

LIST OF MEMBERS

Summary

131

ROYAL SOCIETY OF SOUTH AUSTRALIA

LIST OF NEW MEMBERS

Apam, J. R., B.E., A.M.Aust..M.M., Dept. of Mines, Box 38 P.O., Rundle Street,

Adelaide, S.A,

Ancove, P. C., R.D.A., M.A.LA.S., Dept. of Agriculture, Gawler Place, Adelaide, S.A.

von Bexrens, D., Naracoorte High School, Naracoorte, S.A.

Bursserr, A. H., M.Sc., A.M.AustJ.M.M., F.G,§., Dept. of Mines, Box 88 P.O,,

Rundle Strect, Adelaide, $.A,

Bonsai Miss S., BA. (Mus.), Dip. S.T., Bedford Park Teachers College, Bedford

Park, S.A,

Cannick, R., Ph.D., Mawson Inst. for Antarctic Research, University of Adelaide.

Davy, R., B.Sc. (Hons.) Ph.D., L.RB.LC., F.G.S., C/o AM.D.EL., Conyngham

Street, Frewville, S.A.

Dyxr, N, W.. A.C.A.A., Mem. Inst. Inst. & ‘Coni.. Aust,, Seaton, S.A.

Ernst, L. K., 50 Airdrie Avenue, Findon, S.A,

Famsurn, W., B.Sc. (Hons.) A.M.I.M.M., F.G.S., Dept. of Mines, Box 88 P.O.,

Rundle Street, Adelaide, S.A.

TlENnnincsen, M.. B.E., F.S.A.S.M., A.M.LE., Aust,, A.M.Aust.LM,M., Dept. of Mines,

Box 38 P.O., Rundle Street, Adelaide, S.A,

Hiren, M. N., B,Sc., Dept. of Mines, Box 38 P.O., Rundle Street, Adelaide, S.A.

Huronson, A, R., 218 Shepherds Hill Road, Bellevue Heights, S.A,

Jenxins, R. J. F., B.Sc. (Hons.), Dept. of Geology, University of Adelaide.

Kennepy, Miss G. R., B.Sc., Ph.D., Flinders University, Sturt Road, Bedford Park,

S.A.

McGowan, B., B.Sc., (Hons.), Ph.D., Dept. of Mines, Box 38 P.O., Rundle Street,

Adelaide, S.A.

Mutter, P. G., B.Sc., (Hons,), A.M.Aust.LM.M., Dept, of Mines, Box 38 P.O.,

Rundle Street, Adelaide, S.A.

Mooxcrort, E,, B.Sc., Dept. of Mines, Box 38 P.O., Rundle Street, Adelaide, S.A.

Moutns, M, S., F.R,E.S,, 14 Chisholm Street, Greenwich, N.S.W. 2065.

Otsen, A. M., M.Sc,, Dept. of Fisheries and Fauna Conservation, Adelaide, S.A,

Wu.son, R. B., B.Sc., (Hons.), 22 Chetwynd Street, West Beach, S.A.

Wo aston, Miss E, M., Ph.D., Botany Dept., University of Adelaide.

LIST OF LECTURES AND EXHIBITS, 1967-68 AND

AWARDS OF THE SIR JOSEPH VERCO MEDAL

Summary

LIST OF LECTURES GIVEN AT MEETINGS DURING

THE YEAR 1967-68

Sept., 1967 Dr, H. B.S. Womenstry: “Aspects of Coral Reef Biology”.

Oct., 1967 Mr. C, A. Martin: “Moomba, a South Australian Gas Field”,

Noy., 1967 Mr. T. R. N, Lorman; “The work of the National Parks Commission

of South Australia’.

April, 1968 Mr. D. BR, Cunme: “The Industrial future of South Australia”.

May, 1968 Prof. C. M. Donacn: “Rural development in the Northern Territory”,

June, 1968 Prof. R. W. R. Ruriann: “A structural view of Continental Drilt”.

July, 1968 Mr. R. C, Spiuce: “The Role uf Research in Industry”.

Aug., 1968 Prot |. BR. M. Ravox; “Waves and tides of the South Australian

oast”.

EXHiBITs

Dr. B. G. Forses: Some recent maps issued by the Geological Survey of South

Australia.

Mr. M. J. Tyuen: Relationships between musculature and vocal sac structure m

frogs.

AWARD OF THE SIR JOSEPH VERCO MEDAL

1929 Prov, WALTER Howain, F.G.S,

1980. Jonw Mc. Brack, A-L.S.

(931 Pro, Sm Doucias Mawson, O.B.E., D.Sc., B.E., F.R.S.

1933 Pror, J. Burros Cre.anp, M.D,

1935. Pror, T, Hanvey Jownston, M.A., D.Se.

19388 Pror. J. A, Prescott, D.Sc., F.A.C..

1943 Hinnenr Womemnsey, AIS. FR,E.S.

1944 Prov, J, G. Woon, D,Sc., Ph.D.

1945 Craw T, Mamaan, M.A. BE. D.Se. F.GS.

19486 Herienr M. Hare, O.BL.

1953 L. Krrma Wann, LS,0., B.A,, B.E.. D Se.

19356 N. B. Tsnvaze, B.Sc.

1957 ~-C, S. Prrer, D.Se.

1959 GC. G. SterHens, D.Sc.

1960 H. IL. FirnAysox

1961 KR. L. Spretrr, Ph,D-

1962 IT, G. ANpnewanrHa, M.Ag.Sa.. D.de., TAA,

1963 N. H. Lupproox, M.A., Ph.D. D.G., F.G.S.

1963 RB. ¥, Sourneorr, D.Se., M.D,, B-S.. D.T.M. & I.

1966 Pror. A. R. ALpeRMaw, D.Sc., Ph.D., BGS,

1967 IL, D. Pryor, M.Sce., Dip.For.

1968 BR. C. Spnicc, M.Sc.

For stimulating and versatile contributions to the geology of South Austrilia

in the fields of sedimentation, stratigraphy, structural geology, and regional

napping. Significant in these are the discovery and description. of the

Ediacara medusoid fauna in the Pound Quartzite. formerly considered to

be Early Cumbrian but now regarded as of Precambrian ae; initiation of a

systematic. study of the Adelaide System and sediinentation in the Adelaide

Geosyncline; description of coastline migrations and geochronology of the

Quaternary of the South-East Province and of submarine canyons off the

Sonth Australian coasts; direetion of renewed interest in structures in the

northeast of South Australia, thus encouraging petroleum exploration in

the area in which gasfields were subsequently diseavered. Mr. Sprivg is the

anthor of some 62 scientific papers, 12 of which have heen published in

the Transactions.

CONTENTS

D. R. Smytu and C. M. Pumporr: A field study of the Rabbit Bandicoot,

Macrotis lagotis, Marsupialia, from Central Western Australia -

R. M. E. Wexsourn and R. T. Lance: An analysis of vegetation on

stranded coastal dunes between Robe and Naracoorte, South Australia

R. Scuoppe: Further Taxonomic notes on the species of Millotia Cassini

CASteraCeae:) > — yk EN Petey i hl ee te Mea oe

G. Beni: A new species of Ptilotus from South Australia - - -

J. A. Harris: Age structure, growth rate and spawning cycle of a popula-

tion of Yellow-eye Mullet Aldrichetta forsteri (Cuv. and Val.) from

the Coorong Lagoon, South Australia - - - - - -

M. Smyru: The distribution and life history of the Skink, Hemiergis

peronii (Fitzinger) - : ¢ : : > é - ‘

L, M. Ancex and P. M. Mawson: Helminths from some lizards mostly

from SeuthAnetraliss te ced tee Sak a tee.) ben hie

I. M. Tuomas: Two species of Saccoglossus (Enteropneusta) from South

Australia - er 3 ee eee eek E .

E. M. Campsetu: Lunettes in Southern South Australia - = aya

C. R. Twmate: Origin of Wave Rock, Hyden, Western Australia - -

Obituary: Thomas Draper Campbell IN, am Ney be eee

General Account, Library Account - - A272 ‘ : J

Research Fund - - - - : F 2 : Z 2

Report on Activities of the Council 1967-68 — - = - = F a

Officers for 1967-68 - - - - 2 : i = Se a, ES,

List of Members - - > = - = = m ~ -

List of Lecturers and Exhibits, 1967-68 - - - E - s <

Awards of the Sir Joseph Verco Medal - - - - - - .-

115