A NEW FOSSIL HOMOLID CRAB (DECAPODA, BRACHYURA), MIDDLE

TERTIARY, SOUTHEASTERN AUSTRALIA

by RICHARD J. F, JENKINS*

Summary

JENKINS, R. J, F, (1977).—A new fossil homolid crab (Decapoda, Brachyura), middle Ter-

tinry, southewstern Australia. Trans. R, Soe, 8. Aust. VLC), 1-10, 28 February, 1977.

Two new fossil decapod localities are reported in, respectively, the Oligocene and Miocene

of the Mount Gambier area, and a new species of homolid crab, Paromola pritchardi sp, nov,.

is. described from fragmentary remains collected at the older of these occurrences. The descrip-

tion of this form provides an opportunity for a review of the genus.

latroduction

In 1953 Professor M. F. Glaessner discovered

fossil decapod remains in the Gambier Lime-

stone (Sprigg 1952) near Mount Gambier,

South Australia (Fig. 1)..A second discovery of

fossil decapods jn the same formation, and also

near Mount Gambier, was made in 1955 by Dr

Mary Wade. The Gambier Limestone ts of Late

Eocene to Miocene age and occurs within the

Gambier Embayment, a deep tectonic-sedimen-

tury depression which forms the western pan

of the Otway Basin in southeastern Australia

(Ludbrook 1969), The formation reaches a

thickness in excess of 150 mj; iL consists largely

of the fragmented remains of bryozoans and

often includes abundant foraminifers (Lud-

brook 1961, 1969; Abele 1967),

The discovery made by Glaessner is in the

row of building stone quarries on sections 26,

28, 29, 30, 144 and 145, hundred of Blanche,

12 km west of Mount Gambier. Foraminiferal

assemblages collected from section 28 and

studied by McGowran (1070)! and myself sug-

gest 4 dating within the Globigerina labia-

crassata zone of Lawibrook & Lindsay (1969)

of approximately Zone P. 19/20 of Blow

(L970). With reference to Berggren (1972),

this is late Barly Oligocene. Coccolith studies

made on the same samples by Mr S, Shafic

Were also suggestive of a late Early Oligocene

or late Rupelian age (R. J, F. Jenkins 1974).

The fossil decapods from this locality are the

richest and most diverse assemblage yet known

from the Palaeogene of Australia, Either

Glaessner or I* have identified representatives of

Pagurus Fabricius Trizopagurus Forest, Munida

Leach, Dynomene Latreille, Paromela Wood-

Mason & Alcock, Ebalia Leach, Lyreidus de

Haan, Leptomithrax Miers, Tutankhamen

Rathbun, Qvalipes Rathbun, Nectocarcinus A.

Milne-Edwards, Pseudocarcinus H. Milne-

Edwards, Carcinoplax H, Milne-Edwards, and

Homoioplax Rathbun, Three other unidentified

genera are also present.

The decapod remains occur most numerously

in the interval of well bedded, coarse grained,

pink and yellow, bryozoal limestone which

immediately overlies the homogeneous, even

grained, white bryozoal limestone cut for build-

ing blocks, These limestones are part of the

“middle member” of the Gambier Limestone

{McGowran 1973).

The occurrence discovered by Dr Wade is

in the quarries on sections 601 and 606, hun-

dred of Blanche, 7 km south of Mount Gam-

bier, Foraminiferal studies made by MeGow-

ran! indicate an age within the later part of the

the Globigerina woodi woodi zone of Ludbrook

& Lindsay (1969) or the Globigerina woodi

connecta zone of D, G. Jenkins (1967); these

* Depurlment of Geology and Mineralogy, University of Adelaide, North Tce, Adelaide, S. Aust, 5000,

' McGowran, B. (1970).—Age of six samples of Gambier Limestone, Unpublished Geol, Survey Report,

Dept Mines, S, Arvest. (459), 1-8.

* Jenkins, Bo J. F. (1972) —Austrelian fossil decapod Crustacea: faunal and environmental changes.

Ph.D. thesis,

2 RICHARD J. F. JENKINS

Y KILOMETRES

BURNDA 4 <

"MURRAY.

Aaa Nt SIN

oo 200 300 400 K/LOMETALS Area oO

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1oo 200 300 MILES

MACOONNELL

ATED INT

Fig. 1, Southeastern Australia and the Mount Gambier area. The fossil decapod occurrences described

are in the quarries on the numbered sections shown on the map of the Mount Gambier area.

datings are in the vicinity of the lower part of

Zone N,6 of Blow (1969), or middle Early

Miocene. The decapod fauna is less prolific

than at the first locality; it includes representa-

tives of Axitus Leach, Paguristes Dana, Paro-

mola, Lyreidus, Ovalipes and Nectocarcinus

and an unidentified parthenopid.

The crabs mostly occur in the fine grained

bryozoal limestone at the bottom of the quar-

ries. Two fragments identified as Paromola cf.

pritchardi are from section 606, one from un-

certain level and the other in coarse grained

bryozoal limestone from probably high in the

exposure, The rocks in the quarries are part of

the upper member of the Gambier Limestone,

The repository of the fossil specimens studied

is the palaeontological collection of the South

Australian Museum (catalogue numbers pre-

fixed “P” in the text). Observations were also

made on dried specimens of the extant Paro-

mola petlerdi (Grant 1905) in the collection

of the South Australian Museum (numbers pre-

fixed "C"),

Systematics

Order Decapoda

Infraorder Brachyura

Section Dromiacea

Superfamily Homoloidea

Family Homolidae White, 1847

Type-genus: Homola Leach, 1815.

Remarks: Workers such as Ihle (1913), Gor-

don (1950), and Williamson (1965) have con-

cluded from neontological studies that the

division between the Latreilliidue (type-genus

Latreillia P, Roux, 1830) and the Homolidue

‘is less clear than previously supposed and unite

these two families. This unity is rejected by

Wright & Collins (1972) on palaeontological

grounds; they consider that one of the most

important diagnostic features of the Homolidae

Footnote added in proof: The following publication was not seen.

Serene, R. & Lohavanijaya, P, (1973)—The Brachyura (Crustacea: Decapoda) collected by the Naga

Expedition, including a review of the Homolidae. Naga Rep. 4(4), 1-187.

A NEW FOSSIL HOMOLID CRAB 3

is the presence of dorsal linede homolicae, and

indicate that these structures are absent in Let-

reillia and not present in other Cretaceous

forms winch they refer 10 the Latreilliidae,

They conclude that the Homolidae and Lat-

reilliidae “have probably been independent

stocks since Upper Jurassic times, albeit

develapme to some extent in parallel”. This

viewpoint is accepted herein,

Wright & Collins (1972, p. 31) consider that

Latreillapsir Henderson. 1888, which does have

Iltiede, is probably not allied to Larreillia, but

is a hamolid,

Genus Paromola Wood-Mason & Alcock,

1891

Type species: Dorippe cuvieri Risso, 1816, by

monotypy,

Paramola Wood-Mason & Alcock, 189L: 267:

Rathtun, 1937; 68; Bouvier, 1940; 190;

Gordon, 1950: 222; Griffin, 1965; 86 (but

not the pew species thereunder described },

Alvarez, 1968; 301.

Nomotle (Paromola) Alcock, 1899: 156; 1901:

64; Ihle, 1913: 69 fin key); Sakai, L956:

47,

Thelziope (Molofia) Barnard, 1946; 371; 1950:

341.

Diagnosis; Carapace urneshaped or sub-

rectangular, longer than broad, widest across

branchial! regions; rostrum a simple spine

Nanked on either side by a single supraorbital

spine (occasionally with small side branches)

of equal or greater size; lineae homolicae con-

spicuious, Well inside lateral margins; surface

usually granulate with scattered spinules,

spines and tubercles, smooth in one extant

species, Merus of third maxilliped elongate

with a dentate prominence or a spine near

middle of length of outer margin. inner margin

of joint usually denticuljte. Mert of pereiopods

usually spmose along their length. Palms of

chelae of ist pair of pereiopods smooth or

granulate. Extant species with 13-14 gills plus

5-6 epipods.

Remarks: tn the literature Paromola has often

been confused with two other genera of homo-

lids, Homola Leach, (815 (= Thelxiope

Rafinesque, 1814) and Carrei/lopsis Henderson,

1888. These genera all have an urn-shaped or

sub-rectangular carapace and slender, elongate

pereiopods.

In Paromela and Homola the meri of the

pereiopods are spinose. The branchial formula

of extant species of Homola is 13-14 gills +

6 epipods, similar to or little different from

that in Paromola (see Bouvier 1940, p, 191-

193; and Gordon 1950, p. 220-221), Dif-

ferences between the two are indicated in Table

The extant, New Zealand Paromola spini-

piatia Griffin, 1965, which has two prominent

spines above each orbit, a conspicuous spine on

each epigastric region, and the palms of the

chelipeds spinose, is referable to Hamola.

The genua Larreillepsis has as its type-

species the extant, Indo-West-Pacific Latreillop-

sis bixypinosee Henderson, 1888. This, and a

second living species, the Japanese Larreillop-

sty laciniata Sakat, 1936, are distinguished from

Paromola by their branchial formula of 10 gills

plus four epipods (Gordon 1950, p. 220).

However the gill structure is not preserved in

fossils. External morphological differences

between these two species and members of

Paromola ate given in Table 2,

The following previously described extant

species have been included in Paromola or

appear referable to this genus;

Paramole cuviert (Risso 1816)

Mediterranean (excluding the Adriatic) and

eastern Atantic, from Angola, Cape Bojador

and the Azores nomh to Cork, the Shetland Ts.

TABLE L

Summary af differences distinguishing Paromola Wood-Meson & Alcock, 1891, from Homola Leach, 1865.

Charter

fomols

Paromola

Supraorbylal spines

Two spines project above each orbit, a

Jaleral rostral spime near base of rostrum,

A single spine projects above cach

orbit

und 4 second spine more lateral

Rostrum

Byigastric tibercles or spines

rostral apne

Chelae of Ist pair of perciopads

Usually bidentate, less commonly single

A prontinent wbercle or spine is situated

on. each epigastric region behind lateral

Palms casually spirnoce

Invariably single

Epigastric regions usually without

conspicuous tubercles or spines

Palms cither smooth or bearing

pointed granules

4 RICHARD 5. F. JENKINS

TABLE 72

Summary of external skeletal differencex distinguishing Paromola from Latreillopsis

Choracter

Surfuce of carapace

tubercles

Width of carapace Apparently mature

swollen

Third maxilliped

Ist pair of perelopods (cheli-

peds)

Latreillopsis

Upper surface wrinkled with few distinet

individuals

across hepatig regions, which are very

Merus quadrats in shape; both merus

and ischium without spines

Merus bearing only a lerminal spine

Puromola

Usually spinose with granules and

pointed tubercles between

Except in juveniles, carapace

widest seross branchial rezions

widest

Merits elongate, with dentate pray

minence or spine about midway

along its outer margin; external

distal angle of ischium sometimes

produced to a spine; inner margin

of both joints usually denticulate

Merus spinose along length in

most species

and west coast of Norway; 150-1320 m, rarely

in shallower waters,

Paromoala prafundorum (Alcock & Anderson

1899)

Travancore coast of India: 786 m. Maldive

area; 256 m Eastern coast of Africa; 1362 m,

Paromola petierdi (Grant 1905)

Figs 31 & 4G-H.

Paromola petterdi—Gordon, 1950; 220,

Southern and southeastern Australia, from near

Grafton south to Bruny L, Tasmania, and west

to Eucla; 91-1460 m, North Island of New

Zealand from the Cavalli Is, to Banks Penin-

sula; 183-541 m. The specimen photographed,

3, C 83, is from 32 km S,W. of Cape Everard,

Victoria, ata depth of 164 m,

Paromola rathbuni Porter, 1908

Isla de Mas-Afuera, Juan Fernandez, Chile.

Latreillopsis multispinosa Uhle, 1912

Latreillopsis multispinosa The, 1912; 78, pl. 4.

figs 19-21, Kei Is,; 204 m

This species is: referable to Paramola because

of the numérous long spines on its carapace and

the form of the third maxillipeds, which have

an elongate merus with a luteral spine and a

terminal spine on the ischium, It markedly

resembles and is evidently a near relative of

Paromola acutispina (Sakai 1962) from Japan,

Paromolu japonica Parisi, 1915

[= Latreillopsis hawaiiensis Edmondson, 1932]

Japan: Tanega Shima L, Izu Peninstila, Sagami

Bay: 183-392 m. Hawaii; 55 m, According to

Sakai (1936) the species inhabits a rocky

bottom.

Paromola alcocki (Stebbing 1920)

[Apparently > Latreillopsis major Kubo, 1936;

? = Homola (Parhomola) majora Edmondson,

1951.]

Southern Africa: Algoa Bay, South Africa;

Mozambique; 73-312 m. Maldive area: 229

m. Japan |Paromola major (Kubo)]: Izu Pen-

insula, Sagami Bay, Tokyo Bay; 100-200 m-

tHawaii = [Homola (Parhumola) — majora

Edmondson}; 12-107 m, The Japanese form

inhabits a muddy bottom (Sakai 1936),

Puromola faxoni (Schmitt 1921)

Off San Diego, California: 122-370 m,. A

remarkable photograph of this species in ils

natural environment at a depth of 370 m

(Church 1971, p. 113) shows the subchelate

hind limbs holding a piece of sponge above

the back of the animal.

Paromola macrockira Sakai, 1962

Japan; Tosa Bay and Kii Peninsula,

Hamola (Moloha) acwtispinosa Sakai, 1962

Homola (Moloha) acutispinesa Sakai, 1962;

147, pl. 4 fig. 4.

Japan: Tosa Bay.

Characters of this species which indicate that

it is referable to Puromela are the single large

spine above each orbit, single rostrum,

absence of epigastric spines and the smooth

palms of the chelipeds.

One previously described fossil species can

probable be referred to Paromela;

Hamolopsis japonicus Yokoyama, 1911

fHomolopsis japonicus Yokoyama, 1911: 12, pl.

3, fig. 4.

Paleocene or Eocene: Miike Coalfield, Japan.

A NEW FOSSIL HOMOLID CRAB 5

The single median portion ot a carapace from

which this species was deseribed unfertunately

has the frontoorbital region damaged, bu

closely resembles P. priréhardi and P, petterdi

in the shupe of the other regions and in the

positioning of the major tubercles.

The modern species belonging to Paremela

can be divided into three informal species-

groups which may be characterized as follows:

|. FP. cuvier? group. Carapace more or less

covered by granules and spinules and bear-

ing short to moderately long spines ow lateral

and anterior-dorsal aspects. Palas of cheli-

peds smooth or bearing pointed granules,

particularly in mature individuals. P. cuviers,

P. petterdi, P, rathbuni, P, japonice, P.

aleoeks, P. faxoni, P, macrochira,

2. PF. prefundorwn group. Carapace mainly

smooth, with one hepatic spine and ane cer

ticle on branchial margin. Palms of chelipeds

smooth. . profundorum.

3, FP. multixpinesa group. Carapace bearing

elongate spines, between which jt is mainly

smooth, Palms of chelipeds smooth, P, mul-

tixpinasa. Po aeutiypinosu.

The early Tertiary Paromola

{ Yokoyama)

crviert group.

Paramols prifchardi sp, nov.

Figs 2, 3A-G, & 44-6

Name: Natmed after Pritehards Quarry, of sec-

tion 28, hundred of Blanche, South Australia,

Material: Seventeen incomplete specimens of

vunous pas of the carapace and four speci-

mens of isolated abdominal tergites, Holotype,

Pi5631. Median part of carapace with rostrum

and supraorbital spines lacking,

japonicus

is a fossil member of the P,

Oecnrrence: Gambier Limestone in quarries on

sections 26, 28 and 30, hundred of Blanche.

Age; Late Early Oligocene.

Description: Curapace subrectangular, gently

convex above, extremely deep in lateral axpect:

regions well marked, delimited and subdivided

by mioderately deep grooves; preater part of

sutlace covered by variably sized granules and

spinules, nine short spines on dorsum in

advance of cervical proave, lateral aspects of

varnpace Omamented by short spines and blunt

§pitules. Portion of carapace between /ineue

homoliewe 1.5 limes as long as wide, broadest

across mesobranchial regions.

Rostrum a single, forwardly directed, slender

spine about a sixth lequth of carypace, slightly

deflexed basally, smoothly upeurved distally,

Fig. 2. Paremoale pritchard! sp. nov, reconstruc-

tion of carapace; lettering indicates

repsons: QO, supraorbital, Eg, epigastric;

Pg, protegastric, 1, anteromedial Jobe, 2,

anterolateral lobe, 3, posterolateral Jobe;

H, hepatic; Mg, mesogustric; M, metagas-

tric; Eb, eprbranchial; (6, inner-branchial

lobe; U, urogastric; C, cardiac; Mb, meso-

branchial; Mt, metabranchial; I, intestinal,

Approximately x3_

Orbits forwardly directed and with wo Jarge

supraorhital spine above; inner part of supra-

orbital margin smoothly concave and with a

narrow border; supraorbital spine directed

obliquely upwards, forwards and slightly out-

wards, about twice length of rostrum, with a

lateral spinule at about half length; a short

spine on lower corner of lateral margin of orbyt,

Epigastric regions relatively small, slightly

raised, with only a few granules. Protogustric

regions each subdivided jnto three lobes by an

oblique Y-shaped groove, anteromedial lobe:

with a central spine and two spinules on

posterior part. anterolateral lobe with a single

prominent spine; posterolateral lobe with #

prominent spine on lateral aspect, a lesser spine

an inner portion, and several small spinules

between. Hepatic regions strongly inflated,

bearing a erescentic row of four acute spines

dn anterolateral aspect and a group of spinules

behind, Mesogastric region with a median spine

6 RICHARD J,

set at centre of a circlet of five, or a triangular

arrangement of three small spinules; a pair of

granulate ridges adjacent to posterior margin of

tegion. Cervical proove strongly impressed,

containing slit-like posterior gastric pits at .5)—

52 length of carapace, Metagasinc region in

form of two oblique, elongated, granulate lobes

and with a pair of more prominent granules

Positioned submedially. Urogastric region

saddle-shaped, its lateral margins marked by

two incised, crescentic grooves, Cardiac region

moderately inflated, subtriangular, with three

prominences, two side by side before and one

behind. Intestinal region depressed, progres-

sively broadened rearwards. Epibranchial

regions obliquely elongated, with a spine just

outside lineae homolicae and an irregular line

of spinules on lateral aspect. Inner-branchial

lobes on either side of Urogastric region

obliquely elongated and with one more

prominent granule. Branchiocardive groove

well marked, Mesobranchial regions gently ta-

flated, each with a line of al least three short

spines just outside finede homolicae, most

anterior of these spines the largest and situated

on a slight ridge behind fateral portion of

branchiocardiac groove. Metabranchial regions

relatively small, two-lobed, with a spinule on

posterolateral portion of Inner lobe. Sub-

branchial margin with a narrow border Paos-

terior margin fairly wide, raised, with median

third indented,

A spioule at anterolateral corner of buccal

frame,

Muscle attachment scars only faintly marked

an interior surface of cyrapace except for the

two small depressions forming the gastric apo-

demes,

Third and fifth segments of male «hdomen

each with a prominent median lobe and a4

spinuile at centre of cach of the lateral lobes,

Fifth segment subrectatgular, with posterior

angles slightly produced; median lohe

apparently bearing a few scattered granules and

with a spinule on anterior portion; lateral lobes

obliquely sulcate and apparently ornamented

by coarse granules.

& JENKINS

Fourth sezment of female abdomen with the

lateral lohes slivhtly inflated and curved down-

wards (iF abdomen was straightened behind

crab) jind the surface nearly smooth except

for a few, scatlensd weak granules; an obscure

tubercle on anterior portion of median lobe,

Measurenents: Holotype (P15631), length of

carapace excluding rostrum, 13 mm; width of

median part of carapace between lneae fronu-

lice at level of mesobranchial regions, 12 mm-

Paratype (P15632). length of carapace exclud-

ing rostrum, 23 mm; length of rostrum (incom-

plete), 3.6 mm; width of median part of cana-

ace between Iinede hornolicue at level of meso-

ranchial regions, 16 mm, The largest indi-

vidual known is represented by an incomplete

carapace (paratype P) 5638) approximately 1.3

times the size of that of ['15632,

Remmaks: The precise arringement of the

minor spinules on the carapace of P, pritehard/

is very Variable and the relief of the regions

also ¥aries slightly in different specimens.

FP. pritchard’ belongs to the PL envier

specics-group and closely resembles P. pererdé

and FP. alcecki, It seems slightly more similar

to P. perterd than to P. alcock/, but is possibly

uncestral ta both, tt differs fram P, perterai in

the more rectangular shape of Ils carapace, ts

more uplurned rostrum, und in the less pro-

nounced sculpture of the mesobrunchiul

regions. The posterior gastric pits are situated

at about .50-—52 the length of the carapace m

P. pritchardi, but at about .41—43 the length

in FP pesterdi. As well, the fossil species his a

prominent spine on the anterolateral lobe of

the protogastric regions, while only a small

lubercle ts sifuated in this position In P.

petterdé,

The fossil species differs from P. alcehi

again in its more rectangular carapace, and

if having longer supreorbital spines and many

fewer spinules present on the branchial regions.

P. alvocks has the posterior gastric pits situated

at about .A3 the length of the curapace-

P. pritchard: apparently differs from the

Eocene or Paleocene Paromeler japonicus

Fig. 3. A-G—Paremole pritchardi sp. nov. A, holotype, P15631, median par of carapace, dorsal view,

x3; B, paral

laters! part

. P15632, median part of carapace, dorsal view, x2,5; C-D, paratype, P15637,

carapace, with spinule(s) on lower corner of Lateral margin of orbit, left side, C,

dorsal view, x3, D, lateral view, x3; E-F, paratype, P1566, fragment broken from lateral part

of carupace, feft side, E, dorsal view, x3, PF, lateral view, x3; G, paratype, P15639, fragmentary

rermaius of median part of carapace with rostrum and one supraorbital spine present, view of int-

terior surface, x2,

H—Paramole ef. pritchard? PISROG, fragment of median part of curipuce, view of interior sur-

face, x24.

I—Peromela perterd? (Geant 1905), Specimen C83, d. dorsal view, x1.4.

A NEW FOSSIL HOMOLID CRAB 7

2 .

3 RICHARD J, F. JENKINS

(Yokoyama) jn lacking a distinct transverse

ridge over the anterior part of each meso-

branchial region and im bearing several

additional spines. The dilferences which occur

between FL japonicus and P. pritcherd!. and

between these fossils and the modern species,

FP. petterdi and P. alcocki, seem relatively

minor and are indicative of an extremely stow

rate Of evolitionary change. If this rate of

change is at all comparable to rates of evolu.

tionary change experienced by Paronpola prior

to the Eocene or Paleocene, then the genus

miy date from a moch carlier time, probably

fram within the Mesozoic. The fossil record of

the Homolidae extends back to the Late Juras-

sic (Glaessner 1969), None of the other fossil

venera of homolids yet described appears likely

to be the direct ancestor of Parortela.

Paromola cf. pritchardi

Figs 3H & 4F

Material; A fragment of the median part of a

carapace, P15806, and a fragment of the lateral

part of a carapace, P15805.

Qccurrence: Both specimens from Gambier

Limestone in quarry on section 606, hundred of

Blanche; P15806 from a loose piece of rock

3.5 m below the ground surface.

Age: Middle Early Miocene.

Measurements: P15806, Width of median part

of carapace between lineae homulicae, approxi-

mately 13 mm. P1S805, height of lateral part

of carapace approximately 8 mm,

Remarks; These remains are too fragmentary

to be positively identified to the level of species,

but in all features closely resemble the corfes-

ponding parts of P, pritcharsdi,

Palaeoecology

The abundance of planktonic foraminifers st

the Gambier Limestone (though fot at the

fossil decapod focality 12 kin West of Mount

Gainbier) is indicative of an open marine

environment of deposition (Ludbrook 196!).

The scarcity of terrigenous detritus in the

formation, particularly the middle member,

shows that it accumulated |A very clear waters.

Sleniler branching forms of bryozoans com-

prise the major component of the limestone

(Abele 1967), In the present day seas of

fouthern Australia, living bryozoans alound at

depths of 90-220 m, and their remains are

accumulating, as sediments over wide arcas of

the continental shell and the upper part of the

continental slope (Conolly & von dev Borel

1967, Wass, Conolly & Maclotyre 1970),

Many of these sediments strikingly resemble

those composing (he Gambier Limestone in the

kinds of bryezaans present, the proportions of

foraminifers and other skeletal remains (such

as Molluses, echinoids, and serpulid worms),

their degree of sorting, and in the associated

sedimentary structiires (ctulrrent ripples and

mounds). It seems reasonable to suppose that

the Gambier Limestone is a fossil equivalent

nf these deposits and that Wt formed in waters

of comparyble depth. (approximately 90 to 220

m), This View contrasts la some degree with

that of Abele (1967) who noted that the shape

ul certain foraminifers present in the forma-

tion js characteristic of forms Which live

attached to seaweed, He inferred that deposi-

tian occurred between approximately 4h and

100 rm depth.

The range of depth of deposition suggested

aboye overlaps the depth range in which the

extant species of Petramole are most frequently

recorded, between approximately 100 and 500

Mi. Thus P. pritchardi probably lived at similar

depths as modern meinbers of the venus. The

living species to which the other fossil decapods

known from the Gambier Limestone are most

closely allied, also occur typically on the outer

part of the continental shelf or on the upper

part of the continental slope. Photographs of

the sen bottom in the wreas just mentioned

(Conolly & von der Borch 1967; Wass, Conolly

& Macintyre, (970) show living bryozoans

cocurring in forests and associated with

spatiges, ur itore sparsely distributed on open

areas of sediment, Peremoly seers well

adapted to live In bryozoan forests, its long

Fig. 4. A-E—Puremoala pritchardi sp. nov. A, paratype, P15643, hepatic region of carapace, sight side,

with # spinule(s) on the part of the murgin corresponding to the afMterolateral corner of the buc-

cal frame, lateral view, x2.5; B, paratype, P| 5635, incomplete Jaleral part of carapace, right side,

lateral view, x3; C, paratype 3 P1564, tergite of third segment of abdomen, view af interior

surface (fOp anterior), x6; D, paratype d, P15641, tergite of fifth sewnent of abdomen, view of

interior surface (Lop anterior), x3;

external surface (top anterlor), x2.9.

paratype 9, P15634, tergile of fourth segment of abdomen,

F—Paraniola cf. pritchardi PAS805, fragment of lateral part of carapace, left side, lateral view,

x35.

G-H—Paramole petterdl (arant, (905). Specimen C8], 2. G. anterior-ventral aspect, x6, H. dor-

sal View, X2.5,

fe)

jaa)

10 RICHARD J. F. JENKINS

legs and high stance (Church 1971: fig. on p.

113) probably enabling it to step over the

bryozoans.

Acknowledgments

The writer is indebted to Professor M. F.

Glaessner, University of Adelaide, and Dr

pod occurrences which they discovered; Pro-

fessor Glaessner is also thanked for his con-

structive criticism of the manuscript. Dr B.

McGowran, University of Adelaide, kindly

examined and provided age data on various

foraminiferal samples.

Mary Wade, Queensland Museum, for This research was supported by an Australian

encouraging the study of several fossil deca~ Commonwealth Postgraduate Award.

References

ABELE, C. (1967).—Bryozoal sedimentation: IxHie, J. E. W. (1913).—Die Decapoda Brachyura

Gambier Limestone. In: J. McAndrew, der Siboga—Expedition. I. Dromiacea. Siboga

M. A. H. Marsden and B. Marshall, eds, Sum-

mary papers for section C—Geology, 39th

A.N.Z.A.A.S. Congress, Melbourne, January

1967, A7-A8.

Atcock, A. (1899).—An account of the deep-sea

Brachyura collected by the Royal Indian

marine survey ship “Investigator”. Calcutta.

ALoocx, A. (1901).—Materials for a carcino-

logical fauna of India, No. 5, The Brachyura

Primigeria or Dromiacea. J. Asiat. Soc. Beng.

68, 3(3), 123-169.

ALVAREZ, R. Z, (1968).—Crustaceous decApodos

Ibéricos. Investigacién presq., Barcelona.

BARNARD, K. H. (1946).—Descriptions of new

species of South African Decapod Crustacea,

with notes on synonomy and new records.

Ann. Mag. nat. Hist. (11) 13, 361-392.

BARNARD, K. H. (1950).—Descriptive catalogue of

South African decapod Crustacea. Ann. S.

Afr. Mus. 38, 1-837.

BERGGREN, W. A. (1972).—A Cenozoic time-

scale—some implications for regional geology

and paleobiogeography. Lethaia 5, 195-215.

Biow, W. H. (1970).—Validity of biostratigraphic

correlations based on the Globigerinacea.

Micropaleontology 16(3), 257-268.

Bouvier, E. L. (1940).—Décapodes marcheurs.

Faune Fr. 37. (Paul Lechevalier: Paris.)

Cuurcu, R. (1971).—Deepstar explores the ocean

floor. Natn. geogr. Mag. 139(1), 110-129.

Cono_ty, J. R. & VON DER Borcu, C. C. (1967) —

Sedimentation and physiography of the sea

floor south of Australia. Sediment. Geol. 1(1),

181-220.

Epmonpson, C. H. (1932).—A giant Latreillopsis

from Hawaii. Occ. Pap. Bernice P. Bishop

Mus. 9(24), 3-9.

GLAESSNER, M. F. (1969).—Decapoda. Jn R. C.

Moore, ed., Treatise on invertebrate paleon-

tology (R) Arthropoda 4 (2), R399-R533,

AS552-R566, addendum R626-R628. Univer-

sity of Kansas & Geological Society of

America.

Gorpon, I. (1950).—Crustacea Dromiacea. I.

Systematic account of the Dromiacea collected

by the “John Murray” expedition. II. The

morphology of the spermatheca in certain

Dromiacea. Scient. Rep. John Murray Exped.

9, 201-235, text-figs 1-26, pl. 1.

GriFFIN, D. J. G. (1965).—A new species of

Paromola (Crustacea, Decapoda, Thelxio-

pidae from New Zealand. Trans. R. Soc. N.Z.

7(4), 85-91.

Exped. 39b (71), 1-96, 1-4.

Jenkins, D. G. (1967).—Planktonic foraminiferal

zones and new taxa from the Lower Miocene

to Pleistocene of New Zealand. N.Z. J. Geol.

Geophys. 10(4), 1064-1078.

Jenkins, R. J, F. (1974).—A new giant penguin

from the Eocene of Australia. Palaeontology

17(2), 291-310.

LupsBroox, N. H. (1961).—Stratigraphy of the

Murray Basin in South Australia. Bull. geol.

Surv. S. Aust. 36, 1-96.

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L. W. Parkin, ed., Handbook of South Aus-

tralian geology. Geol. Surv. S. Aust. chapter

5, 172-203, figs 83-101.

Lupsroox, N. H. & Linpsay, J. M. (1969).—

Tertiary foraminiferal zones in South Aus-

tralia. Jn: P, Brénnimann and H. H. Renz,

eds, Proceedings of the first international con-

ference on planktonic microfossils, Geneva,

1967, 2, 366-374. (Brill: Leiden.)

McGowran, B. (1973)—Observation bore No. 2,

Gambier Embayment of the Otway Basin:

Tertiary micropalaeontology and stratigraphy.

Mineral Resour. Rev., S. Aust. 135, 43-55.

RATHBUN, M. J. (1937).—The Oxystomatous and

allied crabs of America. Bull. U.S. natn, Mus.

166, 1-278

Saka, T. (1936).—Studies on the crabs of Japan.

I. Dromiacea, Sci. Rep. Tokyo Bunrika Diag.

(B) 3, Supp. |, 1-66.

Spricc, R, C. (1952).—The Geology of the South-

East province, South Australia, with special

reference to Quaternary coast-line migrations

and modern beach developments. Bull. geol.

Surv. S. Aust. 29, 1-120.

Wass, R. E., Conoity, J. R. & Macintyre, R. J.

(1970).—Bryozoan carbonate sand con-

tinuous along southern Australia. Marine

Geol. 9(1), 63-73.

Wixuiamson, D. I. (1965).—Some larval stages

of three Australian crabs belonging to the

families Homolidae and Raninidae, and obser-

vations on the affinities of these families

(Crustacea: Decapoda). Aust. J. mar. Fresh-

wat, Res. 16, 369-397.

Woopmason, J. & Atcock, A. (1891).—Natural

history notes from H. M. Indian Marine Sur-

vey steamer “Investigator”, Commander R. F.

Hoskyn, R.N., commanding, No. 21. Note on

the results of the least season’s deep-sea

dredging, Ann. Mag. nat. Hist. (6)7, 1-19,

186-202, 258-272.

WricHT, C. W. & Coxruins, J. S. H. (1972).—

British Cretaceous crabs. Palaeontogr. Soc.

[Monogr.], 114 pp.

ASTRAEUS (COLEOPTRA: BUPRESTIDAE): A DESCRIPTION OF THREE

NEW SPECIES AND NEW LOCALITY RECORDS

BY S. BARKER

Summary

Three new species of the Australian buprestid genus Astraeus are described, and new data on the

distribution of several species are presented.

ASTRAEUS (COLEOPTERA: BUPRESTIDAE): A DESCRIFTION OF THREE

NEW SPECIES AND NEW LOCALITY RECORDS

by S. BARKER*

Summary

Barxer, 8S. (1977).—Astraeus (Coleoptera: Buprestidac): a description of three new species

and new locality records. Trans, R, Soe, §, Aust, WI(1), LI-14, 28 February, 1977,

Three new species of the Australian buprestid genus Asirdeus are described, and new data

un the distribution of several species are presented.

Introduction

Following my review of the genus As/raeus

(Barker 1975) more material is available

including three undescribed species and several

range extensions. I have also referred to the

work of Cobos (1955, 1973) who placed

Astracus close to the newly defined tribe

ACHERUSINI Cobos, which ocurs in South

America, whereas | followed Carter (1929) in

placing Astrdeus in the BUPRESTINI,

As yet no information is available in the

literature on the life history of Astraeus,

because apart from the knowledge that the

larvae are trunk and root borers and that many

feed on Casuarine species, little is known of

the biology of Astraeus,

All specimens referred to are lodged in the

collection of the South Australian Museum,

New Locality Records

Astracus (Depollus) irregularis van de Poll

3¢& 19 19 km E of Kimba, S, Aust. on

Casuarina helmsi, 8. Barker, 291.1976.

This is the first record of the species and sub-

genus outside Western Australia to which |

stated it was confined (Barker 1975, p. 107).

Astracus (Astraeus) obscurus Barker

4a &1919 km E of Kimba, 8. Aust. on

Casuarina helmsi, 8. Barker, (2.xi1,.1975; 3 &

& 2 9 Pindar Mill, Tallering Stn, Pindar,

W.A. on Casuarina dielsiana, S. Barker,

27.x1i.1975.

This extends the range by approx. 500 km N

and 1,300 km E.

Astraeus (Astraens) smythi Barker

53 & 2% 4 km west of Glasshouse Moun-

tains, Qld, on Casuarina littoralis, 8, Barker,

5.xi, 1975,

Astraeus (Astraeus) watsoni Barker

PIG. 4A

1 do, Badjaling, W-A. on Casuarina huege-

liana, S, Barker, 20.xii.1975.

Size. Male 10.9 x 3.9 mm.

General remarks. A. watsent was described

from female specimens. A male is now avail-

able and its genitalia is figured below, On the

basis of external features and shape of male

genitalia this species shows closest allinity with

A. macimillani, A, carnabyi, A, badeni, A, jan-

soni, A, oberthuri and A, carteri,

New Species

Astraeus (Astraens) mourangeensis sp. nov.

FIGS |, 4B

Types.

Holotype: ¢& 4 km SW Mourangee Stn, Edun-

galba, Qld on Cuswarinu sp., B.E. & 8. Adams,

26,xii.1974, SAM FT 20 985.

Allotype: ¢ 4 km SW Moitirangee Stn, Edun-

galba, Qld on Casuarina sp, EE. & 8. Adams,

26.xi1.1974. SAM 1 20 986.

Colour,

Male. Head blue-green at the apex, dark blue

ut the base and sides with golden reflections.

Antennae blue-green with golden reflections,

Pronotum wrquoise at the sides with golden

reflections, dark blue in the middle with purple

reflections. Elytra black with blue reflections,

each elytron with the following yellow mark-

ings: a basal spot; a clubbed-shaped fascia

before the middle covering the humeral fold

and running towards the suture but not touch-

* Department of Zoology, University of Adelaide, North Tce. Adelaide, S. Aust. 5000,

12 S. BARKER

5 mm

Fig. 1. Astraeus mourangeensis sp. nov.

ing it; a fascia after the middle touching the

margin but not reaching the suture; a preapical

spot. Undersurface dark metallic blue with blue

reflections; Ist tarsal segment testaceous with

dark brown tips; 2nd, 3rd and 4th tarsal seg-

ments dark brown with metallic blue reflections

on the supper surface; hairs silver.

Female. Head black with blue and purple

reflections, Antennae black with purple reflec-

tions. Pronotum black with purple reflections

in the middle, blue and purple reflections at

the sides. Elytra as in the male except that the

first fascia is broken to form two spots. Under-

surface black with purple reflections. Legs:

femur and tibia black with purple reflections;

tarsi as in the male; hairs silver.

Shape and sculpture. Head with even, shallow

punctures; a thin median keel; hairy. Pronotum

with shallow even punctures; with a thin

median longitudinal glabrous line which

extends forwards from the basal crypt but does

not reach the middle, from its end a wide

shallow depression extends forwards almost to

the anterior margin; rounded at the sides from

the base, tapered before the middle to the apex;

apical edge rounded and projecting forwards

in the middle; hairy. Elytra costate, the inter-

vals flat; slightly concave at the sides from the

base to before the middle then rounded and

tapered to the marginal spine which is sharp

but barely curved; humeral fold well developed,

angled. Undersurface evenly punctured, the

punctures shallow in the middle, slightly larger

and deeper at the sides; sparsely haired. Over-

all the body shape is long and narrow.

Size. Male 6.6 x 2.7 mm.

Female 8.0 x 2.9 mm.

Distribution. Queensland.

General remarks. On the basis of external

morphology and shape of male genitalia this

species shows closest affinity with A. mastersi

and A. samouelli. A. mourangeensis differs

from these species; in outline being compara-

tively narrower; in colour of the head and pro-

notum; in that only part of the first tarsal seg-

ment is testaceous.

Specimens examined. Types only.

Astraeus (Astraeus) blackdownsensis sp. nov.

FIG. 2

Type.

Holotype: ? Blackdowns Tableland Forestry

Reserve, Qld on Casuarina inophloia,

3.xi.1975, E.E. Adams & S. Barker. SAM 1

20 987.

Colour.

Female. Head blue-green. Antennae black with

green and blue reflections. Pronotum with a

heart-shaped dark purple mark in the centre,

green at the anterior margins, blue laterally.

Elytra black with the following yellow mark-

ings on each elytron: a basal spot; a spot before

the middle near the suture but not touching it;

a fascia after the middle not touching the

suture or lateral margin; a small preapical spot:

on the margin a spot near the shoulder cover-

ing the humeral fold, midway between this

spot and the fascia there is a laterally elongate

mark commencing at the margin and extending

for four intervals between the costae. Under-

surface: prosternum metallic blue-green,

remaining undersurface and legs metallic blue;

hairs silver.

Shape and sculpture. Head uniformly and

shallowly punctured; with a median longi-

tudinal keel; hairy. Pronotum with shallow

punctures in the middle, deeper and closer at

the sides; a broad median longitudinal

impressed line clearly defined at the apex run-

THREE NEW SPECIES OF ASTRAELS 13

5mm

Fig. 2. Astracus hlackdownsensis sp. nov,

ning 4 of the distance to the base; at the sides

rounded and tapered from base to apex; hairy.

Elytra costate, the intervals flat but slightly

wrinkled; sides parallel to before the middle

then gently rounded and tapered to the apex;

marginal and sutural spines sharp and

moderately Well-developed; humeral fold

moderately developed and angled. Undersur-

fuce evenly, shallowly but closely punctured;

hairy.

Size. Female 10,1 x 3.8 mm,

Distribution. Queensland,

General remarks. The elytral pattern is similar

to A, watseni but the species differs being

smaller, the elytral intervals are wrinkled and

the body is comparatively hairless, It also

differs in colour,

Specimens examined, Type only.

Astraeus (Astraeus) crockeri sp, nov,

FIGS 3, 4C

Types.

Holotype: of Juranda rockhole 106 km § Balla-

donia, W.A, on Callitris preissti, 15.xi1,1975,

S. Barker. SAM I 20 988.

Allotype: @ Juranda rockhole 106 km § Balla-

donia, W.A. on Callitris preissii, 9.xti,1974, S.

Barker, SAM T 20 989,

Paratypes: 5 ¢ & 5 9 Juranda rockhole 106 km

S Balladonia, W.A, on Callitris preitssii,

9.xi1,1974 & 15,.x11.1975, S. Barker.

Colour. Head and pronotum metallic purple

Antennae black with blue reflections. Elytra

black with purple reflections, cach elytron with

the following yellow markings: a basal spot:

before the middle a fascia covering the humeral

fold, concave towards the base and rumning

close to the suture but not touching it; after

the middle a fascia commencing on the margin

concave towards the apex running towards the

suture but not reaching it, Undersurface purple.

Legs with blue and purple refiections, Hairs

silver,

5mm

Fis. 3. Astracus crocker! sp. nav

l4 5. BARKER

A B

LIimm

Fig. 4. Outline diagram of dorsal surface of para-

meres of male Astraeus (Astracus) species

A: A. watsoni; B: A. mourangeensis; C:

A. crockeri.

Shape and sculpture. Head with shallow even

punctures, no median keel, slightly excavated

between the eyes, hairy. Pronotum with shallow

even punctures, basal crypt very elongate and

with a short impressed line projecting forwards

from it and continuous with the basal end of

a median Jongitudinal glabrous line which does

not reach the anterior margin: at the sides

rounded and tapered from base to apex; hairy.

Elytra costate, the intervals flat; more or less

parallel-sided from the base to the middle then

rounded and tapered to the strongly developed

marginal spine, sutural spine well developed;

humeral fold moderately developed and angled.

Undersurface shallowly but evenly punctured;

hairy, Overall the body shape is rounded when

seen from above.

Size, Males 6,4 = 0,2 x 2.8 © 0.1 mm (6),

Females 6.8 = 0.1 x 2.9 + 0.1 mm (6).

Distribution. Southeast Western Australia.

General remarks. On external characters this

species shows closest affinity with A. badeni,

although male genitalia are similar in shape to

those of A. fraseriensis, It differs from A.

hadeni in being smaller in size, the intervals

between the costae on the elytra are smooth

hence the elytra are shinier than in A. badeni

in which the costal intervals are wrinkled. Also

the humeral fold is better developed as are the

marginal spines. The colour also differs. In the

specimens of A. crockeri | have examined, none

has q preapicul spot on each elytron although

in A. badeni this is a variable character. Named

after Mrs A. BE, Crocker,

With the addition of the above new species

the key to Astraeus (sensu stricto) (Barker

1975, pp. 114, 115) needs the following modi-

fications. Add under the appropriate couplet—

6. Only ist tarsal segment testaceous

mourangeensis sp. nov.

11. Head green, pronoltum with heart-shaped

purple mark in centre, green in front, blue at

sides — _ blackdownsensis sp. nov.

21. Humeral fold ipdorapely developed and

angled* 4 _ crockeri sp. noy.

* See Barker (1975) Fig. 1.

Acknowledgments

I am indebted: to Mr B. Levey for advice;

to Mr G, F. Gross and Dr E. Matthews, South

Australian Museum for advice and assistance;

to Mrs A. E. Crocker and family, Balladonia

Sin, W.A. and Mr E, EB. Adams and family of

Mourangee Stn, Qld for hospitality and assis-

tance; to Dr E, Wollaston, Department of

Botany, University of Adelaide, for identitica-

tion of plant specimens. I was in receipt of a

grant from the Interim Council of the Aus-

tralian Biological Resources Study,

References

Bareer, S, €1975)—Revision of the genus

Astraens LaPorte & Gory (Coleoptera: Bup-

restidae). Trans. R, Soc. S. Aust. 99, 105-142,

Carter, H, J, (1929}).—A check list of the Aus-

tralian Buprestidae Aust. Zool, 5, 265-304,

Coros, A. (1955),—Hstudio sobre los Ptosionites

de CH. Kerrernans (Coleoptera, Buprestidae).

Bull, fnst, R. Soe. Nat. Belg, 31, 1-

Copos, A. (1973).—Revisién del género Tylau-

chenia Burm., y afines (Coleoptera, Bupres-

fitiae), Archiv. Inst. Aclimatacidn 18, 147-

173.

EUCALYPTUS CYANOPHYLLA, A NEW SPECIES FROM SOUTH

AUSTRALIA AND VICTORIA

BY M. I. H. BROOKER

Summary

A new species of mallee eucalypt, belonging to series DUMOSAE, and distributed in the Murray

Mallee region of South Australia and northwestern Victoria, is described and figured, and its

affinities discussed.

KLUCALYPTUS CYANOPHYLLA, A NEW SPECIES FROM SOUTH AUSTRALIA

AND VICTORIA

by M, I. H. Brooker*

SUMMARY

Beoower, M. 1, H. (1977) —Eucalyptus cyanophylla, a new species from South Australia and

Victoria, Trans, A, Soe, §, Aust, WIL),

15-18, 28 February, 1977.

A new species of mallee eucalypt, belonging to series DUMOSAE, and distributed in

the Murray Mallee region of South Australia and northwestern Victoria, is described and

figured, and its affinities discussed.

Description

Eucalyptus cyanophylly Brooker sp.

(SLE:H).1 Figs 1-3.

Eucalypio dumoso A.Curm, ex Schau, affinis a.

qua foliis latioribus et schistacioribus, alabastils et

fructibus grossioribus, et florescentia differt.

Frutex “mallee” ad 6 m_altus, cortice basin

versus fibroso, supra laevi. Glandulae oleosae in

medulla. Lignotuberum formans. Cotyledones reni-

formes, Folia plantulae petiolata, lanceolata vel

ovala, 1-6 x 04-3 em, Folia juvenilia petiolata,

ovata, atroviridia, 7-12 x 4—7 cm, Folia adulta

petioluta, lato-lanceolata, schistucea vel glauca,

10-16 x 2-3 com. Inflorescentiae axillares 7(11)-

florae. Pedunculi crassi, 05-12 em longi,

Alabastra breviter pedicellata vel subsessilia, 0,8—

13 x O.5-0.8 om. Hypanthiurm cylindricum vel

obconicum, rugosum vel costatum. Operculurm

hemisphacricum Vel turbinatum, costatum. Fila-

menta in alabastro primum erecta demum inflexa,

Antherae oblongae vel obovatae, sub-basifixae, ver-

satiles, in rimis longitudinalibus dehiscentes,

Locul: 4-5, Ovula verticaliter eae Fructus

cylindrica vel obconica, 0.7-! x 0.7-0.9 cm,

rugosa vel costata. Discus declivis, Walvae non-

exsertuc. Semina rufa, nituntia, reticulo non-pro-

fundo,

Holotypus ca, 3 km northeast of Berri, South

Australia (34°)5'S, 140°37'R) B. Copley 3799,

5.viii. 1972 (ABD), Isotypus: FRI,

A species with affinity to E, dyumosa A

Cunn, ex Schau, but differing in the broader,

bluer leaves, coarser buds and fruits, and in the

flowering time (late summer-autumn for £.

dumosa; Winter-spring for EB. cyanophylla),

A mallee to 6 m tall with grey-brown, fibrous

bark towards the base, smooth above. Oil

glaids in pith, Capable of forming lignotubers,

Ceryledons reniform, Seedling leaves petio-

late, lanceolate to ovate, 1-6 x 04-3 cm.

nov.

Juvenile leaves petiolate, ovate, dark green, 7-

12 x 4-7 cm, Adult leaves petiolate, broad-

lanceolate, bluish grey (o glaucous, 10-16 x 2-3

cm,

Inflorescences axillary of 711) buds,

Peduncles thick, 0.5-1.2 em long. Buds shortly

pedicellate to subsessile, O.8—1.3 x 0,5-0.8 em,

Hypanthium cylindrical or obeonical, rugose or

ribbed, Operculum hemispherical or turbinate,

ribbed, Filaments in bud at first erect then

inflected. Anthers oblong to obovate, sub-

basitixed, versatile, opening in Jongitudinal slits,

Locules 4—5, Ovules in 4 vertical rows,

Fruit cylindrical or obconical, 0,7—1 x O.7=

0.9 cm, rugose or ribbed. Dise sloping inwards,

Valves to rim level,

Seed red-brown, |ustrous, with a shallow

reticulum.

Collections examined: South Australia—Overland

Corner, 2.xii.1913, J. B. Cleland (NSW): Ala-

woona, Dec. 1913, J, B, Cleland (AD); Morgan-

Renmark, July 1914, W. Gill (NSW); Berri, Jan,

1921, J, B, Cleland (AD); Mindarie, S.iv.1947,

©, D, Boomsma (Woods & Forests Dept Adelaide,

FRI): Berri, 20.%.1962, B, Dangerfield (AD);

udjucent to northwest corner of Berri Irrigation

Area, 11.x%.1965, D. E. Symon 3789 (NSW);

Winkie, Dalziel Rd, 23.xii.1967, B. Copley 1672

(AD, FRI); Renmark-Berri, 18.viii 1968, L A. S,

Johnson (NSW 47194); 20 km from Barmera

towards Overland Comer, 27.viii 1968, M, E, Phil-

bps (NSW); | km N of Alawoona, 6.Vi.1970,

. Anderson (AD): Overland Corner,

Th vi.1970, A, G. Spooner (AD); Berri, July 1970,

F. van der Sommen (FRI); 25.9 koi E of Waikerie

on road to Kingston, 3.iv,1975, M. I. H. Brooker

4905 (FRI, AD, MEL, NSW, PERTH); 24 km

yaw Karoonda, 24.vii.1975, F. van der Sommen

tf ).

+ CSIRO Division of Forest Research. Banks St. Yarralumls. A.C.T. 2600.

' Code derived from Pryor & lohnsan (1971).

16 M. I. H. BROOKER

Vig. 1. Eucalyptus cyanophylla sp. nov. A—Seedling, x .9. B, C—Buds and fruits from type, x .9. D, E

Buds and fruit from Berri, July 1970, F. van der Sommen, x .9. F—Adult leaves, x .6.

EUCALYPTUS CY ANOPHYLLA NEW SPECIES 7

Fig. 2. Eycalypiuy cyanophylla sp. nov. A—Anthers, x 15. B—Bud section, x 3. C—

Ovules, x 14, D—Seed(s) and chaff (c), x 15.

j38°

@ Distrihution of E. cyanophytla

Fig. 3. Distribution of Eucalyptus ¢yanophylla sp,

nov,

Victoria—6.4 km § of Meringur, Apr. 1971, H.

Gorge (NSW); ca. 2 km E of state border

between Renmark and Mildura, 3.iv.1975, M. I. H.

Brooker 4904 (FRI, MEL, AD, NSW, K).

Discussion

Eucalyptus cvanephylla is the species of the

Murtay Mallee of South Australia and of

northwestern Victoria (Fig. 3) which has been

incorrectly referred to as EF. pileata Blakely

by Burbidge (1947), Black (1952), and Willis

(1972). The type of &, pileata is from Des-

mond, Western Australia and the typical form

extends eastwards as far as southern and

western Eyre Peninsula in South Australia.

Related forms on Yorke Peninsula should be

interpreted as intergrades between E. pileata

to the west and E. dumosa and E. anceps

(Maid.) Blakely to the east. &. pileata differs

strikingly from E£. cyanophylla in the narrower,

bright green, glossy leaves which are often held

more or less erect on the branchiets.

Both &. pileata and E. cyanophylla belong in

the series DUMOSAE (Pryor & Johnson 1971)

which is segregated in the proposed subgenus

Symphyomyrius by the association of several

constant features, viz., reniform cotyledons;

strict inflection of the staminal filaments each

of which bears fertile, versatile anthers; glandu-

lar pith; placentae with four vertical rows of

oyvular structures; and lustrous, reddish seed

with a shallow reticulum. These characters have

been discussed by Carr & Carr (1969) and

Brooker (1971, 1972).

The species name is given for the con-

spicuously blue-grey leaves which distinguish it

from E. dumosa and other mallees growing

18 M. I. H. BROOKER

near or within its area of distribution. Willis

(1972) commented on the “silver aspect” of

the trees so noticeable in the field.

My observations agree with those of Mr

C. D. Boomsma (pers. comm.) that E. dumosa

does not ocur within the area of distribution

although it overlaps along the margins as might

be expected.

Key

A revision of part of Black’s Key (1952, pp.

616-617) for the South Australian species of

the section DUMARIA (Pryor & Johnson

1971) is as follows:

CC.1 Leaves alternate, thick and stiff.

DD. Seed reddish-brown, lustrous with a shallow

reticulum.

EE. Buds and fruit quite sessile, hypanthium

not wholly tapering if at all . E. conglobata

EE. Buds and fruit subsessile to pedicellate,

hypanthium tapering.

FF. Leaves green, more or less glossy.

GG, Buds and fruit more or less sessile, oper-

culum conical to hemispherical, smooth

or .Tibbed nates teeter E. anceps

GG. buds and fruit pedicellate, operculum

beaked or hemispherical, usually pro-

minently ribbed ...........0....00..... E. pileata

FF. Leaves grey-green or bluish gray, dull.

HH. Leaves grey-green, 1-2 cm wide, Flower-

ing in late sumer and autumn ................

E. dumosa

HH. Leaves bluish grey 2—3 cm wide. Flower-

ing in winter and spring ..........00.0...00.....

E. cyanophylla

DD. Seed grey and deeply pitted; or blackish-

grey, more or less smooth on the dorsal side

and with sharp ribs on the ventral side.

Il. Seed grey and deeply pitted.

JJ. Operculum flattened-hemispherical, as wide

or wider than the hypanthium, hypanthium

not conspicuously ribbed ........ E. concinna

Jj. Operculum obtusely conical or hemispheri-

cal and pointed, rarely as wide as, usually

narrower than the hypanthium, hypanthium

obscurely or conspicuously ribbed.

KK. Fruit small, to 0.5 cm diameter, leaves

narrow lanceolate .......... E. brachycalyx2

KK. Fruit larger, more than 0.6 cm diameter,

leaves lanceolate .........-...0:n..06 E. rugosa

II. Seed blackish-grey, more or less smooth on

the dorsal side and with sharp ribs on the

ventral side.

LL. Fruit barrel-shaped, 1.5-2 cm long, pedi-

cellate, smooth or with shallow ribbing;

fruiting peduncle reflexed or rarely erect .

E. pimpiniana

LL: Fruit cylindrical or urceolate, 1-2 cm

long, pedicellate or sessile, smooth,

shallowly or coarsely ribbed; fruiting

peduncle reflexed or erect.

MM. Fruit pedicellate, cylindrical or urceo-

late, 1-15 cm long, smooth or

shallowly ribbed; fruiting peduncle

erect or reflexed ............,. E. incrassata?

MM. Fruit pedicellate or sessile, cylindrical,

1.5-2 cm _ long, ribbed;

coarsely

peduncle erect .. d .. E, angulosa

CC.1! Leaves Pepatts, ‘Slagtons, hasigs: connate at

Le : . E. gamophylla

1CC only, as in Black (1952).

2E. brachycalyx and E. rugosa are intergrading

species. E. rugosa is usually coastal and is more

robust in leaves, buds and fruit.

8. incrassata and E. angulosa are intergrading

species. E. angulosa is usually coastal and is more

robust in leaves, buds and fruit.

Acknowledgments

I wish to thank Mr C. D. Boomsma for his

encouragement and assistance in the prepara-

tion of this paper, and Mr G. Moss for the

drawings and map.

References

Back, J. M. (1952).—‘‘Flora of South Australia.”

Part 3, 523-683 (Govt Printer: Adelaide).

Brooker, M. I. H. (1971).—Studies in the genus

Eucalyptus, Series Dumosae. Nuytsia 1, 210-

216.

Brooker, M. I. H. (1972).—Four new taxa of

Eucalyptus from Western Australia. Nuytsia

1, 242-253.

BursBipGe, N. T. 1947).—Key to the South Aus-

tralian species of Eucalyptus, Trans. R. Soc,

S. Aust. 71, 137-163.

Carr, S. G. M. & Carr, D. J. (1969).—Oil glands

and ducts in Eucalyptus L’Herit. I. The

phloem and the pith. Aust. J. Bot. 17, 473-

513.

Pryor, L. D. & JOHNSON, L. A. S. (1971).—“A

Classification of the Eucalypts.” (Aust. Nat.

Univ.: Canberra.)

Wits, J. H. (1972).—“A Handbook to Plants in

Victoria.” (Melb. Univ. Press: Melbourne.)

THE GENUS CYCLOSTRONGYLUS JOHNSTON & MAWSON

(NEMATODA: TRICHONEMATIDAE)

BY PATRICIA M. MAAWSON

Summary

Cyclostrongylus is redefined. The type species of the genus Oesophagonastes (O. gallardi), is a

synonym of C. wallabiae, so this genus falls and its species are transferred to Cyclostrongylus.

Cyclostrongylus spp. Considered valid are C. wallabiae (type sp.), C. gallardi, C. kartana (s. O.

kartana), C. leptos (s. O. leptos) and C. parma (s. O. parma). Of the three other species formerly

assigned to Cyclostrongylus, C. clelandi belongs to another genus, C. dissimilus belongs to

Macropostrongyloides, and C. medioannulatus (of which no specimen can be found) is regarded as

sp. inq.

THE GENUS CYCLOSTRONGYLUS JOHNSTON & MAWSON

(NEMATODA: TRICHONEMATIDAE)

by Patricia M, Mawson*

Summary

Mawson, P, M. (1977).—The genus Cyclostrongyluy Johnston & Mawson (Nematoda; Tricho-

nematidae). Trans. R. Sue. 8S. Anse, W1(1), 19-20, 28 February, 1977.

Cyclostrongyluy is redefined, The type species of the genus Oeswphagonastes (QO. vallurdi),

is a synonym of C. wallabiae, so this genus falls and its species are transferred to Cyelo-

stroneylus. Cyclostrangylus spp. considered valid are C. wallabiae (type sp.), C. gallardi, C.

kartana (s. O. kartana), C. leptoy (s. O. leptos) and C. parma (s. O. parma). Of the three

other species formerly assigned to Cyelostrongylus, C. clelandi belongs to another genus, C,

dissimilis belongs to Macrapastrongyloides, and C, medieannulatus (of which no specimen can

be found) is regarded as sp. ing.

Introdoction

The genus Cyclostroneyvlus Was erected in

1939 to include four species, C. wallabiae, C.

clelandi, GC. gallardi and, doubtfully, C, dis-

similis. C. medioannulatux was added by John-

ston & Mawson (1940). These species were

similar in having a cuticular collar waround the

anteriot end, a deep buccal cavity and, in the

first three, an oesophagus of distinctive shape.

The Walls of the buccal cavily showed different

degrees of thickening.

The types of these species, and fresh material

of C. dissimilix and C. clelandi, have been

examined, and it is now possible to clarify the

position of the genus.

Results

The most striking fact emerging from this

study is that the type species of Cyclostrongylus

(C. wallabiae) is identical with the type species

of Cesephagonastes, O, gallardi (Johnston &

Mawson 1942), described from the same host

Species and from a relatively close locality in

New South Wales, Qesophagonastes, now

becomes a synonym of Cyclostronyylus, and the

species assigned to Oesophagonastes must be

transferred to Cyclostrongylus,

Some of the species originally placed in

Cyelostroneyluy vary considerably from the

type, and are not now gonsidered as valid

species of the genus. These are:

C, clelandi in which the shape of the

oesophagus and of the cephalic papillae are

quite different, and in which the walls of the

buccal cavity do not appear to be sclerotized

at all. A new genus will be necessary for this

species (in preparation).

C. dissimilis described from a single damaged

male specimen is now referred to Muacropo-

wtrongyloides beewuse of the shape of the

ocsophagus, tail, and bursa, It differs from other

M. spp. in having very long spicules. The speci-

men Was apparently moribund when collected,

and the buccal, capsule is atypical, Several

specimens of Mucropostrongyloides have

recently been found in which the buccal cap-

sule i in a similar condition.

In the case of C. medioanhulatus, the speci-

men labelled as type is a female Rugopharynx

dustraliy, obviously placed in the tube in error.

No representative of ©. medioanniilatus has

been found, and the species must he regarded

as a species inquirendum,

A revised diagnosis of Cyclostrongylus and

a key to the valid species follows.

Trichonematidae: Small worms, anterior end

with more or less well developed cuticular

collar pierced by amphids and cephalic papil-

lac; well developed usually transversely striated

buccal capsule; ocsophagus with anterior cylin-

drical part followed by constriction surrounded

by nerve ring, before terminal bulb; cervical

* Department of Zoology, University of Adelaide, North Tce. Adelaide. S000.

20 PATRICIA M. MAWSON

papillae setiform, near nerve ring. Male: bursa

not deeply lobed, externo-dorsal ray arising

separately or with laterals, dorsal ray bifurcate

each branch giving off a lateral stem. Female:

tail more or less conical, vulva near anus.

Parasites of stomach or oesophagus of

macropod marsupials. Type species C. wal-

labiae Johnston & Mawson, 1939 (syn. Pharyn-

gostrongylus gallardi Johnston & Mawson,

1942; Spirostrongylus gallardi: Mawson, 1955;

Oesophagonastes_ gallardi: Mawson, 1965).

From Macropus bicolor, N.S.W.

Other species:

C. gallardi Johnston & Mawson, 1939. From

M. rufogriseus, N.S.W.

C. kartana (Mawson 1955), (syn. Spiro-

strongylus kartana; Oesophagonastes kar-

tana: Mawson, 1965). From M. eugenii, S.

Aust., and M. rufogriseus, Qld.

C. leptos (Mawson 1965), (syn. Oesophago-

nastes leptos). From Macropus dorsalis, Q\d.

C. parma (Johnston & Mawson 1939), syn.

Pharyngostrongylus parma; Spirostrongylus

parma: Mawson, 1955; Oesophagonastes

parma: Mawson, 1965. From Macropus

parma, N.S.W.

This genus differs from Rugopharynx

Monnig mainly in the presence of a cuticular

collar, in the shape of the oesophagus, and in

the shape and character of the bursa.

Key to species of Cyclostrongylus

1. Buccal capsule wall wider anteriorly than pos-

GOST OP EY: 83 ys 1,55 OG ee SERS ober Shes phe.

Wall of buccal capsule not markedly wider

PUENIOT NY | seca he Fld Ath Metis axtstin, atacanh ec 4

2. Cuticular collar not well developed .....................

C. parma

Cuticular collar well developed ...............00.000.. 3

3. Buccal capsule short, about equal to its external

diameter at anterior end .............. C. wallabiae

Buccal capsule long, at least twice external

diameter at anterior end ................ C. kartana

4. Cuticular collar well developed; buccal capsule

wider than long ...................000006 C. gallardi

Cuticular collar not well developed; buccal cap-

sule wider than long ......................... C. leptos

References

JoHNsTon, T. H. & Mawson, P. M. (1939).—

Strongylate nematodes from marsupials in

New South Wales. Proc. Linn. Soc. N.S.W.

64, 514-536.

JOHNSTON, T. H. & Mawson, P. M. (1940).—New

and known nematodes from Australian mar-

supials. Proc. Linn. Soc. N.S.W. 65, 468-476.

Jounston, T. H. & Mawson, P. M. (1942).—The

Gallard Collection of parasitic nematodes in

the Australian Museum. Rec. Aust. Mus. 21,

110-115.

Mawson, P. M. (1965).—Notes on some species

of nematodes from kangaroos and wallabies,

including a new genus and three new species.

Parasitol. 55, 145-162.

REVISION OF THE AUSTRALIAN TERTIARY SPECIES ASCRIBED TO

LIMATULA WOOD (MOLLUSCA, BIVALVIA)

BY M. F. BUONAIUTO

Summary

Limatula crebresquamata Tate (Late Eocene-Miocene) and Limatula jeffreysiana Tate (early

Miocene) are revised. The Late Eocene L. margaritata sp. nov. and the Pliocene L. ludbrookae sp.

nov. have hitherto been mistaken for L. jeffreysiana. The Early Pliocene L. subnodulosa Tate is

shown to be a synonym of Limea (Gemellima) austrina Tate. A brief discussion and revision of the

Tortachilla Limestone is given and a new procedure for S.E.M. photography is described.

REVISION OF THE AUSTRALIAN TERTIARY SPECIES ASCRIBED TO

LIMATULA WOOD (MOLLUSCA, BIVALVIA)

by M. F. BUoNAluToO*

Summary

Auwanaiuto, M. F, (1977).—Revision of the Australian Tertiary species ascribed to Limatula

Wood (Mollusca, Bivalvia), Trans, R, See, S. Aust, 101(1), 21-33, 28 February, 1976.

Limatula crebresquamata Tate (Late Eocene-Miocene) and Limatula jeffreyslana Tate

{Barly Miocene) are revised. The Lute Rocene L. mergaritata sp. nov. and the Pliocene L, Ind-

hroakae sp, nov. have hitherto been mistaken for L, jeffreysiana, The Early Pliocene L, sub-

nodulosa Tate is shown to be a synonym of Limea (Gemellima) austrina Tate. A brief discus-

sion and revision of the Tortachilla Limestone is given and a new procedure for §8,6.M.

photography is described,

Introduction

Hitherto only three fossil species of Limatula

Wood were known or recognized in the Aus-

tralian Tertiary: L, jeffreysiana (Tate), now

known to be Early Miocene in age, the Late

Oligocene-Early Miocene Lb, crebresquameata

Tate, und the Early Pliocene Limarula sub-

nodulasa Tate, here believed to be a worn

specimen of Limea (Gemellima) austrina Tate.

Observations made during a current revision of

the Eocene Molluscan faunas have revealed that

two specimens of the series of L, jeffreysiana

borne on the tablet SAM T972. from Tate's

collection, represent two other species: the Late

Eocene L, mergaritata sp. nov, (T972-M) and

the Pliocene L, ludbrookae sp. nov, (T972-D).

The material here examined is in the Tate

Collection and Molluse Collection housed in

the South Australian Museum (SAM), which

remains the property of the Department of

Geology and Mineralogy, University of Ade-

laide.

Optimal S.E.M, results were obtained by pre-

treating the specimens by exposure to osmium-

tetraoxide vapour for twelve hours, followed hy

coating with carbon and gold-palladium, Car-

bon or silver dag or tragacanth glue did not

influence the resultx, and problems of high

charging were eliminated other than where

there was iniperfect specimen-stub connection

or coating, It produced excellent resolution

even of very rough surfaces at high magnifica-

tions, and represents an extreme simplification

of Robertson’s (1971) technique,

Systematic descriptions

CLASS BIVALVIA Linné, 1758

SUBCLASS PTERIOMORPHIA Beurlen,

1944

ORDER PTERIOIDA Newell, 1965

SUBORDER PTERIINA Newell, 1965

SUPER-

FAMILY LIMACEA Rafinesque, 1815

FAMILY LIMIDAE Rafinesque, 1815

GENUS Limatula Wood, 1839

Diagnosis, Shell small, oval, higher than long,

inflated, Without Wmbonal ridges; auricles small,

subequal; margins not gaping; hinge edentu-

lous; ornaments of primary radial riblets and

secondary concentric costellae, more conspicu-

ous on the dorsal and ventral regions; concen-

tric ornaments can develop into primary in the

anterior and posterior regions; median sulcus

can occur (after Cox & Hertlein, 1969, p.

NGR9).,

Limatula margaritata sp. nov,

FIGS |, 6-9

Derivation of name. From the Latin mareari-

tatus, beaded, because of its beaded ribs.

Holotype. SAM P18343, figs 6-7, 9.

Type-formation, Tortachilla Limestone, Late

Eocene.

* Department of Geology & Mineralogy, University of Adelaide, North Tce, Adelaide, 8. Aust. 5000.

22 M. F. BUONAIUTO

Type-locality. Maslin Bay, Willunga Sub-Basin.

St Vincent Basin.

Material. 172 specimens (21 RV, 22 LV, 129

VV) generally very badly preserved; the topo-

type SAM 1972-M from Tate’s collection,

Description. Shell small, oval, higher than long,

inflated, slightly inequilateral; umbo central, in-

flated with little protruding orthogyrate beaks.

Margins: anterior and posterior subelliptical,

winged; ventral very elliptical. Margin connec-

tions: postero-ventral imperceptible; antero-

ventral rounded, angular. Auricles small, sub-

equal, longer than high, with protruding ends.

Longitudinal shell section convex with maxi-

mum at the posterior ridge. Regions: anterior

flatter and steep; posterior convex and steep;

dorsal and ventral more convex and steeper to

the ventral margin. Commissure region crenu-

late. Cardinal area narrow and rather long,

resilifer deep, hinge edentulous.

Ornament. About 40 radial triangular costae

with narrow trapezoid trough-shaped_inter-

spaces, wider to the anterior and posterior

regions. The costae fade to the auricles; marked

concentric grooves separating concentric weak

costellae; the costellae thicken to the auricles.

Costa-costella intersections bear triangular

beads. Auricles with concentric costellae and

growth lines.

Observations. This form was included by Tate

in L. jeffreysiana which is Miocene. A topotype

is mounted on the tablet SAM T972 labelled

Limatula jeffreysiana (Tate). Distinctive differ-

ences between the species are tabulated in the

comparative synopsis in Table 1. The holotype,

although rather juvenile, was chosen because it

is the only specimen in a good state of pre-

servation, and has a sure stratigraphic location.

Stratigraphic range. Tortachilla Limestone to

Blanche Point Transitional Marls (lowermost

member of Blanche Point Marls); Late Eocene.

Limatula jeffreysiana (Tate, 1885)

FIGS 1-5

1877 Lima (Limatula) subauriculata Tenison

Woods, p. 113 (non Montfort). 1885a Lima

jeffreysiana Tate, p. 208 (nom. nud.). 1885a Lima

subauriculata: Tate, p. 213 (non Montfort). 1885b

Lima jeffreysiana Tate, p. 230. 1886 Lima (Lima-

tula) jeffreysiana: Tate, p. 119, pl. 4, fig. 8 (pars).

1896 Limatula jeffreysiana: Pritchard, p. 128. 1897

Lima (Limatula) jeffreysiana: Harris, p. 311. 1899

Lima (Limatula) jeffreysiana: Tate, p. 273. 1924

Lima jeffreysiana: Marwick, p. 323.

Material. 11 specimens (4 LV + 5 RV + 1

BV) generally well preserved. (SAM T1972

A-C, E-L; Coll. Tate.)

Description. Like L. margaritata. Differs from

it by greater height, less inflation, narrower

ventral margin, by longer and narrow ears with

more protruding ends.

Ornament. 34-37 triangular thin radial ribs.

more spiny on the ventral region, with broad

concave to flattened interspaces, narrower on

the dorso-ventral region. broader to the an-

terior and posterior, where ribs fade to the

auricles. Very fine growth lines; broadly inter-

spaced concentric costellae, more marked in

the anterior and posterior regions. Auricles

with concentric costellae, Median radial sulcus

shallow and observable only in younger speci-

mens.

Observations. The tablet SAM T972_ bears

specimens of L. jeffreysiana (Tate), together

with specimens here described as L. margaritata

sp. nov. (T972-M) and L, ludbrookae sp. nov.

(T972-D),

Tenison Woods referred the species to the

living L. subauriculata (Montfort, non Mon-

tagu). Tate (1885a, 1885b) distinguished it as

a new fossil species and remarked its close

affinity with the living L. strangei Sowerby

(MacPherson & Gabriel 1962, p. 308, fig.

3501; Cotton & Godfrey 1938, p. 108, fig. 97;

this study, fig. 20-26). Later, Tate (1899) also

referred to L. jeffreysiana a New Zealand fossil

form, mistaken for the living L. bullata Born

(Hutton 1873, p. 33). Marwick (1924, p. 323)

separated the New Zealand form, that was later

named by Finlay L. maoria (Finlay 1927, p.

454, figs 104-6). The holotype has not been

located; it does not appear to be in the Tas-

manian Museum, Hobart (Ludbrook 1967).

The two specimens found in Tate’s collection

are both juveniles and one (T972-L) is broken.

Hence, it is here considered inappropriate to

choose one of them as neotype.

Distribution. Table Cape, Bass Basin (type);

Muddy Creek, “Murray River” Snapper Point,

Blanchetown, “Spring Creek”, Other localities

Fig. 1. Tablet SAM T972 (Coll. Tate) bearing specimens of L. jeffreysiana (Tate). T972-D: a paratype

of L. ludbrookae sp. nov.; T972-M: a topotype of L. margaritata sp. nov. (x 1.1).

Figs 2-5. Limatula jeffreysiana (Tate), plesiotype (SAM T972-A), LV, Muddy Creek; (2) dorsal view

(x 2); (3) ornaments, particular from ventral region (x 4); (4) anterior auricle (x 9.3);

(5) umbonal region and posterior auricle (x 3.8).

Ake ee

TP Spreng Cre

Cafan—

“4

asa,

S OF LIMATULA WOOD

alt Se.

Leos os

‘

—b Irnvecelon Adagio.

SPECIE

. Mas hike Cree

Sef,

VF a

tally

appen Pe*

uchi frum |

Name fey

AUSTRALIAN TERTIARY

[Z la 1.24

M. F. BUONAIUTO

Figs 6-9. Limatula margaritata sp. nov., Maslin Bay; (6) Holotype, SAM P18343, RV, antero-dorsal

view (x 14); (7) holotype, anterior view (x 15); (8) topotype, SAM T972-M (x 3.6); (9)

ornaments, particular from holotype’s postero-ventral region (x 44).

Figs 10-11. Limatula ludbrookae sp. nov.; SAM T972-D, Aldinga; (10) dorso-ventral view (x 6);

(11) ornaments, particular from ventral region (x 16.2).

AUSTRALIAN TERTIARY SPECIES OF LIMATULA WOOD

Figs 12-15. Limea (Gemellima) austrina Tate, holotype of Limatula subnodulosa Tate, SAM T1799,

Muddy Creek; (12) dorsal view (x 8); (13) interior view (x 8); (14) hinge and cardinal

area (x 17); (15) ornaments, particular from dorsoventral region (x 20 c).

Figs 16-19. Limatula crebresquamata Tate, holotype, SAM T978-A, “Spring Creek”; (16) dorsal view

(x 3.75); (17) umbo and anterior auricle (x 11.25); (18) posterior auricle (x 11.25); (19)

ornaments, particular from dorso-ventral region (x 15).

M. F. BUONAIUTO

Figs 20-26. Limatula strangei (Sowerby); (20) type figured by Cotton & Godfrey, SAM 15145, Hard-

wicke Bay, South Australia, LV (x 1.2); (21) hinge and cardinal area, specimen SAM

D9431-B (Coll. Verco) (x 15); (22) dorso-umbonal region, SAM D15146-A (x 10); (23)

posterior auricle, SAM D15146-A (x 40); (24) anterior auricle, SAM D15146-A (x 40);

(25) ventral region, SAM D15146-A (x 10); (26) ornaments, particular from the dorsoven-

tral region, SAM D15146-A (x 80).

AUSTRALIAN TERTIARY SPECIES OF LIMATULA WOOD

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-ANAODODIIO Sulpnioid yA Sa] Bos saovjd sulos ul AWWSI[S padeys-A = payeyur = ‘[euosrqns pipwupnbsaiqasd

ALVI MOLIBU PUe BUC] poy [jam puk saul] Mois = ‘uty ‘YSIYy AIA pp “ramoleu ‘d3aap AIA 01 [kao DINIVUT

ap[[a}soo

d14juasU0D SdA0013 Jolajsod pue

auy puke spua Aq pajeiedas IOL9]UL 34} OF “sou ‘ds

ANAO0F Surpnijoid yi a1qudaoied aey]aIsod = Papeaq ‘iejnsurin = japim ‘padeys- 7 —s paeyul woys pipiLipdio

ALvV1 JAMOLIBU “JASU0] TMs 1nq Surpey ysipunos ‘auy ‘peoig dp ‘amolieu MoTpeYys asoul = Jaye ‘[eAO DINIDUT

aay Jenbaqns UOIsayy S]USUTEUIQ sqry [eIpey saoedsioquy uoneguy aulyno saidadg

ipjnguevlsy sieq IOLI3}SOg ainuesue;d jerpey

pue 1olajuy

uo sqry [eIPed

SoustajavADYD jwasojoydiou fo sisdouds aatvsnduiay

T ATaVL

28 M. F. BUONATUTO

quoted by Dennant & Kitson (1903) are here

omitted because specimens from those localities

were not available for checking.

Stratigraphic range. As known at present, Eatly

to Middle Miocene (Quilty 1966; Ludbrook

1973).

Limatnula tudbrookae sp. nov-

FIGS |, 10-11. 27-38

Derivation of name, From Nelly Hooper Lud-

brook of Adelaide for her devotion to Palae-

ontology,

Holotype. SAM P8360, figs 27-28.

Type-formation. Dry Creek Sands (Late Plio-

cene, Yatalan),

Type-loeality, Salishury Bore, 1942, hd. Munna

Para, sec. 4000), at 100 m depth.

Material, \0 specimens from Salisbury Bore

(6 LV +3 RV +1 VV); 1 LY specimen from

Tate Collection (SAM T872-D}, Two hroken

specimens from Abattoirs Bore-

Deseriptian. Shel) oval, auriculated, very high

and narrow, very inflated, sub-inequilateral;

\imbones with small protruding and prosoclite

beaks. Nonegapingt margins; anterodorsal and

posterodorsal represented by two subequal

auricles, longer than high; anterior subellip-

tical very long; posterior very lony, slightly

more ellrptical: yentral yery elliptical. Margin

confections: ntenoranterodorsal and pos-

lervor-posterodorsal angular and concave: others

imperceptible,

Longitudinal shell section, subtrapezoidal.

yery convex, Regions: anterior and posterior

very declivous, subconvex; dorsoventral con-

vex, more gently declivous, Connections be-

iween [he regions imperceptible. Cardinal area

broad, longer than high, horizontally striated:

resilifer triangular, broad, rather deep with

curved margins. Hinge edentulous. Inner sep-

tum below the cardinal plate, Interior with

marked median rib and fine regular striae.

Maonorniyariin, posterior scar at high middle

posterior position near to the median mb. Pal-

lial line marked, Commissure region smooth

excep! on ventral margin where uw ts highly

erenilated.

Ortament. 29 triangular radial costae with

broader trapezoidal interspaces. From the beak

to the ventral (Malgin a marked broad median

sulcus, Fine concentric growth lines; fine

vrowth rugae in odult-senile stage. At costa-

line/ruga intersections short spines. On the

anterior and posterior region, the costae fade

abruptly and the growth lines and rugae pre-

demimate. Auricles with concentric yrowth Imes

anid fugae,

Observations, This form was initially mistaken

by Tate for L, Jéeffreysiana, The juvenile SAM

T972-D from Aldinga is broken at the umbo

and is the only specitmen available from out-

crop. A search in the uncatalogued part of

Tate's collection still kept in the Departmeut of

Geology and Mineralogy of the University of

Auelaide, led to the discovery of 8 juveniles, 1

adult, and 1 semle specimen from Salisbury

Bore, These specimens corroborate the: distine-

tion of this form from ZL, jeffreysiana on the

basis of rib and interspace shape and shell geo-

metry. The senile was chosen as holotype be-

cause of its perfect preservation, The specimen

of L, jeffreysiana (Tate) reported by Reynolds

(1953) in the Pliocene of Aldinga should be

more probably referred to L, ludbrookae,

Distribytion, St Vincent Basin} Aldinea Bay,

Hallett Cove Sandstone; Abattoirs Hore. Salis-

bury Bore (type), Dry Creek Sanets.

Stratigraphic ringe. Yatalan (Late Pliocene}.

Limutula crebresquamata (Tate 1399)

FIGS 16-19

4 Mais Lime (ULiniwirla erebresyucrmeta Tate, p.

Material, Three specimens borne on the tablet

SAM T978 (3 LV): T978-A, the holotype

hroken and glued up on the untero-ventral

region; T978-B, well preserved, juvenile)

TIT8-C, broken, with the dorsal region, the

umbo und the aurictes missing.

Description. Like the above described species,

but differs by a shorter oval to subtrigonal out

line, more inflation, and the occurrence of ribs

on the unterier and posterior regions. Cardinal

area narrow, longer than high, horizontally

striated; resilifer triangular, concave, rather

deep. Hinge edentulous. Interior with marked

radial ribs and narrower interspaces, Pallial

line and adductor scar imperceptible. Commis-

Figs 27-34. L. ladbrowkae sp, nov., Salisbyry Bore; (27) Holotype (SAM P18360) dorsal view (x

22); (28) holotype, interior (x 2.2); (29) paratype (SAM PI83960) A/LV. darsal view

(S 9): (30) Paratype (SAM PI8360B) LY, interior view (x 9); (31) pirnivpe (SAM

PISSG0A) anterior auricle (x35); (32) paratype A, posterior auricle (xX 37), €33} para

type B, cardinal urea (x 27); (44) paratype A, particular median sulcus (x 72)

<—e

eo _

|. hl SS

- 2

fe)

—

©)

ea}

lal

—

ag SAL Tam, sone

30 M. F. BUONAIUTO

sure region smooth, except the ventral heavily

crenulated.

Ornament. 44 ribs, very high, thin, in some

places dichotomous, bearing wide, thick, chev-

ron-shaped concentric scales, separated by

regular rather broad concentric furrows. Radial

interspaces U-shaped, narrower in the dorso-

ventral region, increasing in width to the

anterior and posterior auricles.

Observations. The morphology of this form

agrees with the diagnosis of Limatula Wood of

Cox & Hertlein (1969, p. N389), except in the

strong radial ribbing of the anterior and pos-

terior regions. The median sulcus is obscured

by the heavy costae and squamae, revealed only

by the inner median sulcus. An inner ridge

just below the cardinal area may represent an

embryonic septum as in L. ludbrookae,

Localities. “Spring Creek” (Tate 1899), Bird

Rock, Torquay (Fleming, in litt. 1974).

Stratigraphic range. Late Oligocene-?Early

Miocene (Janjukian-Longfordian).

Observations. Neither the holotype nor para-

types were figured,

GENUS Limea Bronn, 1831.

SUBGENUS Gemellima Iredale, 1929.

Limea (Gemellima) austrina Tate, 1887

FIGS 12-15, 36-41

1887 Limea austrina Tate, p. 73, pl. 4, fig. 7.

1899 Limatula subnodulosa Tate, p. 273. 1907

Limaea austrina-Verco, p. 315. 1929 Gemellima

austrina—Iredale, p. 166. 1938 Gemellima austrina

—Cotton & Godfrey, p. 107, fig. 93.

Material. 1 specimen (LV) (SAM T1799), the

holotype of L. nodulosa Tate; several hundred

specimens (SAM Lot T17).

Description. Shell small, thick, trigonal, slightly

higher than long, slightly inequilateral, very

inflated; umbo inflated with central orthogyrate

beaks protruding a little. Margins: anterior sub-

elliptical; posterior elliptical, both winged; ven-

tral very elliptical. Margin connections:

broadly angular; the antero-ventral rounded.

Ears triangular, very narrow, and subequal.

Longitudinal shell section very convex.

Regions: anterior and posterior very steep;

dorsal declivous; ventral very steep. Cardinal

area longer than high; resilifer triangular

broad, concave, and shallow; hinge with very

fine vertical teeth; monomyarian with orbicular

adductor scar high in the posterior region; pal-

lial line imperceptible or not easily distinguish-

able from other concentric grooves in the shell

interior; commissure region heavily crenulated.

Ornament. Outer: 25 radial large massive pro-

truding ribs with narrower deep U-shaped

interspaces; fine regularly interspaced concen-

tric costellae; rib-costellae intersections produc-

ing short subtriangular spines; irregularly inter-

spaced broad concentric constrictions. Ears

bearing only concentric costellae. /nner: fine

radial grooves corresponding to the outer ribs;

irregular concentric grooves corresponding to

the outer concentric constrictions.

Observations. The rediscovered holotype of

Limatula subnodulosa Tate, 1899 is just a worn

and polished fossil specimen of Limea (Gemel-

lima) austrina Tate, 1887.

Investigations on several hundred specimens

of a sample from Investigator Strait, 36.6 m

depth, showed that as soon as the disarticulated

valves lose the ligamentary organic matter,

their hinge, composed of very fine vertical

teeth, is abraded very easily; if the abrasion go

further, the crenulated commissure region can

be practically smoothed out and the spines on

the ribs reduced to blunt nodules or worn out

too.

This can explain Tate’s erroneous determina-

tion. Instituting Limatula subnodulosa he re-

marked that the shell displays Limea charac-

teristics, and, although he suspected it was

reworked, he did not consider the possibility

that it could be actually a worn specimen of

Limea. Limea (Gemellima) austrina is the type

species of Gemellima Iredale, considered by

Newell (1969) a subgenus of Limea Bronn.

Study of the above-mentioned sample indi-

cated two main morphs connected by transi-

tional forms. One is shorter and longer, less

inflated, with broader interspaces between ribs.

The other is higher and narrower, more

inflated, with narrower interspaces (subnodu-

losa type).

Environmental observations. Limea (Gemel-

lima) austrina was dredged in S.A. waters at

14.6-366.0 m, alive from 27.45-40.3 m; the

optimum depth for populations seems to be

36.6 m (Verco 1907).

Fig. 35. L. ludbrookae sp. nov. paratype B, prodissoconch (x 135).

Figs 36-41. Limea (Gemellima) austrina Tate, Investigator Strait. (36) LV, juvenile, dorsal view (x

9.5); (37) LV, worn juvenile, dorsal view (x 9.5); (38) worn hinge (x 18); (39) hinge (x

18); (40) particular posterior hinge (x 36); (41) particular dorso-ventral ornaments (x 18).

; the 2)

HAG oy

et.

“

ine

‘*

jam

ie)

cal

ea)

22)

32 M. F. BUONAIUTO

Distribution. Spencer Gulf and Gulf St Vin-

cent, recent deposits; Muddy Creek, Grange

Burn Coquina; Otway Basin; Limestone Creek,

W. Victoria (fide Dennant).

Stratigraphic range. Early Pliocene (Kalimnan)-

Holocene.

Acknowledgments

I am grateful to Dr N. H. Ludbrook for con-

tinuing advice and encouragement; to Dr B.

McGowran (Department of Geology and

Mineralogy, University of Adelaide), to S.

Shafik (Department of Palaeontology, B.M.R.,

Canberra), and to J. M. Lindsay (S.A. Geo-

logical Survey) for stratigraphic information.

I would also like to thank Dr McGowran for

reading the manuscript, Dr C. A. Fleming

(New Zealand Geological Survey), whose

interest in this group initiated this study, and

the Director of the South Australian Museum

for lending specimens studied. The work was

carried out in the Department of Geology and

Mineralogy, University of Adelaide, during

tenure of a University Research Grant.

This paper is dedicated to Mr B. C. Dawes,

Ashland Oil, Canada, remembering our fruitful

and stimulating discussions.

References

Cotron, B. C. & Goprrey, F. K. (1938).—The

Molluscs of South Australia, Part 1. The Pele-

cypoda. (Govt Printer: Adelaide.)

Cox, L. R. & Hertiein, L. G. (1969).—Family

Limidae. Jn Moore R. C. (Ed.) “Treatise on

Invertebrate Paleontology. Part N, Mollusca

6. Bivalvia’. 1. (University of Kansas & Geo-

logical Society of America: New York.)

DENNANT, J. & Kirson, A. E. (1903).—Catalogue

of the described species of fossils (except

Bryozoa and Foraminifera) in the Cainozoic

fauna of Victoria, South Australia and Tas-

mania. Rec. Geol. Surv. Vict., 1(2), 89-147.

Fintay, H, J. (1927).—A further commentary on

New Zealand Molluscan systematics. Trans.

N.Z. Inst., 57, 320-485.

GuILcHeR, A. (1953).—Essai sur la zonation et Ja

distribution des formes littorales de dissolu-

tion du calcaire. Ann. Géogr., 62, 161-179

Harris, G. F, (1897).—Catalogue of Tertiary

Mollusca in the Department of Geology,

British Museum (Natural History). Part 1.

The einai nite Tertiary Mollusca. (Lon-

don.

Hutron, F. W. (1873).—Catalogue of the Ter-

tiary Mollusca and Echinodermata of New

Zealand in the collection of the Colonial

Museum. (Govt Printer: Wellington. )

IREDALE, T. (1929).—Mollusca from the Con-

tinental Shelf of Eastern Australia. No. 2. Rec.

Aust. Mus., 17(4), 157-189.

Jaanusson, V. (1961). Discontinuity surfaces in

Limestones. Bull. Geol. Inst. Univ. Uppsala,

40, 221-241.

Jenkins, R. J. F. (1974).—A new giant penguin

from the Eocene of Australia. Palaeont.,

17(2), 291-310.

Krawiec, W. (1971).—Solution hole breccias. Jn

Multer, G. H. (Ed.) Field Guide to some car-

bonate environments. Florida Keys and

Bahamas. (Farleigh Dickinson University.)

Linpsay, J. M. (1969).—Cainozoic foraminifera

and stratigraphy of the Adelaide Plains Sub-

basin, South Australia. Bull. Geol. Surv. S,

Aust., 42, 1-60.

Linpsay, J. M. & LupBrook, N. H. (1966).—The

Aldingan Stage. Quart. Geol. Notes Geol.

Surv, S. Aust., 19, 1-2

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tiary rocks of South Australia. Trans. R. Soc.

S. Aust., 87, 5-15.

Lupsroox, N. H. (1967),—Tertiary Molluscan

types from Table Cape, in the Tasmanian

Museum, Hobart. Pap. Proc. R. Soc. Tas.,

101, 65-69.

Lupsproox, N. H. (1973).—Distribution and

stratigraphic utility of Cenozoic Molluscan

faunas in Southern Australia. Sci. Rep.

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241-261.

McGowran, B., Harris, W. K. & Linpsay, J. M.

(1970).—The Maslin Bay flora, South Aus-

tralia. 1. Evidence for an early Middle Eocene

age. N. jb. Geol. Paldéont. Mh., 1970 (8), 481-

485.

McGowran, B., Linpsay, J. M. & Harris, W. K.

(1971).—Attempted reconciliation of Tertiary

biostratigraphic systems. Otway Basin. Jn H.

Wopfner & J. G. Douglas (Eds), “The Otway

Basin of southeastern Australia.” Special Bull.,

Geol. Survs. S. Aust., Vict., 273-281

Marwick, J. (1924).—An examination of some

of the Tertiary Mollusca claimed to be com-

mon to Australia and New Zealand. Rep. Aus-

tralas, Ass, Ady. Sci., 16, 316-331.

PrircHarD, G. B. (1896).—A revision of the fos-

sil fauna of the Table Cape Beds, Tasmania,

with description of the new species. Proc. R.

Soc, Vict., 8, 74-150,

Quitty, P. J. (1966).—The age of Tasmanian

Marine Tertiary Rocks. Aust. J. Sci,, 29(5),

143-144.

ReYNOLDs, M. A. (1953).—The Cainozoic succes-

sion of Maslin and Aldinga Bays, South Aus-

tralia. Trans. R. Soc. S. Aust., 76, 114-146.

ROBERTSON, R. (1971).—Scanning electron micro-

scopy of Planktonic Larval Marine Gastropod

Shells. Veliger, 14, 1-13.

Tate, R. (1885a).—Notes of a critical examina-

tion of the Mollusca of the Older Tertiary of

Tasmania, alleged to have living representa-

tives. Pap. Proc. R. Soc. Tas. (1884), 207-214.

TATE, R. (1885b).—Description of new species of

Mollusca of the Upper Eocene Beds at Table

Cape. Pap. Proc, R. Soc. Tas, (1884), 226-

231.

AUSTRALIAN TERTIARY SPECIES OF LiIMATULA WOOD 33

Tate, RK. (1886)—The Lamellibranchs of the

Older Tertiary of Australia, Part 1, Trans. Re

Soc. 8. Aust., & 96-158.

Tate, RK. (1887),—Description of some new

species of South Australian marine and fresh-

water Mollusca. Trans. R. Soc. 8. Ausr., 9

OR-TS

Tare, R- €1899}—A revision of the Older Ter-

vary Motlusea of Australia. Part 1. Trans. R-

Sec. 8. Aust., 23, 249-277.

Veroo, 1. C. (1907).—WNotes on South Australian

Marine Mollusca, with description of new

species—Part VIL, Trans. R. Soc. S. Aust.,

31, 305-315,

Wrntworte, ©, K, (1939}-—Marine bench-form-

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Geormorph,, 2(1), 3-25,

Woons, J. E. (1877)—WNotes on the fossils

referred to in the foregoing paper. Pap. Prac,

R, Soc, Tas. (1876),91-116

Appendix

Stratigraphical observations on Tortachilla Lime~

stone Reynolds, 1953 (Lower Aldingan Stage).

A study of the lithostratigraphy of the fos-

siliferous Eocene beds at Maslin Bay will be

presented elsewhere. Meanwhile a summary is

necessary for adequate stratigraphic characteriza-

uon of Limaiula and other molluscs.

The Tortachilla Limestone (Reynolds 1953)

considered by Ludbrook & Lindsay (1966) and

Ludbrook (1973) to be the lowest rock unit in the

stratotype for the Aldingan stage (Late Eocene),

displays erosional unconformities, The major un-

conformity (Jenkins 1974, figs 1, 3) separating the

lower member (Polyzoal Limestone Member of

Reynolds) from the upper one (Blanche Poimt

Glauconitic Limestone Member ot Reynolds), is

a deeply pitted erosional surface on the topmost

limestone in the Polyzoal Limestone. The abun-

dant subvertical pits are filled by the pladconitic

sands, in places cemented by sparite, of the

Blanche Point Glauconitic Limestone Member. By

analogy with the studies of Jaanusson (1961 p, 232

ef seq.), Krawiec (1971), pp. 128-31), and

chiefly by Guilcher (1953) and Wentworth (1939)

thrs unconformity could be interpreted as produced

by sub-aerial dissolution of the emergent limestone,

i.e. karst. The constant widespread occurrence of

the pits can be explained in the negligible slope of

the formation at the time of emergence, thus pre-

venting the accumulation of beach deposits thick

enough to protect the limestone from the action

of erosive and dissolutive agents.

The discovery of this karst surface leads to 4

stratigraphic revision of the Tortachilla Limestone,

restricting the formation to its previous lower mem-

ber and referring the Glauconitic Limestone Mem-

ber to the Blanche Point Transitional Marls, to

which it belongs in a new episode of sedimenta-

tion. The record of this karst surface ts the evi-

dence of a lacuna that covered a span of time

still unascertainable but longer, however, than has

been considered until now,

A precise correlation of the Tortachilla Lime-

stone in terms of planktonic foraminiferal zone is

not yet possible,

5. Shafik (pers. comm. 1974) stated “the ranges

of the few calcareous nannofossils extracted from

Tortachilla Limestone are confined mainly to the

Middle to Late Eocene",

McGowran & Lindsay (pers, comm, 1974-5)

und Ludbrook (1973) support a probable early

Late Eocene age for this formation. Lindsay

(1969) considered the undifferentiated deposits of

Tortachilla Limestone (or its equivalent) and

Blanche Point Transitional Marls, in the Adelaide

Plains Sub-basin to be early in the Late Eocene.

At present, the only two biostratigraphic con-

trols on the older part of the section at Maslin

Bay are;

—the microfloral asemblage occurring in North

Maslin Sands and belonging to the Proteacidites

confragasus zone, earliest Middle Bocene in age

McGowran, Harris, & Lindsay 1970), but pos-

sibly latest Early Eocene (MeGowran pers,

comm 1975}.

— The Hanrkenina primitiva sub-zone occurring in

the Transitional Marls at Maslin Bay, south-

ward of “Uncle Tom’s Cabin", 80-115 cm above

the described karst surface und estimated to be

Mid-Late Eocene in age (McGowran, Lindsay

& Harris 1971).

DISTRIBUTION AND SEDIMENTS OF MANGROVE FORESTS IN SOUTH

AUSTRALIA

BY A. J. BUTLER, A. M. DEPERS, S. C. MCKILLUP AND D. P. THOMAS

Summary

A survey of forests of the mangrove Avicennia marina in South Australia was conducted in

summer, 1974-75. This paper describes the distribution of the forests and contains detailed maps of

the major stands. Smear slides of the sediments have been examined and on this basis two

geographically distinct types are identified. The dynamic relationship between the sediments and

the organisms growing within them is discussed. Extinct mangrove swamps at three sites are

described and the past distribution of mangroves is discussed. Finally we comment on the

composition of the communities of organisms in South Australian mangrove swamps.

DISTRIBUTION AND SEDIMENTS OF MANGROVE FORESTS IN SOUTH

AUSTRALIA

by A. J. Butier,* A, M. Derers,7 S.C. McKitiupe* and D, P. THomast

inary

Buriex, A. J., Deeers, A. M., McKitiue, 8. C. and Tuomas, D. P. (1977),—Distribution

and sediments af mangrove forests ln South Australia. Trans. Ro Soc. §. Aust, 101(1),

35-44, 28 February, 1977.

A survey of forests of the mangrove @vicennia marina in South Australia was conducted

in slimmer, 1974-75. This paper describes the distribution of the forests and contains detailed

maps of the major stands, Smear slides of the sediments have been examined and on this basis

two geographically distinct types are identified. The dynamic relationship between the sedi-

ments and the organisms growing within them js discussed. ExtincL mangrove swamps at three

sites are described and the past distribution of mangroves is discussed. Finally we comment

on the composition of the communities of organisms in South Australian mangrove swamps.

Introduction

Mangrove forests are most complex and

luxuriant in the Wet tropics of the Indo-West-

Pacific region (Macnae 1968), but extend onto

desert shores and to latitudes at high as that of

Westertiport Bay, Victoria (38°22'), South of

the Queensland/New South Wales border there

are Only two species of mangroves, Avicennia

marina and Aegiceras corniculatum, and south

of Sydney there is only Avicennia marina

(Forst.) Vierh, (Macnae 1966), The stands of

this species at Westernport Bay, Victoria, are

the southern-most mangroves in the world.

In South Australia, the flora and fauna of

mangroves are briefly mentioned by Womersley

& Edmonds (1958). Wester (1967)! surveyed

the distribution of mangroves throughout South

Australia using both aerial photographs and

inspections on the ground. There is evidence

that the South Australian mangroves have been

more extensive than they are today; Cotton

(1949) reported the exposure of an old man-

grove mud-flat under the sand of the beach at

* Department of Zoology, University of Adelaide, N-

Glenelg and he suggested that mangroves lived

“until a comparatively short time ago’ as far

south as Port Noarlunga,

It has frequently been argued that South

Australian mangrove forests are important com-

munities in a number of ways, for example in

the support of fisheries and the stabilization of

sediments, and for these and other reasons steps

have been taken to conserve them.* Whilst it

is Clear that mangroves in various places in the

world have such functions it is now obvious

that “mangrove forests” occur in widely vary-

ing conditions and vary considerably in their

composition and functioning (Davis 1940,

Thom 1967, Bird 1971, Carlton 1974, Lugo &

Snedaker 1974, Walsh e al. 1975°). Thus it is

desirable to obtain information, in South Aus-

tralia, about the dynamics of mangrove ecosys-

tems here.

This paper is merely a preliminary step

towards such knowledge, It is based on a sur-

vey* with the following aims: to check the dis-

tribution of mangroves In South Australia and

Tee, Adelaide, § Aust, SODD.

{Present address: Department of Geology, University of Wollangang, Wollongong, N-S.W. 2500,

| Present address: Department of Botany, University of Adelaide, N. Tce,, Adelaide, S. Aust, 5000.

1 Wester, L. L. (1967),—The distribution of the mangrove in South Australia. B.A. {Hons.) thesis,

University of Adelaide, Unpublished,

* Butler, A. J.. Depers. A. M., McKillup, 8. ©. & Thomas, D. P. ((975).—The Conservation of Man-

grove Swamps in South Australia. Report to the Nature Conservation Society of S.A,

* Walsh, G, &., Snedaker, §. C. & Teas, H, J, (1975) (Bds).—Proceedings of the International Sym-

posium on Biology and Management of Mangroves, Honolulu, October 1974. (Institute of Food and

Agricultucul Sciences, University of Florida, Guinesville.)

36 A.J. BUTLER, A. M, DEPERS, S, C, McKILLUP and D. P. THOMAS

ta record as well as possible their biota, the

nature of the sediments jn which they grow,

the condition of each forest in terms such as

the health of the trees and whether the sedi-

ment be accreting or eroding, the types of com-

munities to landward and seaward, and human

activities in and near mangrove forests. Ln

general, it was beyond the scope of the project

to seck explanations for our observations, This

paper records the distribution of mangrove

forests in South Australta at that time, obser-

vations on the sediments, notes on past distri-

bution and brief comments on the composition

of mangrove communities, More detailed

biological and general notes will be published

elsewhere,

Methods

During the summer of 1974-75 almost all

stands of mangroves in the State were visited at

least once; selected areas were re-Visited for

more detailed inspection. On these trips we

were guided by copies of Wester’s! maps, and

4erial photographs! of most of the mangrove

stands.

In addition to stands recorded by Wester,

we Visited several areas where they might have

been anticipated to oceur. For each location,

map accuracy was checked aguinst the 1972

aerial photographs and also by ground surveys.

Notes were made of human activities, and of

the types of habitats lying to landward and to

seaward of the mangroves. Special note was

taken of the health and size distributions of the

(rees, the extent of leaf-damage, and apparent

sedimentary processes al each frea.

Sediment samples were collected and stored

in polyethylene contatners. Preliminary tests

for carbonate content were made In the field

using dilute hydrochloric acid. In the laboratory,

smears of the samples Were mounted on micrp-

scope slides in Caedex resif and eXamined hy

transmitted light at a maximeam of 400% map-

nification, Surface scrapes and the sedimei

smcars were examined for the presence of

microfiora (digtoms, blue-green algae, other

algae), and collectlons of animals were taken

at each site.

Observations and Discussion

DISTRIBUTION

Stands of Avicenaia marina oecur at the

locations shown in Figure t. All are in sheltered

sites as noted by Womersley & Edmonds

(1958), The most extensive stands (Figs 2-8)

are near Ceduna on nerth western Eyre Penin-

sula, at Franklin Harbour on Spencer Gulf,

aftound the heads of both Gulfs, and near both

Port Pirte und Port Adelatide.” We found no

evidence to extend the past distribution of

mMungroves any further soth than the stand at

Glenelg reported by Cotton (1949),

SEDIMENTS

Generally the South Australian mangroves

grow in carbonate-rich sediments, but the per»

centage carbonate varies considerably, both

within and between mangrove communities,

In 4 mangrove community the seaward side

is flanked by extensive inicrtidal shell-grit sands

with or without seagrasses (¢,2, Heterozostera

or Posidonia) whilst to landward the man-

groves are flanked by samphires and

necusionally by extensive supratidal lagoons.

Beach ridges of sheli-grit and dead Posidonia

ure commonty found here, marking the posi-

lion of a previous coastline, prior to large sea-

level changes,

Sediment types

The mangrove sediments were classified

according to depositional texture using Dun-

ham’s (1962) classification,

Two different types of mangrove sediments

could be distinguished on the basis of grain-

size, One, a wackestone-packstone-boundstone,

is confined to Eyre Peninsula; the second, a

boundstone, is found in northern and eastern

Spencer Gulf and Gulf St Vincent,

(1) The wackestone - packstane - boundstone

type of sedimenc was found in all the areas on

Eyre Peninsula, from south of Whyalla to

Davenport Creek. We shall refer to this as the

“West Const” type. it geverally has particles in

the clay to medium sand size category (Folk

1974, p. 25)_ Boundstone sediments are usually

found within the mangroves away from tdal

channels and ¢recks. They are covered by a mal

of blue-green algae thal hinds the top 2-5 cm

of sediment together. Laminue uf such algae

eau be found in long-established sediments. The

houndstone consists dominantly of clay to sill

sized particles, The Wackeslone-packstone sedi

ments are usually found closer to the main

tidal channels where coarser purticles of silt

to sund size are introduced durine the tides.

4 The aenal photographs were taken by the Deparcmend af Lends, S. Aust. in November (972, for the

Fisheries Department, S. Aust. which gow holds them,

"Maps of all sangrove stands are presented in Butler et al (1975)2,

MANGROVE FORESTS IN SOUTH AUSTRALIA 37

Toy .

PT AUGUSTA

FIG 49¥REDCLIFF. PT

ATALA H

\CEDUNA

ACRAMAN CK

“QySMOKY BAY

STREAKY BAY

SQIVENUS BAY

Oo 20 40 60 8010

EE a

KILOMETRES

PT GERMEIN

PT _ PIRIE

FIG 5

PT BROUGHTON

WALLAROO

PT WAKEFIELD

PRICES Yric 6

PT PRIME

PT GAWLER

FIG g@PT ADELAIDE

@ ADELAIDE

Fig. 1. Distribution of stands of Avicennia marina in South Australia, showing locations of stands

mapped in Figs 2-8.

They contain a lower percentage of clay sized

particles, partly attributable to the winnowing

effect of the tides; the tidal waters suspend the

the fine material and whilst in suspension it is

carried to the backwaters of the mangrove com-

munity where it is deposited.

The “West Coast” sediments consist

dominantly of quartz, carbonate clay, algal and

shell fragments, foraminifera and diatoms.

Most of the quartz is rounded to sub-rounded,

with some particles subangular (Powers 1953).

The percentage of organic carbon, mainly

decomposing mangrove and seagrass leaves,

varies greatly with location within a given man-

grove community, with depth in the sediment

and between communities. Minor constituents

are echinoid spines, aragonite rosettes and

* needles, sponge spicules (silica), radiolarian

tests (opaline silica) and minerals from the

hinterland (e.g. amphiboles and feldspars).

Due to the low-energy depositional environ-

ment in which these sediments are found, it is

deduced that the quartz is introduced from the

extensive beaches and sand-dunes in or near

the mangrove stands. These are, or were,

environments of much higher energy. The

quartz-grains are introduced into the mangrove

community either by long-shore drift in the

beach environment and then via tidal water,

or else by saltation from the surrounding sand-

dunes.

Other constituents in the sediments find their

way into the mangrove communities via tidal

channels, or live and die on the sediments and

hence are incorporated (e.g. diatoms).

(2) The boundstone sediments found in Spen-

cer Gulf north and east of Whyalla and in Gulf

St Vincent, will be referred to as the “Gulf”

type. The particles are predominantly of clay

to fine silt size, although there is local varia-

bility. These sediments too are covered by an

algal mat. The major constituents are similar

to those of the ‘““West Coast” sediments. How-

ever, the quartz grains are rounded to sub-

rounded and clear, whereas in “West Coast”

sediments they usually have rutile and tourma-

line needle inclusions. Some minor constituents,

especially radiolarian tests and sponge spicules,

A.J, BUTLER, A. M. DEPERS, 8. C. McKILLUP and D, P. THOMAS

DENIAL BAY?

NARDIA LANDING

Fig. 3, Franklin Harbour, Spencer Gulf,

are absent. The lack of minerals such as amphi-

boles and feldspars, like the lack of inclusions

in quartz grains, results from the absence of

significant metamorphic rock sources in the

hinterland of the ‘“\Gulf* areas. By comparison,

the hinterland of the “West Coast” areas con-

lains a variety of metamorphic

(Glaessner & Parkin 1958).

Areas studied in mere detail

(1) At Port Gawler in Gulf St Vincent

(Fig. 7), very rapid sedimentation, with a con-

sequent relative drop in sea-level, has left a

very thick pile of sediment in which the present

well-established mangroves grow. Deep tidal

channels supply the area with seawater. Near

the present beach a new mangrove colony has

become established, and from a series of aerial

photographs it is clear that the lower-tide

region of the beach has been progressively

colonized within the last 10 years.

Statistics from the smear slide results (Table

|) show certain trends from colonizing man-

groves to mature stands. Generally, quartz con-

lent decreases as does the grain size of quartz

(from medium sand size to fine silt size), car-

bonate clay increases, the contents of algal and

shell fragments decrease along with their grain

size (medium sand size to medium silt size),

and organic carbon increases. Clearly the trends

are not statistically significant in several cases,

because of wide variability. However, it was

sources

MANGROVE FORESTS IN SOUTH AUSTRALIA 39

Fig. 4. Northern Spencer Gulf.

not possible to obtain more data and it seems

unwarranted to carry out a more complete

statistical analysis of these data. The apparent

trends suggest that as mangroves begin to

colonize, usually in a shell-grit grainstone of

fine sand size, the stabilizing effect of the trees

and pneumatophores allows sedimentation of

much smaller particles to commence. The final

result is an algal-covered fine-grained sediment.

The depositional texture of the sediments

changes from grainstone to wackestone-pack-

stone then to boundstone. A major factor in

these changes would appear to be the activity

of species of the blue green alga Oscillateria

and the golden brown alga Vaucheria.

fy werure

> SAND DUNES

er"

Lt ARK

Fig. 6. Northern Gulf St Vincent.

All of the mangrove sediments are under-

lain by a coarse grainstone composed of shell

fragments from gastropods, bivalves and

forams. The influence of this layer is not

known, but we suggest that it is important in

the growth of the mangroves. Being a coarse

40 A. J, BUTLER, A. M. DEPERS, S. C. McKILLUP and D. P. THOMAS

TABLE 1

Statistics from examination of smear slides of sediments from S.A, mangroves

Port GAWLER

Type of No. of Algal + shell Org

colony samples Quartz Clay fragments Forams Aragonite Cc Diatoms Others

Juveniles up 10 Mean 37.00 14.90 42.30 1.50 0.20 2.70 1.40 0.00

to 50 cm high 8.D, 8.13 7.59 11.18 0.71 0.42 3.05 1.43 0.00

Saplings up to 5 Mean 37.00 8.60 45.60 4.00 2.00 2.01 0,80 0.00

2m high S.D. 13.51 9,24 2.88 3.67 1.73 2.73 0.84 0.00

Mature trees 12 Mean 30.00 28.33 24.58 1,27 0.42 13,75 2.09 0.00

2 m and more S.D. 13.98 18.56 13.65 0.37 0.66 9.32 2.02 0.00

DAVENPORT CREEK-CEDUNA

Type of No. of Algal +- shell Org

colony samples Quartz Clay fragments Forams Aragonite Cc Diatoms Others

Juveniles up 2 Mean — 5.00 27.50 59.00 1.00 0.51 3.50 3.50 0.00

to 50 cm S.D. 0.00 3.54 2.83 0.00 0.70 2.12 2,12 0.00

Trees 2m 27 Mean 25.00 22.74 34.45 1,82 1.12 12.22 1,78 1.30

and more S.D. 22.63 15.67 27.46 1.84 1,74 12,92 1.88 2.30

—_—_ eee

layer it is also very porous and permeable, so

that seawater can move through it freely. This

may be important in the functioning of the soil

ecosystem and in the nutrient-balance of the

trees,

Within the mangrove stands deep burrowing

by the crab Helograpsus haswellianus (White-

legge) and other organisms, especially various

polychaete worms, bioturbate the sediment

extensively. The result is that the algal lamina-

tions are destroyed and a mottled texture is

commonly found in cross-sections of the sedi-

ment. Through these crab holes the water is

able to permeate.

(2) In the Davenport Creek area near Ceduna

(Fig. 2) extensive, mobile, carbonate-rich sand-

dunes are present. These are very coarse-

grained (medium sand to coarse sand size) and

are composed of about 95% shell fragments

and 5% rounded to sub-rounded medium sand

size quartz. The prevailing southwesterly wind

blows the dunes directly onto the mangrove

community to the north-east of the beach, and

the mangroves are gradually dying near the

sand-dunes due to the “blanketing” movement

of the dunes. Not far from its mouth the tidal

channel called “Davenport Creek” has cut

through the highly organic mud of a former

mangrove forest which appears to have been

killed by saltation with marine sand.

The mangrove sediments from Davenport

Creek are rich in carbonate and quartz (Table

1). Except for the variation in organic carbon

the changes from colonizing to mature stands

are less clear, but we assume that a process like

that described in (1) above occurs here. The

sediments here are much coarser-grained than

at Port Gawler, and again a coarse grainstone

underlies the mangrove sediments.

In the tidal channels, extensive burrowing

by polychaete worms has left a mound-covered

terrace, not seen in any of the other areas

studied, but the crab Helograpsus is rare; this

seems to be general for “West Coast” sedi-

ments.

Within the mangroves are to be found a

series of stranded beach ridges (probably

cheniers). Since these represent previous shore-

lines (the ridges are built up during storms and

contain extensive beds of dead Posidonia sp.)

we infer that this area has been subjected to

a number of sea-level changes. There are three

such ridges visible and probably several more

buried under the dunes. They are grainstones

composed of coarse gastropod-bivalve-shell

fragments, now overlain by a soil profile sup-

porting small bushes and grasses. Similar

stranded beach ridges are found in the other

Mangrove areas (e.g. Yatala Harbour, Port

Pirie, Port Gawler).

(3) Yatala Harbour, south of Red Cliff Point

in Spencer Gulf (Fig. 4) was studied in detail

prior to this survey.6 This area exemplifies the

“Gulf” type of sediment described above.

ee eSeeeeSFSFSsSSSSSSmmsFhFhFesesesese

8 Depers, A. M. (1974).—Sedimentary facies at Yatala Harbour and a geochemical comparison with

Port Pirie sediments, Spencer Gulf, S.A. B.Sc. (Hons.) thesis, University of Adelaide. Unpublished.

MANGROVE FORESTS IN SOUTH AUSTRALIA 4)

a Mature

COLONISING

Ez, DEAD

i CONSERVATION PARK

SAND DUNES

hip. 7. Middle Beach und Port Gawler, Gulf St

Vincent,

Fig. 8. Port Adelaide, Gulf St Vincent.

A thin sequence of Holocene sediments

occurs in a prograding carbonate shoreline. To

seaward, the area is flanked by shell-grit grain-

stones and Heterozostera seagrass banks; in

deeper water are banks of Posidania seagrass.

On the landward side are extensive supratidal

carbonate lagoons and samphires.

The mangroves grow in grey sediments con-

sisting of from 60% to 100% carbonate mud

as estimated by areas on smear slides. Terri-

genous clay can be as high as 35% and organic

matter up to 20%. Smear slides of the sedi-

ments show that the carbonate is dominantly

precipitated calcite rhombs (45-87%) with

minor aragonite rosettes (<1%) and some

dolomite rhombs (<1%). The remainder of

the carbonate fraction (6—30%) consists of

mainly algal fragments and smaller amounts of

foraminifera, echinoid spines and bryozoan

fragments. Most of the particles are clay to

medium silt sized. Quartz grains are generally

rounded to sub-rounded, with the occasional

ungular grain present. Similar sediments have

also been found at Port Pirie.

The algal mats which grow on bare mud and

hold the surface sediments together are par-

tially laminated in section. Subaerial exposure

causes them to crack and curl, The mud crab

Helograpsus plays an important role in_ bio-

turbating the sediment. The tidal channels are

lurge and usually contuin water at low tide.

Some of the channels are very rich in decaying

organic matter, especially masses of dead leaves

of Poxidonia sp., and smell strongly of hydro-

gen sulphide. In one channel, 12 cm of solid

42 A, J. BUTLER, A. M. DEPERS, 8. ©. McKILLUP and BD. P, THOMAS

black peat was found, The tidal channels com-

Monly contain 4 channel-lag wackestone, Cons

sisting of a concentration of bivalve and gas-

tropod shells.

Extine Mangrave Stands

Extinct mangrove stands have been recorded

al Cilenela (Cotton 1949), at Webb Beach near

Parham, and at Davenport Creek (above). We

examined only one, Baker's Creek at Webb

Beach, in any detail, Here we fouind two well

preserved platforms of sediment bound by algal

mats, one overlying the other with about 15 em

of shell fragment grainstone between them,

Dead tree slumps Jie within the platforms.

Smear slides showed that the sediments gre

similar in composition to those found in other

areas, Both are carbomate-rich; they are

extremely cich in organic carbon, and this is

true of all the extinct stands we have seen.

Prebably the algal layer protects the sediment

and, because it does not break apart, decam-

position is extremely slow,

There is some evidence of considerable sea

level changes in this area (S, Curr, pers.

comm.; Ward & Jessup 1965) and the presence

of two mud platforms one above the other may

also indicate such changes, But the evidence

from the extinct mangrove platforms is not

conclusive, because Mangroves can live over

a range of altitudes in the interlidal zouc (But-

jer unpubl, ),

To the south of the extinct stand is a small

stand of living mangroves in a fidal channel

relatively protected fram the dominant sen

swell,

The deaths of the two Webb Beiich stands

seem likely to be che result of encreachnenc

of shell-grit facies over the mangrove bound-

stone. Extensive dunes on the landward sicte

also Could have had an effect on the mangroves.

Sedimentary Dynamics—Conclusion

Within the mangrove community, a dynamic

relationship exists between the sedinient and the

plants growing in and on it, Some sediment

must be present for colonization by the plants,

but once they have colonized rootlets, pneu-

matophores and algal mats stabilize the sedi-

ment; algal mats Facilitate the entrapment aud

precipitation ef carbonate grains (Buthurst

L971: Carlen (974; Gebelein 1969: Neumann

et al, 1970; Seoffin 1970), This relatlonship is

a delicate one, a4 Gach supports the other,

It is clear that the persistence of a mangrove

forest Jepends greatly on sedimetitary pro-

cesses, There are places (2g. Part Clinton)

where erosion of the sediment is leaving the

trees Withoul Support and they are dying with:

out replacement. In others (e.g. part of Daven-

port Creck) saltation or encroachment by

dunes is Killing the forests; it appears that in

sume such cases mature trees survive but seed-

lings eannot establish, so that the forest

eventually disappears, We cannot cite clear

cases OF the opposite, where death of man-

Broves results in erosion of the sediment, but

is possible,

EXTINCT MANGROVE FORESTS, AND

NOTES ON HISTORY

We have found sheets of mangrove mud con-

taining dead stumips at Davenport Creek, west

of Ceduna, and al Webb Beach, Gulf St Vin-

cemt. Cotton (194%) reported such a mudflat

briefly exposed at Glenelg, and suggested “from

fauital studies” that mangroves existed “until a

comparatively short time ago” as far south as

Port Noarfunga-

Cotton postulated that mangroves are

gradually relreating northwards in Giulf St

Vincent. Independently of that, he also surge

fested because some of the mollusc shelly in

the mud were larger than present-day specl-

mens that when the forest at Glenelg Auurished

conditions were a little warmer than at present,

We have found, in conversation, a poptilar

local belief thal mangroves are retreating

northwards because conditions are becoming

cooler, so that they cannot survave further

south. We do wot think this is the best inter-

‘pretation of the facts.

The distribution of the known extinct stands

&% nol consistent wilh uw simple retreal up the

Guilt. Cotton postulated that the mungrove

forest at Glenelg Was contemporary with the

red sund-dunes which lay behind the recent,

White dunes and that it fourished 1000-3000

years ngo, finally being killed by “sand-silting™,

As noted atrove. we think that the death or

reduction of forests in several ureas cau be

explained mm terms of sedimentary proccsses-—

encroachment by dines, saltation or erosion,

It is beyond the scope of this paper to dis-

cuss broad pallerns of changes in sea level,

climate, wind, wave and current pallerns and

coastal morphology into which these cases

might fit, but we think im clearly not u simple

case of mangroves retreating north as the

climate Cools. That would be Inconsistent with

the tact that they @ecur much further soulh at

Westernport Bay, Victoria, Ruther, we see mon-

graves us living i very dynamic sediments with

MANGROVE PORESTS IN SOUTH ALISTRALIA 43

siltation, saltation and erosion taking place in

different stands according to the local condi-

tions at this time.

We have also encountered (“popular belief”

that mangroves were much more extensive in

South Australia oo the arrival of while settlers

than they ure now, for exumple, that they

occupied the beach at Glenelg and occurred at

Port Noarlunga. It is said they were cleared

because people feared malaria. This belief may

be accounted for by a misreading of Cotton's

(1949) paper, by a misidentification of the

plants, or by its being true, We decided to find

what we could from the writings of early

seutlers.

Cotton (1949) noted thut “a sketeh of [Lhe

Cilenele) area by Colonel Light in about 1835

depicts the beach pretly well as aft present’,

Light (1839: also quoted by Bull 1884) gave

a bearing and latitude which clearly placed him

off Ouler Harbour when he noted “to the north-

ward and eustward mangroves growing Lo the

water's edge”. These must have been the man-

eroves of Torrens Island, St Kilda and north-

wurds tu Port Gawler, At another time sailing

northward long Holdfast Bay he recorded

“hurd sandy beach the whole way",

Certainly extensive areas of tidal swamps

in the Port Adelaide revion und to the south

have heen filled 7 Much of this has been dene

within reeent memory, bul the beach at Glenelg

Way not creattd in this way; the mangroves

there appear to have been buried more than 130

veurs ago. We cannol be surc about Port

Noarlunga,

COMMUNITIES GF ORGANISMS IN

S.A. MANGROVE SWAMPS

Lists of Mory and fauna collected will be

presented elsewhere but the following remarks

bused on those lists are worth recording here,

We [our no trends across the State that

would be ipteresting, biogeographieully: rather,

any of the species could be expected if con-

ditions Were appropriate, This is not surprising,

ug all the mangrove forests Fall within the Flin-

dersian Province defined by Womersley &

Bdimoands (L958). By definition, A vicennia Was

always present; mast commonly there was an

extensive Saltmarsh to landward, dominated hy

Salieornia or with Salicornia and Arthree-

yeni codominunt, snd bare mud Mats or sea-

aiass beds usually lay to seaward of the man-

proves, The fauna vaned between sites,

Ij doves not appear that there is a unique

assemblage of organisms which might be called

a “typical South Australian mangrove com-

munity", Le. an assemblage of species which

nearly always oo¢ur together. Even the animal

species most commonly associated with man-

proves, such as the crab Aclograpius lNay-

welliaays unc the snail Bombiciunt auratus,

are somelimes rare Or absent, and they do oecur

commonty in the absence of mangroves,

These observations mdicate that the distribu-

lion and abundance of species in the tidal

swamps depends primarily on the requirements

of the individual species, and on Factors such as

substrate type and height of scdiment above

mean sea level, rather than on the presence of

the mangrove itsell. Uhis is in yeneral agree-

ment with the conclusions of Clarke & Hannon

(1971) on plant zonation in tidal swamps. in

the Sydney district, and of Macnae (1968) on

the distribution of animals within mangrove

forests.

All aureus of owngroves in South Australia

have features in common, because A virerinia

pueine has certain requirements (Clarke &

Hannon 1967, 1964, L970, 1971; Farrell &

Ashton 1974"), But the mangrove is Aexible in

ils requirements and ip mast respects if appears

tu have wide tolerances. This is reflected in the

variable communities of organisms that live

wilh it,

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T.P. (1970).—The composition, structure

and erodability of subtidal mats, Abaco.

Bahamas. J. Sed. Petrol. 40, 273-297.

Powers, M. C. (1953).—A new roundness scale

for sedimentary particles. J, Sed. Petrol. 23,

117-119,

ScorFin, T, P. (1970),—The trapping and binding

of subtidal carbonate sediments by marine

vegetation in Bimini Lagoon, Bahamas. J.

Sed. Petrol, 40, 249-273.

THom., B. G. (1967).—Mangrove ecology and del-

taic geomorphology: Tabasco, Mexico, J,

Ecol, 55, 301-343.

Womers.ey, H. B.S. & EpMonpbs, S. J. (1958).—

A general account of the intertidal ecology of

South Australian coasts, Aust. J. mar. fresh-

wat. Res. 9(2), 217-260.

Warp, W. T. & Jessup, R. W. (1965)—Changes

of sea-level in southern Australia, Nature 205,

791-792_

THE AGE OF AYERS ROCK AND THE OLGAS, CENTRAL AUSTRALIA

BY C. R. TWINDALE AND W. K. HARRIS

Summary

Large areas of the desert plains of the southwestern part of the Amadeus Basin, central Australia,

are underlain by Cainozoic terrestrial deposits which rest on an irregular land surface eroded in

folded Proterozoic and Cambrian strata. This pre Cainozoic surface embraces the broad massifs

now surmounted by the Olgas and Ayers Rock, and the broad bedrock depression separating the

two, as well as many minor valleys and hills. The main depression is part of an ancient valley

system which drained to the southwest.

THE AGE OF AYERS ROCK AND THE OLGAS, CENTRAL AUSTRALIA

by C. R, Twipave* and W, K. Harrist

Swinmary

Twroane, ©. R. & Harris, W. K. (1977). —The age of Ayers Rock and the Olgas, central Aus-

tralia. Trany. R. Soe. §. Ause. W1(1), 45-50, 28 February, 1977.

Large areas of the desert plains of the southwestern part of the Amadeus Basin. central

Australia, are underlain by Cainozoic terrestrial deposits which rest on an irregular land surface

edded in folded Proterozoic and Cambrian strala, This pre Cainozoic surface embraces the

broad massife now surmounted by the Olgas and Ayers Rock, ahd the broud bedrock depression

sepiraling the Lwo, ws Well as many minor valleys and hills. The main depression is part of

am uncient valley systern which drained to the southwest.

The lower parts of the old landscape have been buried by lacustrine, alluvial und acolian

sediments. The age of the basal Cainyzoic strata deposited in the lower parts of the old relief

provides a minimum age for the erosional surface including the upland precursors of the

comlemporiury inselbergs, though it is emphnsised that the present steep-sided morphology of

these bornhardts is a comparatively recent development.

Palynological evidence from carbonaceous sediments lying directly on the old lund surface

indicates « Middle Paleocene age. Thus the Olgas and Ayers Rock viewed as upland masses

canoot be younger than this, and probably evolved during the later Cretaceous, They were

Upland remnunts standing above a late Cretaceous surface of low relief that extended over

wide areas of central Australia, the northern Flinders Ranges and northwest Queensland.

Introduction

Since their discovery by Buropeans just over

a hundred years ago (Gosse 1874, Giles 1875,

1889), Ayers Rock and the Olgas complex

have fascinated earth scientists and the lay

public alike. Both uplands are bornhardts or

domed inselbergs (Figs 1 and 2). Ayers Rock

is a single, isolated, monolithic dome thal

stands 877 m above sealevel and 340-350 m

above the surrounding desert plains (Fig. 3),

The Olgas, on the other hand, consist af a

group of topographic domes of varied mor-

pholagy, some being hemispherical, others

towers with rounded crests, and yet others are

the pitons emoussées or bevelled towers. of the

French literature (¢.g. Mainguet 1972), The

highest, Mt Olga itself, rises 1,069 m ahove

sealevel and SQ0-550 m above the plain level.

Both inselbergs are noteworthy by virtue of

their size alone, but they gain further dramatic

impact from their splendid isolation, from the

steepness and intricate etching of their bound-

ing slopes and their consequent abrupt rise

from the surrounding essentially flat desert

plains, and from the marked contrast between

the latter and these towering red massifs.

From a geomorphological point of view the

bornhardts are unusual in thal they are both

eroded from folded sedimentary formations,

Bornhardts developed in granitic rocks are

fairly commonplace and Widely distributed

(see Wilhelmy 1958; Twidale 1971, 1976a)

but Ayers Rock has been sculptured from very

steeply dipping Cambrian arkose, and the

Olgas group from the moderately dipping, mas-

sive and coarse Mount Currie Conglomerate of

the same age (Tate & Watt 1896, Jokilk 1952,

Forman 1966, Wells ct ul, 1970), deposited in

the piedmont of the Musgrave Block, and with-

in the intracratonic Amadeus Basin (Fig. 2).

The processes responsible for shaping these

spectacular Jandforms have been discussed else-

where (Twidale 1977, Twidale & Bourne

1977), However the age of the inselbergs as

topographic forms, as ef any landforms, is

critical, for this provides a framework for the

climatic and tectonic conditions under which

the features evolved, and an essential perspec-

* Depariment of Geouraphy, University of Adelaide, North Tce, Adelaide. S. Aust. 5000

| Geolovical Survey of South Australin, Box 191, Eastwood, S. Aust. 5063,

ott 4.

oe 7 ee

Nie ———

Ce Ms ae

TEP

tf a Fe ake

Le Sarl

C. R. TWIDALE & W. K. HARRIS

ee a

. sme

— =

woe allt:

Fig. 1. Oblique aerial view from the southeast of Ayers Rock (foreground) and the Olgas in the dis-

tance. Note the steep dip of the strata in Ayers Rock, the flat desert plains, and the approximate

location of bore G 394855-44 (circled). (S. Aust. Tourist Bureau.)

tive when problems of survival are contem-

plated (Twidale 1976b).

Geomorphological setting

Drilling has confirmed inferences drawn

from their morphology, namely that the plains

that surround the two residuals are deposi-

tional. They are underlain by Cainozoic

sequences comprising lacustrine, alluvial and

aeolian beds, the thickness of which varies.

Though the plains surface slopes gently down

from west to east and displays only minor local

relief amplitude due to the development of

dune ridges on the one hand and playa depres-

sions on the other, the surface underlying the

terrestrial deposits and eroded in folded Pro-

terozoic and Cambrian beds is irregular. To

the west and north of Ayers Rock, for instance,

fresh arkose is nowhere more than a few

metres beneath the surface (Fig. 3) and there

are indeed outcrops of fresh arkose not only in

the northern piedmont zone but also 700-800

m west of the base of the Rock. Immediately

to the south of the inselberg the Cainozoic

cover is more substantial (20-35 m) but about

300 m south of the cliff line it suddenly

thickens as it passes over a (?) fault-line scarp

separating the arkose from the older Protero-

zoic strata. The surface cut in the older rocks

is here irregular, the Cainozoic being com-

monly 70-90 m thick but bores have pene-

trated through 181 m without entering the Pro-

terozoic (Fig. 3).

To the west of the Olgas the Cainozoic cover

is thin, and there are many outcrops of the

Mount Currie Conglomerate in the form of low

domes and platforms. To the east the younger

sequence is thin near the dome complex (Fig.

3) but its thickness increases to the east where

it buries a broad depression which is more

than 100 m deep and is believed to be part of

an old valley system draining to the southwest

(R. E. Read, pers. comm.). This depression

separates the broad platforms or massifs sur-

mounted by the two bornhardts. Thus the

Cainozoic sequence has inundated the lower

parts of an irregular land surface cut in the

Proterozoic and Cambrian sediments. As there

was a broad valley depression prior to their

deposition, it follows that the two adjacent

higher massifs also predate the deposition; and

though there is reason to believe that neither

AGE OF AYERS ROCK 47

ARUNTA BLOCK

“COONNETT RA

AMa

- 2 = AL Nea EUS

L. Amadeus

Ayers Rock

ALICE SPRING

PEDIRKA

AMt Conner

BASIN

DESERT

EVERARD RA

®)QODNADATTA

ARCKARINGAS

Fig. 2, Generalised tectonic map of southwestern part of the Northern Territory showing locations men-

tioned in text.

Ayers Rock nor the Olgas were then as steep

sided and of such dramatic appearance as they

now are (Twidale 1977), there must have been

ancestral uplands in the same locations. Thus

the age of the basal Cainozoic provides a mini-

mum age for these residuals. Bremer (1965)

surmised that Ayers Rock is of some antiquily

but was unable to cite any specific evidence on

the point.

Age of the basal tertiary strata

According to Wells et al. (1970) the Caino-

zoic sequence of the southwestern part of the

Amadeus Basin includes lacustrine strata of

early Tertiary age, but they were unable 10 be

more exact. In a general way, however, their

estimate is borne out by the fact that some of

the Cainozoic beds have been duricrusted,

both ferruginous (laterite) and siliceous (sil-

crete) carapaces having been developed in the

course of prolonged deep weathering. The sil-

erete of central Australia is generally con-

sidered to have formed during the early-middle

Tertiary, reaching its climax in the Miocene

(Wopfner & Twidale 1967, Wopfner et al.

1974) so that the lake sediments beneath these

duricrusts appear in general to be of early

Tertiary age.

Fortunately it is now possible to give a more

precise age for the basal Tertiary sediments.

Lignites resting directly on the old land surface

eroded in the folded Proterozoic and Cam-

brian rocks occur in four bores near Ayers

Rock and the Olgas (Fig, 3), A sample (S.A.

Dept. Mines Sample No. 84065) of one lignite

from bore G394855-44 located in the broad

vulley between the Olgas and Ayers Rock (Fig.

3), and from a depth of 81-84 m beneath the

surface, contains a diverse palynomorph assem-

blage, which includes the following strati-

graphically useful species: Anacolosidites acu-

tullus Cookson & Pike, Beaupreaidites elegansi-

formis Cookson, 8. verrucosuy Cookson,

Cumarozenosporites bullatus Harris, Cyathi-

48 C. R. TWIDALE & W. K. HARRIS

} “

{ 2

{ oa

| 7 ee

| A i ("=

. 3 ant

4 a = — < aa 77 266 an \ /

; 39 1 {

; J iy y 066 4a oy \ ost

“5 4 4 Py “

a 3B an - y " a ane

Aa s. 7 wr #63 . a a LL pene Fi

qe a 2 oaeousise Bt Lt

EO aALS pak OEP ne! "

ey a9 \

PT at ral . sao. i f ‘ \

etal SoD = - Pol oe a 936 ok 06 | os i

——J. C 2 pamilse- 43 az

s=yen * = se * a ns

cL Sy “a ’

st LIT 39638 570.

THE OLGAS > ‘he E pi

1 ae 4

‘ ae a

: =

ha * bore depth to bedrock

a © bore depth of bore

oe \ e + bore lignite within sequence

f+ bore lignite dated

tracks

— — interred tauit

— — park. boundary

Fig. 3. Drilling sites in the Ayers Rock and Olgas region. Depth to Proterozoic or Cambrian bedrock in

metres. Bores with lignite starred. Position of G 394855-44 shown. (After Water Resources

Branch, Dept of Northern Territory, Alice Springs).

b p e

Fig. 4. Spores from carbonaceous sediment from

depth of 81-84 m in bore G 394855-44.

a Latrobosporites ohaiensis (Couper)

Stover, b Quadraplanus brossius Stover,

¢ Gambierina rudata Stover, d Tetracol-

porites verrucosus Stover, e Tricolpites

reticulatus Cookson. All x500.

dites splendens Harris, Ephedripites notensis

Cookson, Gambierina rudata Stover,

Gephyrapollenites wahooensis Stover, Herko-

sporites elliottii Stover, Krauselisporites papil-

latus Harris, Proteacidites angulatus Stover, P.

fromensis Harris, P. kopiensis Harris, Quadra-

planus brossius Stover, Tetracolporites verruco-

sus Stover and Tricolpites reticulatus Cookson.

These species and other elements not reported

herein have features in common with southern

Australian Paleocene spore-pollen assemblages.

Biostratigraphic schemes for the early Tertiary

have been proposed for the Gippsland Basin

by Stover & Partridge (1973) and for the

Otway Basin by Harris (1971). More recently

these schemes have been extended from the

coastal basins to the continental basins of South

Australia (Wopfner et al. 1974). Paleocene

sediments of the Eyre Formation in the Ero-

manga Basin contain palynomorph assemblages

closely similar to that described here.

The absence of the nominate zone species

which characterise the biostratigraphic zones of

Harris (1971) and Stover & Partridge (1973)

does not preclude correlation and whilst there

are significant differences between this assem-

blage and those of both the Otway and Gipps-

land basins, there are, nevertheless, many ele-

ments in common. In particular Camarozono-

sporites bullatus, Gambierina rudata and

Krauselisporites papillatus are late Cretaceous

to Middle Paleocene forms, Herkosporites

elliottii and Proteacidites angulatus commence

their stratigraphic range in the Middle Paleo-

cene, Tetracolporites verrucosus is an Early to

Middle Paleocene form and Quadraplanus

brossius is restricted to the Early Paleocene of

the Gippsland Basin. Anacolosidites acutullus,

AGE OF AYERS ROCK 44

Beaupreaidites elegansiformis and B, verruco-

wus appear luter in the Paleocene or Early

Eocene of the Gippsland Basin but A, eufullus

occurs in the Middle to Late Paleocene of the

Otway Basin

The weight of evidence thua favours a

Middle Paleocene age for the assemblage which

correlates with the Garnbhicrina edwardsii zone

in the Otway Basin and the Lygistepallenites

halme7 zone in the Gippsland Basin, This is

supported by the absence of the younger Mal-

vecipellis diversus Harris and the older

elements such as Pricelpitey longuy Stover, The

assemblage differs from the correlatives of the

Ganbierina edwardsi7 zone in the Eyre Forma

tion in the very rare occurrence of Notho/agi-

difes spp, and absence of Australopollis

obyeurus Harris. Kemp’s (1976) assemblages

from Central Australia are younger and quite

distinct, There are na indications of marine

influence and abundance of conifer pollen of

the Microcachrys and Podacarpus types indi-

cates high rainfall.

A systematic account and a full discussion

on the implication of this assemblage wil! be

presented elsewhere.

Conclusion

As the lignites resting on the old land sur-

face cut in Proterozoic and Cambrian sedi-

ments are of Paleocene age the old landscape

including the precursors of the present Ayers

Rock and the Olgas cannot be younger than

Palcozene in age. They may thus be part of the

late Cretaceous land surface described from

other parts of central Australia (Mabbutt 1965)

and from other areas marginal to the Great

Artesian Basin (Woodard 1955, Twidale 1956,

1966, 1969, 1976b, Twidale, Bourne & Smith

1976), and contemporary with the crests of the

higher granite inselbergs on Eyre Penjnsuly

(Twidale & Bourfe 1975).

Acknowledgments

The writers are indebted to Mr R. E. Read,

Water Resources Branch, Department of the

Northern Territory. Alice Springs, for supply-

ing the all-important sample of lignite, and for

giving us the benefit of his extensive expenence

and knowledge of the subsurface geology of

the southwest of the Amadeus Basin, The

paper is published by Kind permission of the

Director of Mines, South Australia,

References

Bremen, H, (1965).—Ayers Rock, ein Beispicl

fiir klimagenetische Merphologie. 2. Geo

morph. 9, 249-184.

Forman, D. |, (1966),—The geology of the soulh-

weslern margin of {Ie Anradeus Basin, cen-

tral Australia, Repr Bar Mine Res. Ceal-

Geophys. Aust. 87.

Gipes, FJ, (1875)—"Geowraphie travels m cenr

tral Australia trom 1872 ta 1874," (MeCar-

ron, Bire: Melbourne.)

Gues, EF, J. (1889) — Australia

‘Traversed.” (Sampson Low-: Landon, )

Gosse, W. C. (1874).—Report and diary of Mr

W. C. Gosse's central and western exploring

expedition. 1873, Parl, Pap. ¥, Aust. 48.

Hannis, W. K. (1971).—Tertiary stratigraphic

palynology, Otway Basin, én H, Woptner &

I. Douglas (Eds), “The Otway Basin of south-

cast Austrailia’, 67-87. Spec, Bull. Geol. Survs.

S. Aust. & Vier.

Jokiic, G. PF. (1952) —Geolagical reconnaissance

of 30uth-western portion of Northem Terri-

tory. Repr Hur, Min. Resour. Geal. Geophis.

Anse,

Kempe, BE. M_ (1976). Early Tertiary patlen trom

Nupperby, central Austraha. J, Bur Min

Resor Geel, Geoplivs. Aust. 1, (09-114,

Massury, J. A, (1965}—The weathered land sur-

Sepik centval Australia 2. Geomorph, 9.

Matnuuer, ML (1972)—"Le Modelét des Cis.7

MInstit. Geogr, Natl, Paris.)

Twice

Stover, L. & & Parrringe, A. BD. (1973).—Ter-

liary and Late Cretaceous spores and pollen

from the Gippsland Basin, southeastern Aus-

tralia, Proc, R, Soc. Vict. 88, 237-286,

Tare. R. & Wart SF. A, (1896).—General geology.

fn B. Spencer (Ed.), “Report on the Work of

the Horn Scientific Expedrtion to Central Aus-

tralia”, 26-75, (Melville. Mullen & Slade-

London),

TWIDALE, C. R. (1956).--Chronology of denida-

tion in north-west Queensland. Bull. Geol,

Sac. Aust, 67, 867-882,

Twipace, C, R. (1966).—Chronology of demuda-

tion in the southern Flinders Ranges, South

Australia. Trans. Ro Soe. S. Apst. 90, 3-28,

Twinstx, C. R, (1969),—Geomarphaology of the

Flinders Range, 57-137. Ja DO W. PL Corben

(Ed.). “Natural History of the Fiindere

Range”. (Publ. Libr: Adelside, }

Twiiace, ©. R. (1971) —"Structural Landforms,”

{A,N.U, Press: Canberra_}

Twibate, C. R, (197 6a).—'Anatysis of tanl-

forms,” (Wiley; Sydney.)

Twinare, ©. R. (1975b)—On the survival of

pialaeoforms, Aim, I. Sei, 276, 77-94,

Twipace, C. R, (1977).—On the origin of “Ayers

Rock. central Ausimilia, 2. Geavrrearph. (in

press)

Twrpare, C. RO & Bourne, I A (1975) Epi

yodic exposure of inselbergs. Geal. Soe, Aner

Bull, 86, 1,473-1,48)

Twinace, ©, R. -& Bourse, I, A, (1977),—Horn-

hardts developed in scdimenmey rocks trom

ceritral Australia. Geepr Rev. (submitted).

VOL. 101, PARTS 2, 3, 4 31 MAY, 1977

TRANSACTIONS OF THE

ROYAL SOCIETY

OF SOUTH AUSTRALIA

INCORPORATED

CONTENTS

Mawson, P.M. Revision of the genus Macropostrongylus and descriptions of

three new genera: Popovastrongylus, Dorcopsinema, and Arun-

delia (Nematoda: Trichonematidae) - - - - - 51

Twidale, C. R., Bourne, J. A. and Twidale, N. Shore platforms and sealevel

changes in the Gulfs region of South Australia - - - - 63

Buonaiuto, M. F. Revision of the composite Rea Lima bassi Tenison Woods

(Mollusca, Bivalvia) - - - - - - er i

Tyler, M. J. Pleistocene frogs from caves at Naracoorte, South Australia - 85

Butler, A. J. and Murphy, C. Distribution of introduced egy ae on Yorke

Peninsula, South Australia - - - - - 91

Houston, T. F. Nesting biology of three ailodapine bees in the subgenus Exon-

eurella Michener (Hymenoptera: Anthophoridae) - ~ - 99

PUBLISHED AND SOLD AT THE SOCIETY’S ROOMS

STATE LIBRARY BUILDING

NORTH TERRACE, ADELAIDE, S.A. 5000

REVISION OF THE GENUS MACROPOSTRONGYLUS AND

DESCRIPTIONS OF THREE NEW GENERA: POPOVASTRONGYLUS,

DORCOPSINEMA, AND ARUNDELIA (NEMATODA:

TRICHONEMATIDAE)

BY PATRICIA M. MAWSON

Summary

The genus Macropostrongylus is redefined and revised. Species retained in the genus are M.

macropostrongylus, M. macrostomata, M. yorkei, M. lesouefi, and M. irma. New genera are

proposed: Popovastrongylus for M. wallabiae, M. pearsoni, and M. irma n.sp.; Dorcopsinema for

M. dorcopsis; Arundelia for M. dissimilis. M. australis, M. cornutus, and M. minor are referred to

Cloacina; M. labiatus to Zoniolaimus, and M. baylisi to Macropostrongyloides. The genus

Gelanostrongylus is suppressed. Cloacina daveyi nom.nov. is proposed for C. australis Johnston &

Mawson nec C. australis (Yorke & Maplestone).

REVISION OF THE GENUS MACROPOSTRONGYLUS AND DESCRIPTIONS OF

THREE NEW GENERA: POPOVASTRONGYLUS, DORCOPSINEMA, AND

ARUNDELIA (NEMATODA: TRICHONEMATIDAE)

by ParriciA M. Mawson*

Summary

Mawson, P. M. (1977) Revision of the genus Mucropostrongylus and descriptions of three

new genera: Popovastrangylus, Dorcopsinema, and Arundelia (Nematoda: Trichonema-

tidae), rans, R, Soe. S. Aust. 101(2), 51-62, 31 May, 1977.

The genus Macropostroneylus is redefined and revised, Species retained in the genus are

M. macropostrotieylus, M. macrostoma, M. yorkei, M, lesouefi, and M. irma, New genera are

proposed: Pepovastrongylus lor M. wallabiae, M, pearsoni, and M. irma 0.8.5 Dorcapsinema

for M. dorcepsis: Arundelia for M. dixsimilis. M. australis, M, cornutus, and M. miner are

referred to Cloacina; M, lahiatus to Zoniolaimus, and M. baylixi lo Macropastroagyloides, The

°. australis

genus Gelanosirongylus is suppressed, Cloacina daveyi nom.noy. is proposed for C.

Johnston & Mawson nec C. australis (Yorke & Muaplestone).

Introduction

As a result of the availability of new collec-

tions of nematodes from kangaroos and

wallabies it is now possible to revise some des-

criptions, in particular those of species

attributed to Afacrepastrongyluy Yorke &

Muplestone. Where possible comparison has

been made with type material, and a complete

revision of the genus has been undertaken. New

spectes undoubtedly await description, as the

parasites of macropod marsupials, especially

those in western and northern parts of Aus-

tralia, have seldom been collected system-

atically. It is hoped that the present work will

aid future studies.

Most measurements of specimens have been

omitted from descriptions, they are available

on request from the author or Librarian.

Historical

Although it has not been possible to examine

the type material of M. macropestrengylus and

M. australis. the species for which the genus

Macropostrongvlus was erected by Yorke &

Maplestone (1926), specimens so identified by

Baylis (1934) have been studied. The species

are re-described from this material and from

specimens from the same host (M. agilis) trom

Papua, The revised generic diagnosis is given

below, From Yorke & Maplestone’s figures M.

australis appears referable to Cloacina Linstow

1898; the specimens identified by Baylis are

certainly Cloacina sp. As C. australis (Yorke &

Maplestone) predates C. anstralis Johnston &

Mawson (1938). 4 new name must be given

to the latter, and €. daveyi is proposed.

Baylis (1927) added M. yorkei to the

genus; this is redescribed below from the type

host.

The paratype material of four new species

assigned to Macropostrongylus by Davey &

Wood (1938) has been re-examined, M. cornu-

tust and M. minor belong to Cloacina, as was

suggested from a study of the figures by John-

ston & Mawson (1939). M. labiatus belongs to

the genus Zoniolaimus, close to 2. setifera

Cobb, 1898. MM. macrostoma, partially

described below is a true Macropestrangylus.

Paratype material of M, dercopsixs Baylis,

1940, from a wallaby in New Guinea has been

examined, and is considered so different from

Macropostrongylus spp. as to necessitate the

erection of a new genus, Dorcopsinema, des-

cribed below.

Johnston & Mawson in several papers (1939,

1940) added five species: M. dissimilis, M.

inpma, M. lesouefi, M. pearsoni, and M,

wallahiae, Of these, M. dissimiilis is referred to

* Department of Zoology, University of Adelaide, North Tce, Adelaide, S. Aust. S000.

7M. cernuius has recently been described by Mawson (in press}.

52 PATRICIA M. MAWSON

Arundelia i.e. Mf. wallabiae and M. pearsont

lo Popovastrengylus n.g., and the others

retained in Macropastrongylus.

Yamacuti (1961) placed M, lasiorhini Maw-

son, 1955, from a wombat, as the type of Mac-

rapostrangyloider, Macropostrongylus bavlist

Wood, 1930, is pow transferred to this genus.

Mavrepostrengylus macropostrongylus was

described as having a leaf crown, Popova

(1952) erected 4 new genus Gelanostrongylus

for species which had been assigned to Macre-

postrongylus but in which the leaf crown is

absent, She placed the following species in the

new genius; M, mracrostoma (type species). M.

disstmilts, M_ labitatus, M. irma, M, lesowefi, M,

wallahiae, and M. dorcopsis, However, the

morphology of M, macrostema is essentially

similar to that of AZ. macropestrangylus, and

therelore Gelanastrongylus cannot stand, How-

ever jl certainly appears that there are two dis-

linct groups of species left in Macroposrrongy-

lus, even afler those belonging to other venera,

as noted above, are excepted, The species M.

macrepostrongyhis, MM. miaerasterna, M.-

lesouefi. M. yorker and M-irma form a natural

group, as do M, wallohiae ad Md, pearseni. A

new genus, Popevasironey/ns, is now proposed

for the latter group.

In Macropostrengylus the perioral cuticle

forms eight lobes, the buccal capsule is ridged

longitudinally and ends anteriorly in eight

small projections, and the oesophagus is more

or less ¢ylindrical ending in an elongate bulb,

In Popevastrengyles the perioral cuticle con-

tinues into the buceal eavity without Forming

lobes, the buccal capsule is more or fess cylin-

drica) (or oval in section) without ridges and

without anterior projections, «4nd the

oesophagus is relatively shorter, marrows sud-

denly in its posterior half, and ends in a bulb,

Macropostrongyloides ig distiaguished from

Moaeropostrougylus by the presence in the bue-

cal capsule of four large lecth, by the shape of

the oesophagus. and by the position of the

externo-dorsal ray, Which rises from the dorsal

ray. Muecropestrengylus spp. and Pepove-

stromeylus spp. occur in the stomach of the

host; Macropostrongylotdes spp. in the large

intestine.

Macropostrongylus Yorke & Maplestone

Generic diagnosis (revised } =

Trichonematidae:

Anterior end with four submedian

setigerous papillae and two lateral eleva-

fons bearing amphids; buccal capsule and

mouth more or less laterally compressed;

perioral cuticle forming eight lobes; buccal

capsule folded longitudinally into eight ridges

which, variously thickened, project anteriorly

under the cuticular lobes; oesophagus long,

stender, with oval terminal bulb. Male: dorsal

lobe of bursa longer than laterals, ventral lobes

distinct from laterals and more or less joined

ventrally; externo-dorsal rays arising separately

or with laterals, dorsal ray bifurcating before

mid-length, cach branch giving off a fateral

branch, spicules alate; gubernaculum present.

Female: Tail short, conical, vulva near anus,

Parasites of the stomach of macropod mar

supials, Type species: M, macrepostronpylis.

other species; M. macrosromea: M- yarkei) M,

lesouefi; M- irma.

In Macropostrongyles the anterior end is

simple, without a collar roll. A slightly raised

ridge surrounds the cephalic papillae. The sub-

median papillac: usually setigerous, and the

lateral elevations ate more or less prominent.

Lateral compression of the mouth and buceal

capsule, but not of the entire lateral end, is

variable. Anteriorly the eight longitudinal

ridges of the buccal capsule project as lobes

but these are covered by the corresponding

cuticular perioral lobe, formimg structures

which are apparently erectile (Fig, 2), and the

huceal capsule varies in shape with this, giving

a more or less open mouth, The lobes thus

form a sort of leaf crown, but one quite dis-

tinet in appearance from that in Cleacina and

Murshidia though both are formed from the

peribucesl and perioral cuticle,

Macropostrongylus macropostrongyhius Yorke &

Maplestone

FIGS 1-7

vena & Maplestone, 1926, from Mavropus sp,

Baylis, 1934 fi 129, from M- agilis, Qh.

Johnston & Mawson, 1939 p, 143, M1. agiliv, M,

wellsbyi, Old, 1939 p. 209, Mf, avilis, Old,

—_—ereeeee rr

Fies (-7. Macropestrongylas Macreposirangyles. 1, median view of head; 2. lateral view of head, with

mouth witely opened; 3. en face yiew of head: 4. oesophageal regions 5. bursa; 6 dorsa) rag;

7, posterior end of female; Figs 1-3 10 same scale. Pigs 4. 5, and

to Sate scale. Figs 4, 7, 6

to same scale. Figs. 8-12. Mucropostrongy lis winerestome, B. anterior end; 9, anterior end. on

face; 1M, oesophageal region; 11- bursa} 12, posterior end of female, Pigs ¥ and 11 to «ime

scale.

REVISION OF MACROPOSTRONGYLUS (NEMATODA: TRICHONEMATIDAE) 53

FIGS 1-12

54 PATRICIA M. MAWSON

Host and locality; M, ayilis, from Weam,

Papua New Guinea (BBM-NG.S0820,

BBM-NG-S0798}; Vhylogale branti, from

Weam, Papua New Guinea (BBM-NG-

50850),

The specimens from Papua New Guinea are

shorter than those originally described, but they

agree generally with them, and with those iden-

tified by Baylis (Yeerongpilly N5.28.1.2) and

others recorded by Johnston & Mawson, Some

redescription of the anterior end can now be

made. (Figures were drawn from Papua New

Guinea material from M, agilis,)

Amphids le on apices of two prominent

lateral clevations, Buccal capsule, somewhat

laterally compressed, is not strongly chitinised;

the two largest of the longitudinal ridgrs are

lateral, and the two smallest dorsal and vestral.

Ratio length: spicule length 3.0-3.6, and of

length: oesophagus 3,3-4,2,

Cervical papillae thread-like, about twice the

length of buccal capsule from anterior end,

Egg aa. 80 x 40 um,

Macropostrongylus macrostoma Davey & Wood

FIGS. 8-12, 47

Davey & Wood, 1938 p. 260, from Maerapues

robustus, Queensland,

Macropostrongylus yorker (ted Baylis): Johnsian

& Mawson, 1939 p, 143, p.p., from M, parey!

Gelanestrongylus macrestoma: Popova, L952 p,

176,

The paratype material of this species has

been examined and figured,

The anierior end is similar to that of Av.

macropostroneylus, The main — differences

between these species are

1, Buceal capsule longer and more strongly

chitinised in M. pnaerestoma, ond is anterior

projections more strongly developed ahd

reinforced hy extra sclerotisation in the form

of an encircling bell at ahout ils mislength,

and hy a thickening arownd base, greater

dorsally.

.Oesephuyus swollen in middle third of its

leneth i Af, macrastorna not swollen if

M. snacropostronyy lis,

. Pormof the dorsal ray differs.

Tu Ad. macrapostronevius distance between

Vulva and anus is less Than tail length: in

M. macrostomea. it ig distinctly greater.

Macropostrongylus yorkel Baylis

FIGS 13-19, 51

Baylis, 1927 p, 215, fram Macropus sp, Toews:

ville, Old; 1934 p. 124%, from MW. evilis, Barke-

town, (ld,

Jobnston & Mawson, 193% p. b43; 1979 p. 209,

from MM. agilis, Old.

= ia

Host and locality; Marroprs azilis (stomach),

Tipperary Sun, N.T-

The material identified by Johnston & Maw-

sen is scanty and in poor condition. That

reported from M. parryi by Johnston & Maw-

son (1939) is now referred to M. macrostoma

(q.v.), and the single female worm from Aq.

wellshyi (now Wallabia hicolor wellsbyi), pro-

bably belongs to an as yet undeseribed genus.

The type and paratypes have not been seen,

Nhe following partial redescription is based on

some recently collected specimens of M. agiliy.

Length of male, 6.5-8.6 mm, of female

14.2-20.5. Anterior end outlined by a low

ridge, oval in en face view, with the long axis

dorsoventral, Within this, submedian cephalic

papillae and aniphids. are on slightly raised

cuticular swellings, Bueeal capsule ore

rounded-triangular than oval al its base, the

longitudinal ridges developing in its anterior

half and surrounded near base by a sclerotised

nng, The whole area inside the anterior ridge

probably eversible. Eversion is ussociated with

ain upthrust of the anterior end of the ocso-

phagus, While the buccal capsule appears to

widen, so becoming a longer oval in Lransverse

section (Pig, 14),

Oesophagus long (body length: oesophagus

3.6-4.3 in male, 5.8-6.2 in female), more or

less cylindrical anterior to spindle-shaped ter-

minal bulb, Nerve ring surrounds oesophagus

at about a third to a quarter of its Jength from

head in male, less in female; thread-like cervi-

cal papillae lic about half way between anterior

end and nerve ring, and excretory pore close to

postenior end of oesophagus. Tips of spicules

enlarged and alate. Ratio length; spicule 10.9—

14.0. Eggs measure 98-110 x 53-55 am,

The species is most like M. macrastama, dif-

fering chiefly in the size and form of the buccal

capsule.

Macropostrangylus Jesouefi Johnston &

Mawson

FIGS 20-24

Macropostronovias, Jesoucfi Johnston & Mawson.

1939 p, $25, from Macropus rrfovrisea, Sydney

Zoological Gardens.

Gelanestrangy lias lesouefi: Popova, 1952, p. 769.

No fresh material of this species is available,

The type and paratype material have been

examined, and some redeseription is possible.

The species is distinguished by the very pro:

minent Wweral cuticular clevations, bearing at

(heir apices the Openings of the amphids. Hue

cul capsule lalerally compressed only in some

REVISION OF MACROPOSTRONGYLUS (NEMATODA: TRICHONEMATIDAE) 55

Figs 13-19.

50 pm

FIGS 13-24

Macropostrongylus yorkei. 13. head of male; 14. head of female, in mouth wide open posi-

tion; 15. head, en face; 16. oesophageal region; 17. lateral view of bursa; 18, dorsal ray; 19.

posterior end of female. Figs 20-24. Macropostrongylus lesouefi, 20, anterior end, lateral

view; 21 and 22. anterior end in dorsal view, with mouth in closed and open positions, res-

pectively; 23, oesophageal region; 24. tail of female. Figs 13-15 to same scale. Figs 17-18 to

same scale. Figs 19 and 23 to same scale. Figs 20-22 to same scale.

56 PATRICIA M, MAWSON

specimens, suggesting that this is a movement

connected with feeding. Anterior projections of

the capsule strongly developed, those in dor-

sal and ventral positions directed outwards,

Oesophagus long and more or less cylindn-

eal, with elongate terminal bulb, Nerve ring

further back than in other species, almost at

the end of the anterior half of the oesophagus.

Excretory pore just behind nerve ring; cervical

papillae half way between nerve ring and head.

In the only specimen in which the spicules are

intact, they measure 480 4m,

Aus of female closer to vulva than to pos-

terior end of body; vagina short, Eggs 145-155

x 70-75 #m,

Macropostrongylus irma Johnston & Mawsoau

Johnston & Mawson, 1940 p. 363, from Macropus

irmna, Wi A.

Gelanastronevlus fenia: Popova, 1952 p, 769

These specimens ate immature, probably

fourth stage larvae, as the vulva is not patent,

‘Two referred to as “females differing some-

what" are in fact fourth stage larval males

This species should perhaps be declared a

nomen nudum, but it may be possible to recog-

nise jt should fresh material become available

from the same host species. For the tlme being

it is retained.

Key lo species of Macropostrongylus (ex-

cluding MM. trea),

1. Amphids 6n Very prominent cuticular elevations

BPlevations bearing amphids not higher than sub-

median papillae 3

2. Nerve ring about | length of oesophagus from

head; [nteral branches leave dorsal ray

immediately after its bifurcation .,, ,,,

M, macropostrongy lus

Nerve ring at nearly 4 Jength of oesophagus

from head; lateral branches leave dorsal ray

neur edge of bursa : M, lesorefi

3, Buccal capsule longer than its width

M,. macrestoma

Buccal capsule not longer than its width , ;

M. yorkei

Popovastrougylus 1y.gen.

Syn. Mucropostrongylus Yorke & Maplestone

Pp.

Trichonematidae: Antenor end with outicu-

Jar collar bearing four setigerous submedian

papillae and two amphids, buccal capsule and

mouth opening circular to oval; extension of

perioral cuticnle lines buccal cavity and may

project as shelf inside mm: buccal capsule

thickest in its midleneth, anterior border with-

out projections. ocsophagus cylindrical

anteriorly, usually narrowing abruptly in

second half, ending in bulb, Male: spicules

alate, gubernaculum present; bursal lobes dis-

(tinct, ventrals not joined, ventral rays separate

from laterals, cxterno-dorsals arise with

laterals, dorsal ray bifurcate, each branch with

shorter lateral off-shoot. Female; tail long,

vulva near anus. Parasites of the stomach of

macropod marsupials.

Type species: P. wallabiae, syn. Macropo-

strongylus wallabiae Johnston & Mawson,

1939,

Other species: P. pearseni, syn Macropo-

sirongylus pearsoni Johnston & Mawson,

1940, P. irmea, o.sp.

Popovastrongylus wallabiae (Johnston &

Mawson)

FIGS 25-30, 44

Macropostrongylus wallablae Johnston & Mawson,

1939 p. 526, from Wallabie bieolor (M_ walla-

bars) from NS.W.

Gelanastrongylus wallabiae: Popaya, 1952 p. 785.

Host and locality: Macropus rufogriseus,

Logan Village, Qld) Launceston, Tas.

Collections of this species from three hosts

in the same area in Queensland and in one

from Tasmania permit an elaboration of the

original description, in regard to head structure

and shape of the dorsal ray.

The small anterior collar is less obvious in

some specimens than in others, as it appears

partly retractable. In the type specimens 4

narrow shelf is present towards the anterior

end of the boccal cavity, but this is not clear

in all specimens. Figs 25-30 were drawn from

the type specimens. In the new material from

M, rujogrivea the eggs measure 105 x 50 ym,

The length:spicule ratio is 9.0.

Popovattrongylus pearsoni (Johnston &

Mawson)

FIGS 31-33, 48

Macropostroneylus pearson’ Johnston & Mawson,

194¢) p. 95, Mawsan, 1971, 171; fram Petrogale

penteillnta pearsont from Pearson 1,, 8. Aust,

Host and jocality: Macrepys eugenii, Kan-

garoo L, S. Aust.; Mecropus rufogriseus

from Launceston, Tasmania.

Pepevasrrongylus pearseni was redescribed

by Mawson (1971). It is similar in many

features to P. wallabiae, particularly in’ the

structure of the head. As both species have now

been identified from the same host species in

Tasmania (though not as yet from the same

host specimen) the main features distinguish-

ug them are given:

Figs 25-30, Popovastrongylus w 2

ventral views: 29. genital cone, ventral view;

-In P.

REVISION OF MACROPOSTRONGYLUS (NEMATODA: TRICHONEMATIDAE) 57

300 pm

FIGS 25-33

allabiae. 25. Head; 26. oesophageal region; 27 and 28. bursa, lateral and

30, posterior end of female. Figs 31-33. Pepo-

vastrongylus pearsoni. 31, oesophageal region; 32, bursa; 33, genital cone, dorsal view. Figs

26, 30 and 31 to same scale. Figs 28 and 32 to same scale. Figs 29 and 33 to same scale.

_In P. wallabiae terminal bulb of oesophagus

is spherical; in P. pearsoni it 1s more oval.

wallabiae nerve ring surrounds

oesophagus well in front of the point where

it narrows, in P. pearseni it lies at this point.

. Dorsal lobe of bursa is much longer than

lateral lobes in P. wallahiae, but about the

same length in P. pearsoni.

. Shape of the dorsal ray differs (Figs 28, 32).

. Appendages of the dorsal lip of the cloaca,

on the genital cone, differ (Figs 29, 33).

.Spicules rather shorter in relation to body

length in P. wallabiae.

Popovastrongylus irma n.sp.

FIGS 3440, 50

Host and locality: Macropus irma (stomach),

from Perth, W.A.

Males 8.7—-10.1 mm long, females 11.1—13.0

mm. The cephalic papillae, borne on a well

developed cuticular collar, are not prominent.

The buccal capsule, its base thickened by an

outer sclerotised ring, is a little more oval than

circular in transverse section, with the long

axis not exactly dorso-ventral; it lacks an in-

ternal shelf.

3k PATRICIA M, MAWSON

The oesophageal balh ts slightly elongate.

Ratio length: oesophigus ts 7,2-8,4 in male,

78-91 in female. Nerve ring surreands

ocsophagus at the point of narrowing, and

excretory pore is behind this; the thread-like

cervical papillae lie shortly behind anterior end,

Spicules alate and end in u rounded Up,

without enlargement; ratio length: spicule is

6.7-8.6. Bursa voluminous, all lobes of more

or less even length; ventral lobes joined.

Genityl cone bears two small bilobed processes

of dorsal hp of clowea. Bursal rays are shown

in Figs 37, 38.

The female has an unusual constriction

between the yulya and the anus; in older

females the body is markedly swollen in the

region of the vigina, as far back as this con-

striction, Tail conical, ending in a point, Ezes

absent in all specimens.

This species is distinguished from P.

wallabiae and Po pearseni chielly by the

absence of a “shelf in the buccal cavity and

by the shape of the dorsal tay.

Key to species of Popovastrongyl is:

t, Nerve ring surrounds oesophagus alietnetly ant

terior to ils narrowing P, wallablae

Nerve ring surrounds oesophagus al point o

narrowing 2

2. Lining uf buceal capsule forms distinct “shelf

: P_ pearsoni

Lictng of baccal capsule withour “shelf”

Py irvine

Arundefia o.gen.

Trichonematidye: Cloacininae: Small worns

with heavily fringed cuticle; anterior end with

small cuticular collar, four small, bipartite, sub-

median, cephalic papillae; external leaf crown

of 6 elements; lips absent; bu¢cal capsule short,

stoutly built, circular in transverse section; base

of buccal cavity with large hollow oesophageal

projechion, associated with dorsal duct in

oesophagus: oesophagus widening posteriorly

but without bulb, Male: bursa short, wide; ven-

tral rays arising together, ventro- afd miedio-

laterals arise together, postero-laterzl and

externo-dorsals arise separately: dorsal ray

shown itt the 6.8, Micrographs (Fig. 52), They

bifurcates twice, spicule stoutly built, alate:

subernaculum and telamon present. Female:

vulva close to anus, Parasites of the stomach

of macropod marsuprals.

Type species: A_ dissimilis, syn. Macropo-

strongyluy dissimilis Johnston & Mawson,

1939.

A small dorsal tooth associated with an

oesophageal duct has been described in the

buccal capsule in Cloacina dahli \instow,

1898 and in C. murday! Mawson, 1972, but

int Cloaeina an internal leaf crown, arising from

the baccul capsule, is present.

In Popevastronpylus species the buccal cap-

sule is oval to circular in section and a leat

crown is absent, but there is a cuticular lining

inside the buccal capsule, no tooth in the buccal

cavity, and the oesophagus is quite a different

shape. In Mucropostronyylus the shape of the

buceal capsule is quite different.

Arundelia dissimilis (Johnston & Mawson)

o.conib.

FIGS 41-44, 52

Mucrepostrongyluy dissimilis Johnstan & Mawson,

1939 p. 526, from Wallabia bicalor (M_ walla.

batus), NSW,

Hust and localities: Wallabia bicolor, trom

Keyneton, Bemm River, Yarra Valley, and

Dartmouth, Victoria.

This species is apparently a relatively com-

mon though not numerous parasite of Wallabia

bicolor, and has not been found in any

other macroped, Details of the buccal capsule

are Visible in the fresh material so the descrip-

jion can now be amended. Measurements of

the specimens are similar to those of the

original description,

Cuticle strongly ringed; body widest in pos-

terior half, tapering slowly to head, very

rapidly behind vulva, and very little to bursa.

A low, thick cuticular collar anteriorly bears

submedian papillae and unobtrusive, slittike,

amphid openings. Mouth cireular without tips,

Six short, triangular, cuticular projections

around mouth not easily seen in side view but

Figs 34-40, Popovastronyylas irina. 34, bead; 38, bead with buccal capsule dorsoventrally compressed;

36, oesophageal region: 37-38, bursa jn lateral and dorsal views} 39 and 40, Posterior ends

of younger and older females, respectively, Figs 34 and 35 to same scule. Figs 37 und 38 to

same scale. Figs 39 and 40 to sume scale. Figs 41-44. Arundelly dissimilis, 41-43. Head, in

lateral, dorsal and en face views respectively; 44, oesophageal region, Bigs 42-43 to same

scale, Pigs 45-46. Dorcopsinemu dercopsis. 45. Head of male, antero-lateral view; 46, head

of female, sub-lateral view.

REVISION OF MACROPOSTRONGYLUS (NEMATODA: TRICHONEMATIDAE) 59

FIGS 34-46

MAWSON

“FIGS 47-53

REVISION OF MACROPOSTRONGYLUS (NEMATODA: TRICHONEMATIDAE) fl

form) a sort of leaf crown hut do not appear

to arise from the buccal capsule as do the

elements of the leaf crown in Cloacinu species.

Buccal capsule, circular in transverse sections,

shallow, but with thick walls, Large, conpeal,

chitinised structure rises dorsally in buccal

cavity from anterior end of oesophagus; this

is hollow, open at apex, and connected al its

base with u duct in dorsal wall of ocsophagus.

In en face view, a ventral thickening snd

groove in the capsule wall is associated with a

similar but smaller duct from oesophagus,

which has not been seen in any side view of the

anterior end,

Ocsophagus cylindrical im its anterior third.

at end of which lies the nerve ring, and then

widens gradually to its posterior end, No ter-

minal hulb. Thread-like cervical papillae lie

anterior to nerve ring; excretory pore at about

three-quarters length of oesophagus [rom

anterior end.

Size of eggs in the vagina, and newly lanl

in the Vaginal extrusion, is 130-132 x 65-70

pm, much greater than in original material

(possibly measured in the uterus),

Dorcopsinema n,gen,

Trichonemiatidae: Zontalamminae: Large

worms; anterior end with wide collar bearing

cephalic papillae and amphids, perioral cuticle

forming eight lip-like processes; buccal cap-

sule lightly chitinised, more or less cylindrical;

oesophagus long, cylindrical Male: spicules

alate, long: bursa entire, dorsal Jobe long, ven-

tral ritys arising logether, externo-dorsal aris-

ing with laterals, dorsal ray bifurcating and

with two lateral branches from point of bifur-

cation. Female: tail conical, vulva shortly in

front of anus. Parasites of macropod mar-

supials.

Type species: D. doreapsis (Baylis).

Macropostronyylus dorcopsis Baylis, 1940.

The structure of the head does not closely

resemble that of any other species. The lip-like

processes around the mouth are very like those

of Luhiostrongylus and Zoniolaimus but the

cephalic papillae are borne on the collar. In

2. lahiarus Johnston & Mawson (1939) there

is a collar around the anterior end bearing the

cephalic papillae, and surrounding the “lips”,

but the ocsophagus ends ina bulb.

svn,

Dorcopsinema dorcopsis (Baylis) n.comb.

ViGs 45-46, 53

Mocrapastroiaylus dorcopas Baylis, 1940, p. 313,

Som Dorcepsis mullers CD. veterum) trem

Papua New Guinea.

A Male atid a ferale paratype have been

examined, The presence of a very thick collar,

the structure of the buccal capsule (no tongi-

tudinal rides, no anterior projections) and the

shape of the oesophagus differentiate the

species from those of Maecropostrongylus.

The “tosth-like processes" around the

mouth “like a leaf crown” described by Baylis

are tm fact not thi and chitinised like teeth (or

uw leaf crown) hut are more like fleshy lobes,

with broad bases, mucronate at the free ends

and grooved on their outer surfaces, In the

male these processes are almost closed over the

mouth and in the female are drawn back in a

“mouth open” position (Figs 45, 46), As

described by Baylis, the anterior end is sur-

rounded by a wide collar on whieh ate the

small pointed submedian papillae and the

amphids,

Acknowledgments

The greater part of the new material

examined in this work was provided by Profes-

sor Arundel and Dr Beveridge of the Mel-

bourne University School of Veterinary

Science. That from Macropus irma was

obtained through the kindness of Dr de

Chancet of the Animal Health Laboratory,

Perth. Paratype material was lent by the

Sehool of Public Health and Tropical Medicine

in Sydney (Mavrapostrongylis macrostoma,

Mf. cormutus, Mf. miner and M. lebiatus), and

by Mr 8. Prudhoe of the British Museum (Nat.

Hist.) (At. dorcopsiy and M. baylivi). Speci-

mens identified by Baylis as M, macropo-

veongylus and M. australir were lent by Dr

Green of the Animal Health Luboratory at

Yeerongpilly, T am very grateful for all this

help.

The micrographs (Figs 47-53) were taken

by E.T.E.C, Autosean jn the Central Electron

Optical Laboratory of the University of

Adelaide. | am indebted to Dr Karl Bartusek

of this Laboratory for help in taking the micro-

graphs and to P. G. Kempster for developing

and printing them,

Fig. 47. Muatropostroneylas macrastena (<404): Fie 48. Popavavronevine wintlahlae CxA0); Pig. 49. Pe

péarsont (x600); Fig. 50. P. fema (x600); Pu St AL. vorker (2404) Fie. 52. drandelia aliy-

sillis (x1,500)5 Fig, $3, Dareapsaenid darcapsds (x240\

62 PATRICIA M. MAWSON

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todes from Australia. Ann, Mag. nat. Hist.

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Baytis, H, A. (1934) Some parasitic worms from

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Bay is, H. A. (1940) A new species of the genus

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Coss, N. A. (1898) Extract from M.S. Report

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Jounston, T. H. & Mawson, P. M. (1938)

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Jounston, T. H. & Mawson, P. M. (1939a)

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Jounston, T. H. & Mawson, P. M. (1939b)

Strongylate nematodes from Queensland mar-

supials. Ibid. 63, 121-149.

JoHNsTON, T. H. & Mawson, P. M. (1939c)

Strongylate nematodes from marsupials in

New South Wales. Proc. Linn. Soc. N.S.W.

64, 513-536.

’

JOHNSTON, T. H. & Mawson, P. M. (1940) On

a collection of nematodes from Australian

marsupials. Rec. Aust. Mus. 20, 360-366.

Linstow, O. von (1898) Nemathelminthen gesam-

melt von Herrn Prof. Dr. Dahl im Bismarck-

Archipel. Arch. f. Naturg. 1, 281.

Popova, T. I. (1952) Jn K. I. Skrjabin (Ed.), Key

to parasitic nematodes. 3. Strongylata. (In

Russian) Acad. Sci. U.S.S.R.

Mawson, P. M. (1971) Pearson Island Expedi-

tion. 8. Helminths. Trans, R. Soc. §, Aust. 95,

61-64

Mawson, P. M. (1972) Three new species of the

genus Cloacina Linstow (Nematoda: Strongy-

lata) from macropod marsupials. Jbid. 96,

109-113,

Mawson, P. M. (in press) Excerta Parasitologica

en memoria del Doctor Eduardo Caballero y

Caballero.

Woop, W. A. (1930) Some new parasitic nema-

todes from Western Australia. Rep. Director

Inst. Animal Path. Univ. Cambridge (1929-

1930), 209-219.

YAMAGUTI, S. (1961) Systema Helminthum, Vol.

Ill. The nematodes of vertebrates. (Inter-

science: New York.)

York, W. & MApLestone, P. A. (1926) “The

nematode parasites of vertebrates.”

(Churchill: London, )

SHORE PLATFORMS AND SEALEVEL CHANGES IN THE GULF

REGIONS OF SOUTH AUSTRALIA

BY C. R. TWIDALE, JENNIFER A. BOURNE AND NICHOLAS TWIDALE

Summary

The platforms that occur along the shores of Kangaroo Island and the Gulfs region of South

Australia are developed in Precambrian crystalline rocks, Palaecozic sedimentary strata and

Pleistocene dune calcarenite. Evidence from these areas suggests that many platforms cut in granite

and gneiss are etch surfaces, or weathering fronts developed in late Pliocene and early Pleistocene

times, exposed by erosion in Recent times, and only fortuitously situated within the present tidal or

spray zone. Elsewhere the unconformity between crystalline rock and aeolinate has been revealed

by marine processes. Contemporary platforms cut across the structure of contorted sediments and of

the aeolinate are widespread and occur as much as 8 m above present sealevel. Since they must

postdate the aeolinate which is regarded as of last-glacial age, they attest to considerable erosion in

a short time. Only on the coast of Kangaroo Island is there unequivocal evidence of an earlier and

higher stand of the sea 5-6 m above present sealevel.

SHORE PLATFORMS AND SEALEVEL CHANGES IN THE GULFS REGION OF

SOUTH AUSTRALIA

by C. R. Twibace,* JENNIFER A. BouRNE* and NicnHoLas TWIDALeEy

Summary

Twibacr, C. R., Boukne, J. A, & TwibaLe, N. (1977) Shore platform und seulevel changes

in the Gulfs region of South Australia. Trans, R. Soe. §, Aust. 10102), 63-74, 31 May,

1977

The platforms that occur along the shores of Kangaroo Island and the Gulfs region of

South Australia are developed in Preeambrian crystalline rocks, Palacozoic sedimentary strata

and Pleistocene dune calcarenite, Evidence from these areas suggests that many platforms cul

in granite and gneiss are etch surfaces, or Weathering fronts developed jn late Pliocene and

early Pleistocene times, exposed by erosion in Recent times, und only fortuitously situated

within the present tidal or spray zone. Elsewhere the unconformity between crystalline rock

and aeolianite has been revealed by marine processes. Contemporary platforms cut across the

structure of contorted sediments und of the aeolianite are widespread and occur as much as

8 m above present sealevel. Since they must postdate the aeolianite which is regarded as of

last-glacial age, they attest to considerable erosion in u short time. Only on the coast of

Kangaroo Island is there unequivocal evidence of an earlier and higher stand of the sea 5-6 m

above present sealevel,

Introduction

Several of the shore platforms that occur on

the coasts of western and southern Eyre Penin-

sula, and Kangaroo Island (Fig. 1) are

anomalous, albeit in different ways,

In cach of these areas Precambrian ecrystal-

line rocks (granite and gneiss) or Cambrian

sediments, metasediments (mainly schists),

and granites are exposed from beneath the

widespread Pleistocene dune calcarenite. The

latter is commonly referred to as acolianite

(Crocker 1946a) for it was deposited in coastal

dunes and subsequently lithified through

secondary calcification. The old dunes are

extensively preserved in coastal areas of

Western Australia, South Australia and Vic-

toria, They stand up to 50 m above sealevel

and as has long been appreciated the rock

extends well below present sealevel ut many

places (Tate 1879; Sprigg 1961; Cooney 1965);

for example, it is recorded that aeolianite rests

on bedrock of probable Precambrian age at a

depth of some 61 m just east of Elliston.’

There is thus no doubt that the acolianite is

related to a glacial phase or phases of the

Pleistocene but there is as yet no more precise

age determination, Many relic soil profiles are

revealed within the aeolianite sequences in cliff

sections but the permeability of the old dune

rock is such that water readily infiltrates into

the mass, and weathering and soil formation

are rapid, Despite appearances to the contrary

the building of the dunes and the deposition of

the acolianite probably did not occupy a long

period of time. As Fairbridge & Teichert

(1952) state of the Western Australian

aeolianite dunes “the periods of soil formation

were not of long duration and, . , the dune

developments were in rapid, sequence”. The

dune calcarenite exposed in the areas under

investigation was probably all deposited in a

single glacial phase. Certainly there is no

evidence of cut and fill, such as might be

expected had the dune rock presently exposed

been built up in the course of several glacial

periods and subjected to dissection during sub-

* Department of Geography, University of Adelaide, North Tce, Adelaide, 5S, Aust. 5000.

{ Medical School, Flinders University of South Australia,

1. G, Shepherd (1962) Report on groundwater prospects, Hundred of Ward, Elliston Police Station.

Geal. Sury. 8S. Aust, Rept, Bk, $8/32.

64 Cc. R. TWIDALE, JENNIFER A. BOURNE & NICHOLAS TWIDALE

Pt Brown

High Cliff, 7 STREAKY BAY

Smooth Poo!|—€-——Westall Peninsula

Speed Pt

Cape Labatt

EYRE

PENINSULA

-Talia Caves

Wellesiey Pr7 XE ttISTON

Drummond Pt.

Redbanks

Cape Carnot

Cape Wiles

Stenhouse Bay

Cape du Couedic

G-Northumberland (J

Daly Head —

Gym Beach

PORT RICKABY

Brown Pt.

Pl. Turto:

ADELAIDE

Hallett Cove

Wentworth Pt,

Cape Cassini.

Windmill Bay

Fig. 1. Location map,

sequent periods of glacial low sealevel, and to

planation during the interglacial high stands

of the sea.

But which glacial phase: the last, the first,

or some intermediate phase? There is no direct

evidence on this point. On the other hand there

is nothing to show that the acolianite so widely

exposed on the South Australian coast is not

all related to the last glacial (Wisconsin)

period of low sealevel. And there are several

inferences to suggest that they are. For

instance, if the dunes predate the Wisconsin

they ought, judging from the rate of postglacial

erosion, to be more extensively baselevelled

than they now are. There ought to he signs

of interglacial high stands of the sea imprinted

on the dunes, and of these there is no indica-

tion, If it is argued that the evidence has been

destroyed by erosion, again, why have the

dunes survived at all? And if the dunes are of

great antiquity why were they not deeply

eroded during the Pleistocene glacial low sea-

levels? Further reference is made to some of

these arguments below, but the aeolianite is

taken to be of late Wisconsin age in the areas

under discussion.

The shore platforms of the coastal sectors

under consideration are developed in both the

older crystalline and sedimentary rocks and the

younger aeolianite. They pose _ several

problems,

Platforms deyeloped in fresh granitic bedrock

According to Jutson (1940) and Hills

(1949; 1971), shore platforms are poorly

developed in fresh granitic bedrock, yet along

considerable sectors of the west coast of Eyre

Peninsula which is exposed to westerlies sweep-

ing in off the Bight, shore platforms are com-

monly developed in granite and in places

extend some 200 m from the base of the cliffs.

How have they formed and in what way, if at

all, are they related to the contemporary sea-

level?

SHORE PLATFORMS AND SEALEVEL, CHANGES, GULFS REGION

calcrete

DRUMMOND. POINT

etch surlace

high tide platform

intertidal platform

aealianite

<< UnGontarmity

(lerruginised)

~) Weathered Pa

gneiss

ey we: aay ie

Pfft ff fff fresh P€ gneiss

fi / if / y

/ }

/ 1) //, 4)

WELLESLEY POINT

aeolianite

calcrete

Fig, 2a, Diagrammatic com

high tide eM

_-~+~

intermediate ratrorm

25m

mean sea level

site section at Drummond Point, The tiny remnant of calerete on the

} isolated hill (left) is developed on acolianite,

Fig, 2b, Section of the acolianite cliffs at Wellesley Point (see also Fig. 7b),

1. Point Brown, northwestern Eyre Peninsula

Point Brown (Fig. 1) is typical of several

sites on the west coast of Eyre Peninsula in

that Pleistocene acolianite clearly overlies

granite; but this locality, and that at Drum-

mond Point (see below), together provide the

clearest evidence concerning the probable

origin of the granitic shore platforms.

The granite is even grained but coarsely

granular. Tt is gneissic in places and there ure

veins of aplite and quartz, but most of the

exposure is of the coarse-grained “sugary”

granite. This is not fresh for it is iron-stained

and disintegrates. when hit with a heavy harm

mer; but it is cohesive and by no means friable.

Some of the eranite exposures are bouldery,

and concentric zones of flaking are preserved

around some of the boulders, suggesting that

they are corestones or kernels (see Senvenor

1931) Larsen 1948; Linton 1955; Twidale

1971), But elsewhere the surface of the granite,

though criss-crossed by joint crevices and

clefts, is essentially even, and though in detail

composed of many individual joint blocks,

forms a platform that in places extends as

much us 30m from the base of the cliffs,

However, both on Point Brown itself and on

the next headland to the south a _ rego-

lith of weathered granite separates the lresh

granite from the aeolian calcarenite, This

wWeuthering profile must have developed

beneath a near-stable surface of low selief.

Regional considerations suggest that it was the

late Pliocene-Pleistocene Koongawa Surface

(Twidale, Bourne & Smith 1976). The weather-

ing profile, 2-4 m thick, is mottled red, yellow,

white and grey, but the granulay texture is

retained and the joints remain clearly dis-

cermible. The regolith is obviously belng eroded

hy wave action and by pool weathering: there

ate undercut visors on the seaward-facing

bluffs, and alveolar forms and weather pits are

commonplace, The strippmg of this regolith

has exposed the former limit of weathering—

the weathering front of Mabbutl (1961a), The

weathering front is, as might be anticipated,

irregular because of the deeper penetration of

water and alteration along afd near joint

planes. Hence, in some places, corestones are

exposed but where the front is even platforms

are revealed. In other words the platforms are

etch surfaces (see Wayland 1934; Mahbutt

1961b; Twidale 1976),

2. Drummond Point, southwestern Eyre

Peninsula

At Drummond Point (Fig. 1), Precambrian

gneisses with well developed steeply dipping

lineation are exposed beneath the dune cal-

66 C. R. TWIDALE, JENNIFER A. BOURNE & NICHOLAS TWIDALE

. Part of Drummond Point, showing cal-

carenite overlying gneiss. A—calcarenite;

B—gneiss exposed in platforms.

. Smooth Pool showing the calcarenite

cliffs, the massive joint blocks exposed in

the granite platform, and the many

granite boulders and cobbles, which were

originally corestones.

Fig. 3c. Granite platform at Smooth Pool show-

ing large joint blocks, the one in the fore-

ground displaying raised rim and saucer-

shaped depression.

carenite that forms the upper half or three-

quarters of the high cliffs (Figs 2a and 3a).

The unconformity between the Precambrian

and Pleistocene rocks is irregular, but, though

eroded, is nevertheless expressed in many

places as a gently sloping bench standing well

above high tide level? but within the spray zone

(Fig. 2a). A narrow platform serrated in pro-

file has developed at about mid-tide level but

there is another prominent flat located at, or

more commonly just above, high tide level. It

is everywhere coincident in elevation with the

weathering front, the junction between the in-

trinsically fresh gneisses below and_ the

weathered rock above.

The coastal exposures at Drummond Point

suggest very strongly that the high level plat-

form, standing 6 m above the intertidal plat-

form which is located at about mid-tide level,

is of etch character. Here perhaps even more

clearly than at Point Brown is the nature of

some of the platforms revealed: the compara-

tively high platform at Drummond Point (and

the mid- to high-tide platforms at Smooth Pool,

Cape Labatt and Talia Caves—see below) are

etch surfaces and they have no significance so

far as former stands of the sea are concerned.

They have developed only because the weather-

ing front happens to be located within the tidal

and spray zone of contemporary sealevel.

The weathered gneiss consists of quartz

grains and fragments set in a matrix of

kaolinised material, and with abundant iron

oxides. There is, however, no sign of a

ferruginous capping or horizon within the

gneiss; rather are there patches of iron dis-

colouration following lineation within the rego-

lith. Whether there was originally a’ferruginous

carapace that has been destroyed by chemical

attack following burial by the aeolianite is not

known (cf. Twidale & Bourne 1975).

However, though particularly well displayed

there, Drummond Point is not the only site

where there is evidence of the etch character

of some of the platforms cut in pre-aeolianite

crystalline and, in one instance, sedimentary

rocks,

3. Westall Peninsula, northwestern Eyre

Peninsula

The term Westall Peninsula is used of the

complex promontory that lies to the west and

southwest of Streaky Bay (formerly Flinders)

? The evidence of jetsam on beaches, distribution of lichens, etc. at particular sites has been taken in

preference to theoretically derived tidal ranges. Thus “high tide level” implies “near the top of the

evidenced tidal range”.

SHORE PLATFORMS AND SEALEVEL CHANGES, GULFS REGION 67

High Cliff

( WESTALL PENINSULA

V ~

Smooth Pool \=>

Fig. 4. Westall Peninsula showing the extent of

the shore platforms cut in granite and

gneiss. Major joint trends are shown.

township. Most of the exposures, notably in the

60-70 m high cliffs, are of aeolianite but over

long sectors granite and granite gneiss are

exposed at and near present sealevel. At High

Cliff, located 14 km southwest of Streaky Bay,

strong structural control is evident in the plat-

form. The bedrock is subdivided into massive

slabs by prominent joints which dip 2° sea-

wards. However, despite the many minor joint-

block “cuestas” developed in the massive

granite slabs, the outcrop has been eroded and

overall the granite forms a platform extending

as much as 80 m beyond the base of the

aeolianite cliffs, though with clefts well

developed along joints.

The granite outcrops at Smooth Pool have

also been eroded to form a platform which

slopes gently (4°) westwards from the base of

the aeolianite cliffs (Figs 3b and 4). At low

tide the platform extends about 200 m from the

base of the cliffs, but at high tide is covered by

the sea. Over wide areas it is coincident with

horizontal or sub-horizontal joints but in many

places the surface transects the joint blocks, so

that there are innumerable minor platforms

within the larger feature.

In detail the blocks that comprise the plat-

form display a varied morphology. Some have

slightly raised rims, and are in consequence dis-

tinctly saucer-shaped (Fig. 3c). Others are

—-

_—.

— -

a eS

tie ee

Fig. Sa. Flared slope of granite boulder at

Smooth Pool.

Fig. 5b. Granite platform cut in granite at Went-

worth Point, Yorke Peninsula. It stands

below the unconformity between the cal-

carenite above and the granite below but

the platform seen in the middle distance

is essentially coincident with the uncon-

formity.

Fig. 5c. Calcified grus with granite boulders set

in it at Wentworth Point.

slightly humped with broad rises a few centi-

metres amplitude standing above the general

level of the edges of the platforms. Some dis-

play both saucers and humps. The joint-con-

trolled margins of the blocks are uniformly

steep and some are slightly concave. There are

68 R. TWIDALE, JENNIFER A. BOURNE & NICHOLAS TWIDALE

inany Jonse rounded boulders (diameter $0 om

and fess) strewn over the platform und 4lso,

particularly in the region close to cliff-feat,

many larger joint blocks are rooted in the plat-

forms, Some of the latter are distinctly Mared

(Fig. 5a). One block has been ereied to form

a platform on the landward side and the hump

standing above il is slightly fared so that the

total effect is that of an elephant rock (Twidale

& Bourne 1976),

The platform can be traced continuously

round the Peninsula at approximately the same

elevation for several kilometres. The tncon-

formity between the crystalline rocks and the

overlying limestone is obscured by the coarse

blocky rubble that has fallen and accumulated

at the base of the cliffs, but to jude from the

Upper limits of fixed gneiss and pranire boulders

in the oliff-foot regien [t occurs |-2 m above

the present high tide limit,

On Westall Pemnsula, evidence of the etch

ongin of the wide platform is fragmentary and

diffivult to Rnd However, at one site at Smooth

Pool there is a patch of Weathered granite,

motel in red and yellow preserved beneath

the bouldery grarite sind blocky aeolianite

rubble, This and the many loose houlders suz-

gest that thers was wl onc time a regolith cam-

prising corestones set in a matrix of ers over

the fresh rock, Wave action must have stnpped

all but one small remnant of the gris away

though many of the <orestones remain as

boulders. The lNared marwins of fixed boulders

and of joint blocks ave in these terms due to

subsurface weathering (Twidale 1942),

This interpretation is corroborated by the

character of the unconformity between eranite

and aeoljanite exposed on a minor headland

abour one kilometre south of Smooth Pool.

Here there is nhiindant calcified gnis—granite

boulders, fragments snd rock particles set in a

lime matrix, preserved in joints in fhe fresh

sranite. Some 2 m aheove the intertidal platform

cul in granite there is a platform in aeolianite

consisting largely ef coalesced pools in the flat

floors of which granite is exposed.

At Cape Lahatr remarkably smouth plal-

forms in granite and developed on massive

joint blocks are exposed in the interlidal zone

at the base of the high oneolianite cliffs, hur ne

remnants of a former regolith have heen found

there.

4. Talia Caves, Western Eyre Peninsula

The Tale Caves atea provides the only

exposure Gf a gently dipping conglomeratic

sandstone of possible Mesozoic age (Smith &

Kamerling 1969), A platforr eroded across

the tilted sedimentary layers is developed close

to high tide level. A {hin layer of quartz par-

licles set in a lime matelx intervenes between

the acolianite and the still massive sandstone.

As the platform here corresponds in elevation

with the base of this impregnated and

cemented skeletal soil, an etch origin is again

indicaled,

5. Cape Carnoy, southern Eyre Peninsula

Platforms cul in orystalline rocks 4re

exposed at the base of towering acolianite cliffs

and upparently cmerge from beneath these

Pleistocene deposits at several sites in the Cape

Wiles-Cape Carnot area of southern Eyre

Peninsula: but they are jnaccessible. However,

at Cape Carnot and in a sector extending a few

kilometres northwards the coast is less Forbid-

ding and the origin of the platforms can be

investigated, Platforms in gneiss are swept by

the highest tides af Cape Carnot itself, but

only a few score metres to the north, a rego-

lith is preserved bencath acolianite the base of

which stands some 3-4 m above high tide level.

This section strongly suggests that the shore

platforms are eich surfaces from which both

ealcarenite and regolith have been stripped. At

Redbanks, a few kilometres further to the

north. the same relationship obtains between

high platforms in fresh gneiss and regolith

preserved beneath the aeolianite, the regolith

again occurring 3-4 m above high tide level.

The seyolith is only 2-3 m thick and is

ferruginous, It is not a laterite for there is no

distinct horizon development; this regolith. like

those described from Point Brown and Drum-

mond Point is most hkely related to the Koon-

gawa Surface of late Phocene and Pleistocene

age (Twidale, Bourne & Smith 1976).

G, Wemtworri: Point, southern Yorke Peninyula

In the rocky cliffs of Wentworth Point, on

southern Yorke Peninsula, both aeolianite and

granite are displayed. The unconformity

belween the two, which in peneral occurs about

half way up the cliffs, is irregular in some sec-

tors but remarkpbly horizontal and smooth

elsewhere, A regolith up to 2 m thick is

preserved jm places, though whether the

weathering to which it is telated dates from

before the deposition of the dune sand, or

whether ic has developed subsequent to the

burial of the granite is not clear, Nor is there

unequivocal evidence as to the nature of the

surface cul In gramte Is il of epigene origin,

nid if sa is it rented to the Koongawa Surface

SHORE PLATFORMS AND SEALEVEL CHANGES, GULFS REGION

surtace

corestones

in grus

a Fis aie

vaan 5 |

©beach boulders.

with lichen

A bpulder beach

notch at water level

step near pool

salt z ryatald in ae clefts

° high tide ievet

mean sea level+

WENTWORTH POINT

rampart

méan sea

lavel

WINDMILL BAY

platform

5- em.

Fig. 6a, Section through granite platform and cliff at Wentworth Point shown in Fig. Sb.

Fig. 6b. Composition section of Windmill Bay showing raised platform raised stack and raised beach,

and associated period of weathering? Or is it

an earlier Pleistocene shore profile, including

a small platform, which has been preserved and

weathered by virtue of the aeolianite cover?

The answers to these questions are not

apparent,

The granite surface has been cut into a

blocky cliff the upper part of which, just below

the unconformity, is intensely weathered, but

which is quite fresh below this 2 m thick zone

(Fig. 6a). The bluff gives way to an essentially

smooth platform, with pools, and protected on

the outer side by a blocky rampart. The plat-

form is at or just above high tide. level on one

headland but occurs a few metres higher on

others in the locality (Fig. 5b). All, however,

are in the spray zone. At some sites calcified

grus and corestones are preserved (Fig, 5c),

and aeolianite remains in one of several joint

controlled lapies developed on the sheer face

of a massive block of granite gneiss. Thus

although it can be argued that there has been

modilication of the granite surface since the

deposition and stripping of the aeclianite—and

the development of solution notches up to 15

cm deep in the limestone blocks that have

tumbled into pools on the platforms attest to

this—it is also clear that some, at any rate,

of the granite relief essentially predates the

aeolianite,

But the problem of the origin of the plat-

forms and associated features ig not resolved:

are they cpigene or marine? If the former one

would expect the usual assemblage of flared

slopes, widespread Rillen and platforms; the

flared backing slopes are absent suggesting that

the assemblage may be coastal, but the

evidence is not wholly convincing.

Modern platforms and the so-called “ten-foot”

stand of the sea

Shore platforms are developed within the

present tidal range in the acolianite, and in the

various crystalline and sedimentary rocks of

Precambrian and Cambrian age. Some

developed in older rocks are not evidently

coincident with the weathering front and so

differ from those earlier discussed in that they

cannot be explained as ctch forms, The plat-

forms cut in aeolianite must post-date the

material in which they are cut. Those eroded in

crystalline and Precambrian sedimentary rocks

are either related to modern sea level or they

predate the acolianite and have recently been

70 C. R. TWIDALE, JENNIFER A. BOURNE & NICHOLAS TWIDALE

Fig. 7a. Shore platform eroded in dipping Pre-

cambrian siltstone at Hallett Cove.

Fig. 7b. Platforms in aeolianite at Wellesley

Point. A—calcrete horizons; B—platform

eroded below calcrete; C—cliffs; D—

spray pool platform (-++ 3 m); E—inter-

tidal platform. B, D and E transect the

cross-bedding.

Fig. 7c. The “10-foot” platform developed in

oolitic Pleistocene limestone at Gym

Beach, Yorke Peninsula.

exhumed. But if the latter suggestion is correct

they must predate the last glacial. Why were

they not undercut and cliffed by the sea, and

dissected by streams during times of low

(glacial) sea level?

Most of the areas considered here are lack-

ing in surface streams due to the perviousness

and permeability of the calcarenite but in times

past the granitic and gneissic hinterlands would

surely have generated surface drainage. Com-

parisons with the Hallett Cove area seem to be

apposite. Here several small streams drain the

area immediately back from the cliffs. Water-

fall Creek is one of the larger, and it debouches

on to the northern end of the shore platform.

Now if the platform (Fig. 7a) predates the last

glacial, surely the Creek would have cut a

valley below it. It has not, and it can only be

surmised that the platform is related to the

present stand of the sea, Similar arguments

apply to the other intertidal platforms on

Yorke and Eyre peninsulas, The normal plat-

forms are of Holocene age and they are related

to the contemporary sea level; the dunes are

late Wisconsin in age.

If the platforms cut in crystalline rocks are

of postglacial age, they are contemporary with

those eroded in aeolianite. The platforms in

aeolianite are widespread but achieve the

greatest widths (i.e. development in a direc-

tion normal to the coast) in sheltered situa-

tions. For example in an unnamed bay behind

Speed Point northwestern Eyre Peninsula, the

shore platform in aeolianite (Fig. 8a) is about

30 m wide. That developed at the head of

Stenhouse Bay, on southern Yorke Peninsula, is

20 m across, in contrast with the 1-3 m widths

observed on promontories and headlands. In

each case the platforms transect the variable

cross-bedding of the aeolianite and so are

erosional: they are not the structural benches

or ledges of Jutson (1939) and Hills (1971).

The contemporary platforms eroded in crystal-

lines are about 10 m wide at most though the

platform cut in dipping Precambrian sediments

at Hallett Cove (Fig. 1) near Adelaide, extends

30 m from the base of the cliff (Fig. 7a). All

these platforms argue considerable rates of

erosion in postglacial time. They are extending

inland as a result of attack at the cliff base, but

they themselves are suffering undermining by

wave attack at the outer edge and also by the

development of pits (cf Jutson 1949a; Sprigg

1952). But they have all been eroded during

the past 6,000-—7,000 years, the maximum esti-

mate allowed for postglacial time (see Fair-

bridge 1961; Curray 1965; Shepard & Curray

1967).

These intertidal platforms pose another diffi-

culty, for some workers take each individual

platform to represent a separate and distinct

SHORE PLATFORMS AND SEALEVEL CHANGES, GULFS REGION 71

Fig. 8a. Shore platform in calcarenite in sheltered bay behind Speed Point. Note raised rim of outer

edge of platform.

Fig. 8b, Platforms in aeolianite on the exposed coast at Speed Point showing the narrow intertidal

feature and the high platform, the so-called “10-foot” platform, cutting away the bedding in

the aeolianite. Note the line of debris on the beach in the foreground at the same elevation as

the high platform.

stand of the sea (e.g. Teichert 1947). Others

consider that whole flights of platforms are

developing simultaneously in relation to the

contemporary sealevel, and believe that a simi-

lar multiplicity of platforms evolved in relation

to former stands of the sea (e.g. Jutson 1930,

1949b; Hills 1949).

The question is complicated by structure, for

on many coasts lithological differences, and

bedding, have been exploited by marine agents

to produce benches or flats at different levels.

For instance at Daly Head, on the “toe” of

Yorke Peninsula, the platform developed in

biotite schists is 1-2 m lower than that evolved

on the adjacent aplite. Again, both at Wellesley

Point near Elliston and at Robe, in the South-

east district (Fig. 1) a calcrete formed at the

surface of an ancient dune finds expression in

a distinct bench the elevation of which changes

parallel to the coast, simulating as it does the

slope of the former dune. These are coastal

ledges in the terminology of Hills (1971). Jut-

son (1930, 1939) and Bird & Dent (1966)

have cited other examples of structural con-

trol.

Nonetheless, when structural effects are fully

accounted for there remains within the present

tidal zone a multiplicity of platforms extending

through a considerable vertical range and

clearly cut across the local cross-bedding in the

aeolianite, and across jointing and foliation in

crystalline rocks. Many platforms have

developed within the spray zone as a result of

pool weathering (see Wentworth 1938, 1939;

Hills 1949; Kaye 1959). Such flights of plat-

forms are especially well displayed on the

aeolianite: there are for example, three such

at Cape Wellesley which faces the Great Aus-

tralian Bight (Figs 2b and 7b). Each of these

is currently suffering alteration, the lower one

72 C. R. TWIDALE, JENNIFER A. BOURNE & NICHOLAS TWIDALE

Fig. 9a. General view of Admiral Arch showing

the dipping Cambrian schists below and

the aeolianite, with stalactites, above.

Fig. 9b. Unconformity between schists and

aeolianite with schist cobbles and

boulders embedded in the base of the

dune rock.

Fig. 9c. Raised stack in granite at Windmill Bay.

by wave attack, the second by rapid alteration

of wetting and drying, the higher by occasional

wetting and drying by rains and by spray.

However, the presence of sand, shells, and salt

crystals in pools on the highest platform shows

that they receive spray from time to time.

Thus platforms appear to be developing at

the present time in a range that extends up to

8 m above present high tide level. But what,

then, of the so-called “ten-foot” level, of Recent

but relic age, advocated by some workers, for

example Crocker (1946b)?

The problem is very well illustrated at Speed

Point, on the northwest coast of Eyre Penin-

sula. There, on the open coast as well as on

the shores of the sheltered inlet platforms are

well developed at about mid-tide level but there

is also a prominent platform preserved on Jow

calcarenite promontories on the exposed coast

(Fig. 8b). However, the head of the steeply

sloping beach is level with that of the platforms

which stand 3 m (10 feet) above the mid-tide

platform. The presence of debris at the beach

head shows that waves reach to that elevation

so that at least occasionally the high platforms

are affected by wave action; and they are cer-

tainly within reach of spray.

At Brown Point, Crocker’s (1946b) type

section on the western shore of Yorke Penin-

sula, the evidence for the ten-foot platform

being wholly a relic form is equally unconvinc-

ing. The platform extends only 2-3 m_ back

from the cliff edge. Solution hollows and pools

attest the effectiveness of spray water in

weathering the limestone and though the plat-

form is clearly being undermined and

destroyed by wave action it is equally obvious

that the platform is also extending and being

smoothed by pool weathering.

Similar situations obtain at Port Rickaby,

Point Turton (Clark 1928) and Gym Beach

(Fig. 7c), all on Yorke Peninsula (Fig. 1),

and at Cape Northumberland in the Lower

Southeast to name four examples. Are these

3 m platforms relict or modern? There are

three possibilities:

1, That they are relict features, date from early

Recent times and are now essentially being

destroyed, but simultaneously are suffering

planation by pool weathering.

2. That they are contemporary forms which are

due to pool weathering, but which are at the

same time being undermined by wave attack,

illustrating that different marine processes

act at different levels.

3. That some of the 3 m platforms are relict

and others modern.

The evidence favours the second of these pos-

sibilities.

Evidence of higher stands of the sea

Is there then, any definite evidence of higher

stands of the sea in the area under discussion?

Cambrian Kanmantoo schists that are exten-

sively exposed in the cliffs and platforms of the

north coast of Kangaroo Island also occur at

the western extremity, near Cape du Couedic.

At Admiral Arch the schists occur beneath the

SHOKE PLATFORMS AND SEALEVEL CHANGES, GULFS REGION 73

Pleistocene acolianite in the coastal cliffs and

in the lower part of the natural arch (Fig. 9a

und b), The bay heads between promontories

are occupied by shingle beaches composed of

the boulders and cobbles of the Kanmantoo

schist. Within the arch a former cobble and

shingle beach is preserved at the unconformity

between dipping schist and limestone (Fig.

Oh). It stands 5-6 m above high tide jevel and

the present shingle beaches, und surely indi-

cites the stand of the sea previois to the

deposition of the calearenite dunes, probably

during the last glactal,

Windmill Bay is located near Cape

Willoughby on the southeastern coast of Kay-

garoo Island (Fig. 1). In the Bay itself there

is a coarse beach of granite boulders (Fig, 6b)

some of which are fitted (Hills 1970), These

extend 5-6 m up the slope beyond the present

beach, Those on the slope are covered with an

urange fichen (Nanthoria ectanea (ACH)

RAS ex R, FILSON) ond are scattered over

the surface,

The boulders are probably former corestones

derived from the Weathered granite that is seen

in cliff sections a few metres to the southeast

and which probably underlies the slope behind

the beach, But in front of the rocky coast

backed by the granite cliffs is a platform stand-

ing 5-6 m above high tide level, and on it an

old stack with a ootch on its seaward side

also 5-6 m ahove sealevel (Fig, 9c), There is

no aeolianite exposed in the cli? but it rests on

the granite in the hill immediately to the north.

Some evidence of a similar pre-avolianite

platlorm standing 3-4 m above present sea-

level occurs on the north coast of Kangaroo

Island at Boxing Bay and Cape Cassini, where

dedlianite rests unconformably on a surface

eroded in folded Cambrian quartzites. The sur

face al and adjacent to Boxing Bay may be u

yhorve platform pf pre-acolianite age und that

around Cape Cassini could be a combination

of shore platform and coastal plain of similar

age. If so both planate features stand 3-4 ni

above present high tide level and represent a

former sealevel of that order,

Conclusions

The evidence from western Eyre Peninsula

und southern Yorke Peninsula suttyests hist

many of the pltlofras cut in granite and gneiss

are etch surfaces, or exposed weathering

fronts which lorturtousty lie within the present

tidal or spray zone. Shore platforms cut in

fresh granite are of limited extent, as postu-

lated by Jutson antl Hills.

Those granite platforms that oceur at mid

tide level, as at Smooth Pool, are not indicative

of a pre-acolianite stand of the sea similar to

the contemporary sealevel. And those that

stand at High-tide level, or 4 few metres above

it do not represent a former higher stand of

the sea. Pre-acolianite regoliths preserved on

southern Eyre Peninsula may be related to the

same haselevel or sealevel to which the epigenc

surfaces were graded.

These remarks apply equally to the benches

essentially associated with the unconformity

berween the Pleistocene aeolianite and the older

rocks, Thus the extent and perfection of the

granite and gneiss platforms is duc to the effec-

tiveness of weathering in carlier times, The

only granite platforms possibly due to con-

temporary jnavine plocesses are at Wentworth

Point on southern Yorke Penmsula, and even

there the evidence ss equivocal.

Of those platforms cut in acolianite and in

alder, folded, sediments und due wholly to

marine processes, mary occur wilhin” the

modern tidal or spray zone, and are develop-

ing simultaneously at the present time. The so-

called ten-foot platform may be a relic feature

which is suffering modification at present; but

there is evidence that it too is & Contemporary

form. Finally there is evidence of a stand of

the sea 5-6 m higher chan present dating from

pre-acolianite (probably pre late Wisconsin)

times, and no indicahon of significant tectonism

having allected Kangaroo Island in the time

\iterval that has elapsed between the formation

of the now ramsed beaches and other coastal

forms and ihe present.

Acknowledgments

The writers wish to thank Emeritus Profes-

sor E. §. Hills for a ¢ritical reading of the

paper in draft stage, and Mr N, N. Donner of

the SA, State Herbarium for identifying the

lichens moeatioued in the text. The field work

on Yorke Peninsula was carried out with the

ussistance of a grant from the Royal Socicty

of South Australia Endowment and Scientific

Research Fund, for which the recipients are

graceful.

74 CR. TWIDALE, JENNIFER A. BOURNE & NICHOLAS TWIDALE

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REVISION OF THE COMPOSITE SPECIES LIMA BASSI TENISON

WOODS (MOLLUSCA, BIVALVIA)

BY M. F. BUONAIUTO

Summary

The composite species Lima Bassi Tenison Woods is revised. Among the forms referred to Lima

bassi (ranging from late Eocene to Pliocene), four species are recognized: the Late Eocene L.

maslinensis sp. nov., the early Miocene L. bassi s. str., the Middle Miocene (Batesfordian) L.

morganensis sp. nov., and the Late Pliocene L. elianae. A neotype and paraneotypes of Lima bassi

from the type-locality are established.

REVISION OF THE COMPOSITE SPECIES L/MA BASS] TENISON WOODS

({MOLLUSCA, BIVALYIA)

by M. F. Buonaruro*

Summary

Buowato, M. F, (1977) Revision of the composite species Lima bassi Tennison Woods

Mollusca, Bivalvia). Trans. R. Soc, §. Aust, 101(3), 75-83, 31 May, 1977,

The composite species Lima bassi Tenison Woods is revised. Among the forms referred

to Lima bassi (ranging from Late Eocene to Pliocene), four species are recognized: the Late

Eocene L. maslinensis sp. nov., the Early Miocene L. bassi 5. str., the Middle Miocene (Bates-

fordian) L, merganensis sp. nov,, and the Late Pliocene L. elianae, A neotype and paranco-

types of Lima hassi from the type-locality are established,

Introduction

During the present revision of the Eocene

Mollusca from the type section of the Aldingan

stage, different species revealed themselves as

composite: Lima basst Tenison Woods is one

of them, Past authors grouped at least three

distinct forms in it; the Longfordian Lima

baysi, (he Late Eocene L, maslinensis sp. nov.,

the Batesfordian L. merganensis sp, nov., and

a fourth Pliocene form L, elianae from Dry

Creek Sands.

Apart from the revision of Lima bassi and

the description of three new taxa, it is neces-

sary to establish u neotype and paraneotypes of

L, bassi, Since Ludbrook (1967) revised the

Johnston and Wood's types, it is common

knowledge that many of these types were lost

during the first half of this century, The holo-

type of Linta bassi is one of them,

Although authors quote several localities, the

only localities considered here are those from

which the specimens examined were obtained,

Definitions of the parameters here meastired

(after Cox, Nuttall & Trueman, in Moore

1969).

Hi —height of valve distance between two planes,

parallel to cardinal axis, perpendicular to

commissure plane, and tangent to umbonal

and ventral ends of valve.

Lt length of valve as distance between 2 planes

perpendicular to cardinal axis and tangent

io anterior and posterior ends of valve.

Lpa—tength of posterior auricle, as distance

hetween (wo planes tangent fo beak and to

posterior end of auricle and normal to car-

dinal axis,

Lad—length of anterior auricle,

defined as the ahove parameters.

Lea—leneth of cardinal area as distunce between

two planes tangent to auricles’ ends and

perpendicular to cardinal axis.

Hr —height. of resilifer as distance between two

parallel planes, parallel to cardinal axis, and

tangent to its upper and lower ends.

Lr —length of resilifer, as distance between two

planes, perpendicular to cardinal axis, and

tangent to its anterior and posterior ends.

Ts —thickness of valve, as distance between two

parallel planes: former tangent to commis-

sure line, and latter tangent to outer valve

surface.

Hea—height of curdinal area, as distance between

two planes parallel to cardinal axis and tan-

gent to its upper and lower ends.

Tea —dislunce between two parallel planes: for-

mer tangent to beak; latter tangent tu com-

missure line.

analogously

Standard ratios were calculated. The: ratio

Tcea/Ht = Tgé represents the tangent of the

angle ¢ between the geometric generating curve

as defined by Raup (1966) after Stasek

(1966) where the biological generating curve

coincides with the growing edge of the valve,

and the geometric generating curve is the inter-

section of the valve with a plane containing the

coiling axis and tangent to the ventral end of

the biological curve. The angle 4 determines

the degree of maximum opening breadth

between valves.

* Department of Geology and Mineralogy, University of Adelaide, North Tce, Adelaide, 8, Aust, 5000,

76 M. F. BUONAIUTO

FIGS 1-7

REVISION GF THE COMPOSITE SPECIES LIMA BASSI 7

Collections. SAM: South Australian Museum.

GSSA: Geological Survey of South Australia,

Systematic descriptions

CLASS: BIVALVIA Linné, 1755

SUBCLASS; PTERIOMORPHIA Beurlen,

1944

ORDER; PTERIOIDA Newell, 1965

SUBORDER; PTERIINA Newell, 1965

SUPERFAMILY: LIMACEA = Kafiresque,

1815

FAMILY: LIMIDAE Ratinesque, 1815

GENUS: Limo Bruguiére, 1797

SUBGENUS: Lima s.str,

Lima (Lima) bassi Tenison Woods, 1877

FIGS 1-2, 11-13, 17-4

1877 Lime bassil Tenison Woods, p. 112. 1886

Lima bassi—Tate, p, 117 (pars). 1955 Lime

haysi—Ludbrook, p. 35 (pars).

Neolype: RY senile, figs }-2 (SAM 16343/1).

Paraneotypes: 6 RV, 2 LV adults, figs [7-24,

(SAM P18344/2-8),

Strarigraphle Jocation: Freestane Cove Sand-

stone (Longfordian).

Type locality: Table Cape, Tasmania, Bass

Basin (Freestone Cove).

Description; Shell rather thin, subtrigonal, very

inequilateral, liltle inflated, higher than long:

uibones. with acute small prosogyrate beuks.

Margins: antero- and postero-dorsal straight,

the Jatler longer; anterior very long, concave:

posterior very long, concave near the umbo,

convex and very elliptical to the Ventral; ven-

tral very elliptical. Margin connections: untero-

dorsal—-anterior and posterodorsal—posierior

very angular, anterior/ and posterior—ventral

imperceptible. Auricles triangular, small, the

anterior reduced, Longitudinal shell section

regularly but weakly convex. Regions: pos-

terior and dorsoventral convex and gently

declivous; anterior convex and gently declivous

to the dorsum, subconeave and vertical at the

margin. Region connections: imperceptible, the

wo diferent parts of che anterior by & sharp

nm.

Cardinal area narrow, triangular; resilifer

broad, triangular, concave. Hinge taxodont

with two small longitudinal teeth on the

auricles. Interior radially ribbed; pallial line

imperceptible; monomyarian with posterior

adductor sear faint, rather high and marginal,

8-shaped, broader in the upper part, Conimis-

sure region crenulated except near the hinge.

Crnament: Prominent scaly radial ribs with

equal U-shaped interspaces, Between the inter-

spaces concentric flat microcostae; in juvenile—

adiilt the anteriors and the posteriors conver-

gent to the dorsoventral where they overlap

with a shagreen patiern; in adult regularly con-

centric microcostae; in seniles very fine growth

lines only,

Anterior marginal region concentric costae

and weak radial ribs.

Auricles: anterior strong concentric costacy

posterior with strong radial ribs,

Dimensions (mm): (see Table |).

Observations; Tate commented on differences

among specimens from Table Cape, Morgan,

Aldinga and Adelaide. On the basis of concen-

trié Microormaments he distinguished from L.

hassi 5. str. a var. A for the specimens from

Morgan, and «a var. & for the specimens from

Aldinga and Adelaide Bore, These varieties are

here rvised to species as L. morganersis and L,

maslinensis respectively.

Parancotype 2 displays in neanic-juvenile

slages concentric microornament similar to that

of L. maslinensis, abruptly passing to the regu-

lar concentric microornument. Io adult senile

stages secondary radial microriblets can

develop in some interspaces. The other para-

neotypes display vanability in Ht/Lt ratio and

in morphology (Figs 17-22), In reference to

the type-locality, Johnston (1877) quoted a

Lime squamesa in the “Crassatella Beds”, Fur-

ther. Johnston listed L. bass? = L. squenrosa

Lamarck, Banks (ja Gill 1962) revised and

redefined Johnston's “Crassatella Beds" as the

Freestone Cove Sandstone, attributing to it a

Late Oligocene age. Quilty (1966) and Lud-

hrook (1967, 1973) gave evidence of a Long-

fordian age for Table Cape Group of which

the Freestone Cove Sandstone is part, on the

basis of both henthonic and plantonic fora-

Figs 1-2, Lina bassl, neotype, Freestone Cove, Longfordian: (1) dorsal view, (2) tnterior view (x115),

Figs 3-4, L. piorganensis sp. nov., Batesfordian: (3) holotype (GSSA M 3138), near Morgan,

dorsal view (x2.2); (4) paratype (SAM 7T982 EB), Murray Cliffs, interior view {x2.55).

Figs 3-7, &. maslinensis sp. nov. Adelale (Kent Town) Bore, Alitingan; (5) holotype, dor-

gal view (42.2): (6) holotype, interior view (42.2). Paratype (SAM P 18344), Maslin Bay;

17) dorsal view {x2.2).

FIGS 8-16

ie)

eo)

KEVISION OF THE COMPOSITE SPECIES LIMA BASSI ry)

TABLE |

Dimensions (in mm) and ratios of Lima basst

Specimens Ht Li Loa Lan Hr Lr Ts Tea Lea Hea a

Neotype sig) 4s.ks 7.0) sCiTMNSSS—C“‘«‘i;«SSOC*d.:CiSC 8"

Parnnectype 1 23.50 19.35 2,75 245 210 1.90 3.10 OTS 5.15 1.45 1°49°

Paraneotype 2 DAF — 4.60 2.90 2,20 1,70 4.55 13s 7,50 1,95 2°50’

Pararsotype 3 38.17 32,90 5.25 3.70) 3.50 3.00 7.35 1.20 8,95 2.60 1°47"

Paraneatype 4 23.40 22.95 3.75 2.45 205 245 490 ),85 6.70 1,60 1°41’

Tzi—

Specimens List Laa/Lpa Laa/bea. Lpa/bea Lr/Hr Ts/Lt Ts/Hte Tew/Ht Hea/Lea

Neotype -BOS9 HILG 3879 6129 1.1316 1734 1500 02830 2339

Paruneotype | 8234 BOLO ATRS 5371 048 1602 1319 03191 2832

Paratieotype 2 _— 6304 3867 6133 1,294} — L676 04972 2600

Paraneotype 3 8601 7048 AN34 5366 M57t 2234 A921 03137 2905

Parateotype 4 7969 6533 3657 5597 1.4951 2135 A701 02951 2388

minifera. Hutton (1887) synonymized L, hasyi

with &, wvelerata Hulton, 1873 (Boreham

1965) because “Mr Woods’ name stands as

mine is incoryect’. Later authors such as

Suter (1914) accepted Hutton’s name.

Probably aller Hutton, Tate (1899) quoted ZL,

basei occurring also in New Zealand. Finlay

(1924) quoted L, celerata as one of the New

Zealand species corresponding to L. Aasyi.

However, un Awamoau senile specimen of L,

rolorata from Otago, in the Department of

Geology and Mineralogy at the University of

Adelaide, displays specific differences as trape-

zojdal-shaped broad radjal ribs with broader

mlerspaces, as Only fine growth aes in juvenile

stages. and also radial oblique, very fine stna-

titns tn adult and senile siages,

Lima maslinensis sp, nov.

FIGS 5-10

1886 Lima hassi var, B Tate, p, 117, pl. 8, fig.

la-c.

Derivation of Name: From Maslin Bay, locality

of the lowest recorded occurrence of this form.

Holorypes T983D, holotype of var, B, figs 5-6,

Paratypes: T983 A-C, E.

Yype Leeality; Old E & W. Dept Kent Town

Bore, Hd Adelaide, sect, NE Parklands No, |4

Stratigraphic Ranges Aldingan (Late Eocene)

{Ludhrook 1973)

Collections: SAM TYSS A-E, P18344_

Material: 26 specimens (8 LV, 4 RV, 14 VV)

badly preserved, 5 specimens from Tate's Col-

lection (2RV, 3LV)

Description: As L. bassi. Differences: stronger

teeth, the anteriors longer, the posterior

triangular.

Ornament; Primary radial scaly costae with

reclungular section and with equal U-shaped

interspaces. In the interspaces, fine flat trans-

verse microcostae in the anterior and posterior

regions, convergent to the dorso-umbonal; in

the dorsoventral region the microcostae over-

lap with a shagreen pattern, Anterior marginal

region with numerous fainter radial spiny ribs,

Auri¢les; anterior with concentric costae,

posterior with concentric costae and faint spiny

radials.

Dimensions (mm)!

T9853 D—Ht, 37,5; Lt, 25.45; Lpa, 4.70; Laa,

3.55: Hr, 2.30; Lr, 2.90; Ts, 5.65; Tea, 1,45;

Hea, 2,70; Lea, 8.25.

Ratios: T983° D—Lt/Ht, .6876; Laa/Lpa,

7553; Laa/Lea, .4303; Lpa/Lea, 5697; Lr/

Hr, 1.2609; Ts/Lt, .2220; Ts/Ht, .1507; Tga

= Tea/ Ht, .03866; Hea/ Lea, 3273,

Observations. Tate initially separated this form

from Lima bassi Tenison Woods as var. B.

Teeth, microcostal pattern, posterior auricle

and unterior marginal ornaments distinguish

this form at specific level from ZL. bassi,

Tate's holotype of the variety is here chosen

as the holotype of L. maslinensis, although

from it is the subsurface and broken into two

neat pieces, it is the only well preserved speci-

men.

Distribution. St Vincent Basin; Adelaide Plams

Sub-Basin, Kent Town Bore; Willunga Sub-

Basin, Maslin Bay,

Figs 8-10, L. miastinensix sp. nov; (8) anterior ventral ornament (x13); (9) posterior ventral ornaments

(x12); (10) dorsoventral ornaments (x12), Pigs 11-13, Lima hassi, paraneotype 8, Freestone

Cove, ornament: (11) anterior ventral (x13): (12) dorsoventral (x13); (13) posterior ven-

tial (X12), Bigs 14-16. Lima rherganensis sp. nov.,

holotype, ornaments; (14) dorsoventral

¢alQis (15) juvenile dorsoventral afd posterior auricle (x9.5); (16) anterior veniral (x9,5).

M. F. BUONAIUTO

FIGS 17-24

REVISION OF THE COMPOSITE SPECIES LIMA BASSI SL

Lima morganensis sp. nov.

FIGS 3-4, 14-16

1886 Lime bassii var, A Tate, p. 117. pl. 5, fig.

Ra-b, 21897 Lima passil Harris, p, 310 (non

Tenison Woods}.

Derlvation of Name: From Morgan, the town

nearby, after which Morgan Limestone was

named.

Holotype: GSSA M3138, fig. 4.

Puratypes. T982 A-L, fig. 4.

Stratigraphic Location: Cadell

(Batesfordian) (Ludbrook 1973),

Type-Lecaliry; 6.4 km § of Margan, type sec-

tion of Cadell Marl, section G, Hd Cadell

(Ludbraok 1961).

Material: 11 specimens from Tate's callection

(7 RV, 4 LV): 1 RV from GSSA Collection

(Holotype).

Description. As L. bassi.

Ornament: Primary radial subtriangular scaly

ribs With equal V-shaped interspaces; long very

melined chevron-shaped microriblets covering

ribs and interspaces, with their head on the ribs

onentated to the umbo. Posterior auricle with

more marked concentric costae and weak spiny

radial ribs. Anterior auricle with concentric

costae and weak beaded radial ribs,

Dimensions (rm) '

T982 A—Ht, 32.20, Lt, 26.00; Lpa, 5,55; Laa,

3.85; Hr, —: Lr, —; Ts, 4.70; Tea, 1.35; Lea,

9,40; Hea, 2.10; 4, 1°24'. T982 E—Ht, 20,50;

Lt, 15.25; Lpa, 3.60; Laa, 2.70; Hr, 2,20; Lar,

L.50; Ts, —: Tea, —; Lea, 6.30; Hea, 1.95;

“.—

Ratios;

T982 A—Lt/Ht, .8074; Laa/Lpa, .6937; Laa/

Lea, .4085; Lpa/Lea, 5904; Lr/ Hr, —; Ts/ Lt,

1808; Ts/Ht, .1460; tga = Tca/Ht, 04192;

Maris Lens

Hea/ Lea, 2234. T982 E—Lt/ Ht, .7439; Laa/

Lpa, .7500; Laa/Lca, 4286; Lpa/Lea, 5714;

LrfHe, .o81S; Ts/1.1, —; Ts/Ht, —: tga =

Tea/ Ht, —; Hea/ ca, ,3095.

Observations. Tate initially distinguished his

form as oa variety of Lima bassi Tenison

Woods. Shape of the ribs and imterspaces

ornaments of posterior auricle, anterior mar-

ginal region and interspaces and radial costae

separate this form from L- bassi.

The specimen T9282 A is the holotype of

Tate’s var. A. The original illustration of Fig.

8a is inverted, The specimen from GSSA Col-

leclion was chosen as holotype because of ils

good tapographic and stratigraphic location,

Lima elianae sp. nov.

FIGS 28-41

1955 Lima bessi—Ludbrook, p, 36 (pars),

Derivation of Name; After Dott, Eliana Gar-

barina (Mrs Buonaiuto), the author's mother.

Holotype: GSSA M 2384, figs 25-26.

Paratypes; GSSA M 158, figs 28-31; SAM

P19210, Fig. 27.

T'ype-Locality: Observation Bore D, hd Port

Adelaide, St Kilda, 81,38-83.5 m depth,

Type-Formalion: Dry Creek Sands, Yatalan,

Material: The holotype (LV) and an adult

(SAM P 19210) with damaged margins and

a specimen (GSSA M 159) with the ventral

part broken.

Description, As £, hasst,

Ornament: Primary radial subquadrangular

cosiae with equal U-shaped interspaces; in

neanic stage prominent concentric microcostae,

In the adult stage short chevron-shaped con-

centric microcostae with the head to the ven-

tral in the interspaces; the V-microcoslae can

be substituted by narrow belts of normal con-

centric microcostae, meanwhile in the senile

stage they are entirely substituted by fine

growth rugse and fine radial oblique micro-

plicae

Observations. The specimen of L. elianae frorn

Abattoirs Bore is that quoted by Woods

(1931) as Austrolima bassi, The form referred

hy Tare (1890) to L. bassi from Dry Creek

Bore is at present unlocated but almost cer-

tainly belongs to L. efianae,

Unfortunately the only three specimens

available are both from bores and both

damaged. The senile from Observation Bore

1s chosen as holotype because it is the only one

with a sure stratigraphic location.

L. elianae, L. basst, and , maslinensis are

all characterized by subquadrangular radial

costae and chevron-shaped concentric micro-

costae. They might represent a lineage, but the

material available is inadequate to be certam,

eS re ee a EEEEEESEEEEEEEEEene aR

Fins 17-24, Lima bassi, paraneatypes, Freestone Cove; (17) Paranealype 3 (X1,15); (18) Paraneotype

7 (41-7); (19) Paraneotype t (x24);

(x1.15); (22) Paraneotype 4 (x2,2); (23) Paraneotype ® (%2,2):

(x2.4).

(20) Puraneotype 6 (x2.1); (ZL) Parancotype 3

(24) Paraneotype 2

82 M. F. BUONAIUTO

FIGS 25-31

Figs 25-26. Lima elianae sp. nov., holotype: (GSSA M 2384), Observation Bore D, Late Pliocene,

views (x1.0); (25) dorsal; (26) interior, Fig. 27. Lima elianae Sp. nov. paratype (SAM P

19210), Abbattoirs Bore, Late Pliocene, dorsal view (x1.4). Figs 28-31. Lime elianae sp.

nov., paratype (GSSA M 159), Munno Para Bore, Late Pliocene; (28) dorsal view (x1,95).

Ornament; (29) dorsal juvenile transitional to adult; (30) dorsal, adult; (31) anterior,

adult.

Other Localities: Adelaide Plains Sub-Basin.

Abattoirs Bore; bore hd. Munno Para, Sect.

4251, 72,5-78.0 m depth.

Stratigraphic Range: Yatalan (Late Pliocene),

Acknowledgments

! am very grateful to the Director of Mines,

South Australia, and to the Director of the

South Australian Museum, Adelaide, for the

loan of the material here studied; to Dr N. H.

Ludbrook for continued advice; and to Dr B.

McGowran for reading the manuscript.

The work was carried out in the Department

of Geology and Mineralogy, University of

Adelaide, during tenure of a University

Research Grant.

References

Bangs. M. R. (1962) Cainozoic. Marine Succes-

sion. Zn Spry, A. & Banks, M. R. The Geology

of Tasmania. J. Geal. Soc. Aust., 9(2), 233-

236.

Borewam, A. U. E. (1965) A revision of F. W.

Hutton's Pelecypod species described in the

Catalogue of Tertiary Mollusca and Echino-

dermata (1873), N.Z. Geol. Surv. Paleont.

Bull., 37, 1-84.

DennantT. J, & Kitson, A. E. (1903) Catalogue of

the described species of fossils (except Bryo-

zoa and Foraminifera) in the Cainozoic fauna

of Victoria, South Australia and Tasmania,

Ree. Geol. Sury, Vict,, 1(2), 89-147.

REVISION OF THE COMPOSITE SPECIES LIMA BASSI 83

Finuay, H. J. (1924) List of recorded relation-

ships between Australia and New Zealand

Mollusca. Aust. Assoc. Adv. Sci., Rep. 16th

meet., 332-343,

Harris, G. F. (1897) Catalogue of Tertiary Mol-

lusca in the Department of Geology, British

Museum (Natural History). Pt. 1, The Aus-

tralasian Tertiary Mollusca. (London. )

Hutton, F. W. (1887) Notes on some Australian

Tertiary fossils. Proc. Linn. Soc. N.S.W., 2nd

ser., 1, 481-482.

JOHNSTON, R. M. (1877) Further notes on the

Tertiary Marine Beds of Table Cape. Pap.

Proc. Rep. R. Sac. Tasm. (1876), 79-90,

Lupsrook, N. H. (1967) Tertiary Molluscan types

from Table Cape in the Tasmanian Museum,

Hobart. /bid, 101, 65-69.

Lupsrook, N. H. (1961) Stratigraphy of the Mur-

ray Basin in South Australia. Bull. Geol.

Surv. S. Aust., 36, 1-96.

Lupsrook, N. H. (1955) The Molluscan fauna of

the Pliocene strata underlying the Adelaide

Plains, Pt. Il. Pelecypoda. Trans. R. Soc. S.

Aust., 78, 18-87.

Moore, R. C. (1969) Treatise on Invertebrate

Paleontology. Pt. N. Mollusca, Bivalvia.

(Univ. Kansas & Geol. Soc. Am.)

Quitty, P. G. (1966) The age of Tasmanian

Marine Tertiary rocks. Aust. J. Sci., 29(5),

143-144.

Raup, D. M. (1966) Geometric analysis of shell

coiling. General problems. J. Paleont., 40(5),

1178-1190.

SrasexK, C. R. (1963) Geometrical form and

gnomonic growth in the bivalved Mollusca, J.

Morph. 112(3), 215-229,

Suter, H. (1914) Revision of the Tertiary

Mollusca of New Zealand, based on type

material. Pt 1. N.Z. Geol. Surv. Palaeont.

Bull., 2.

Tate, R. (1886) The lamellibranchs of the Older

Tertiary of Australia. Pt 1. Trans. R. Soc. S.

Aust., 8, 96-158.

Tate, R (1899) A revision of the Older Tertiary

Mollusca of Australia. Pt 1. /bid, 249-277.

Tate, R. (1890) On the discovery of Marine

deposits of Pliocene age in Australia. Ibid,

23(2), 172-180.

Woops, J. E. TENison (1877) Notes on the fossils

referred to in the foregoing paper. Pap. Prac.

Rep. R. Soc. Tasm. (1876), 91-116.

Woops, N. H. (1931) Pelecypoda from the Abat-

toirs Bore, including twelve new species.

Trans. R. Soc. S, Aust., 55, 147-151.

PLEISTOCENE FROGS FROM CAVES AT NARACOORTE, SOUTH

AUSTRALIA

BY M. J. TYLER

Summary

Pleistocene cave deposits at Naracoorte, South Australia, have yielded 166 frog ilia assigned to five

species (two questionably) now found in southeastern Australia. They represent the first Pleistocene

frogs known in Australia. The significance of the absence of any member of the Litoria aurea

species complex in the Naracoorte Pleistocene fauna is discussed.

PLEISTOCENE FROGS FROM CAVES AT NARACOORTE, SOUTH AUSTRALIA

by M. J. TYLER*

Suminary

Tyiter, M. J. (1977) Pleistocene frogs from caves at Naracoorte, South Australia. Truss.

R. Soe, S. Aust, 101(3), 85-89, 31 May, 1977.

Pleistocene cave deposits at Naracoorte, South Australia, have yielded 166 frog ilia

assigned to five species (two questionably) now found in southeastern Australia, They repre-

sent the first Pleistocene frogs known in Australia, The significanve of the absence of any

member of the Literia aurea species complex in the Naracoorte Pleistocene fauna is discussed,

Introduction

It is generally accepted that glacial periods

during the Pleistocene were associated with

moist climatic conditions. Por the frogs of Aus-

tralia these moist periods provided (he oppor-

tunity for them to colonise or cross formerly

arid arcas and, as a result of custatic changes,

reach previously isolated islands on the exposed

continental shelf.

Until recently there have been no known

fossil frogs in Australia, and the solitary species

recorded (Australobatrachus ilius Tyler) is of

mid-Miocene age (Tyler 1974, 1976). There-

fore, to date, information about the frog fauna

during the glacial periods has been interpreted

from current geographic distributions and from

deduced patterns of speciation.

In fate 1974 and carly 1975 Dr R .T. Wells

permitted me to examine boxes of sievings

from Victoria Cave at Naracoorte, South Aus-

tralia. ‘The site had provided vast quantities of

late Pleistocene vertebrates reported by Smith

(1971, 1972), Van Tets (1974), Van Tets &

Smith (1974) and by Wells (1975), The exist

ing frog fauna within and surrounding the

Naracoorte area is well known (Tyler 1966;

Woodrulfe & Tyler 1968), and the site was

therefore ideal for an intensive search for fossil

frogs of limited antiquity,

The material provided by Dr Wells was exa-

mined for the presence of ilia: bones previously

demonstrated to be of considerable value tn the

identification of genera and species, and pre-

viously surveyed to establish the characteristics

of Australian species (Tyler 1976),

Following the recovery of numerous ilia in

the Victoria Cave deposit, additional specimens

Were oblained from the adjacent locality of

Henschke’s Quarry Cave. Subsequently Mr

N. Pledge located further specimens from

Henschke's Quarry Cave sievings.

In reporting the nature, extent and identity

of the material obtained, the object is to estab-

lish the existence of the first known Pleistocene

frog fauna in Australia, and {0 explore the

palueoclimatic and biogeographic implications

of its nature,

Methods

Descriptions of the Victoria Cave site and of

the techniques used for separating the fossils

from the silt appear in Smith (1971) and Wells

(1975), Rapid isolation of the ilia from the

bone fragments and particulate matter involved

the establishment of a search image, In prac-

tice the only macroscopic confusion was with

some small vertebrate ribs.

As yet in Henschke’s Quarry Cave only fos-

sil birds have been reported (Van Tets 1974).

Descriptive terminology of the ilium is that

of Tyler (1976), With the object of extrupolat-

ing the size of the donor animals from the ilial

fragments, various measurements were recorded

of bones from existing species. As a result an

association was found to exist between the body

length, and the distance separating the anterior

limit of the dorsa) acetabular expansion from

the inferior limit of the ventral acetabular ex-

pansion (DAE-VAE) as depicted in Figure 1.

All fossil material reported here has been

deposited in the South Australian Museum, and

* Department of Zoology, University of Adelaide, North Tce, Adelaide, S$. Aust, 5000,

86 M. 1. ‘TYLER

DAE

L-

VAE

Fig. 1. Acetabular portion of generalised frog

ilium illustrating measurement of span

between dorsal acetabular expansion

(DAE) and ventral acetabular expansion

(VAE).

the registration numbers are those of its Palac-

ontology Department.

Fossil fauma

Family HYLIDAE

Litoria ewingi (Duméril & Bibron)

Henschke’s Quarry Cave: P. 18878, P. 19505,

2 left ilia; P. 18940, P, 19504, 3 right ilia.

Victoria Cave: P, 16801, 21 left iha; P. 16802,

12 right ilia,

The ventral acetabular expansion forms a

characteristically thin but exceptionally well

developed plate, approximately equidistant

from the acetabular margin throughout its

length. The dorsal acetabular expansion of this

species is of moderate height; the dorsal pro-

tuberance is small but usually quite distinct

from the dorsal prominence as a laterally dis-

posed projection (Fig, 2e).

The size of the fossils appears to be well

within the range of modern individuals from

the same locality. In each fossil specimen the

ilial shaft is incomplete to a greater or lesser

extent,

The maximum DAE-VAE distance is 2.8

mm which, as demonstrated in Table |, can be

equated with a snout to vent length of 35 mm.

Such a body size is well within the range of

modern individuals from this area, and is the

mean for adult males in southern Victoria

(Watson, Loftus-Hills & Littlejohn 1971)-

Family LEPTODACTYLIDAE

Limnodynastes tasinaniensis Gunther

Henschke’s Quarry Cave: P, 19506, 1 left

lium; P. 18853, 18939, 2 right ilia.

TABLE |

Cumpariven al measeremerts of the largest fossil

Litoria ewlngi with modern representatives.

DAE-VAE iliul shaft snoubvent

distance length lenath

Fossil 2.8 mor 16.6 mm Cinconiplete) 735 mem

itodern =. 2.2 mm 12.3 mm 33 mn

modern = 28mm 13.7 mtn 35 mm

modern 2.7 mm 13.4 mem a5 mie

Victoria Cave: P. 16803, 14 left ilia; P. 16804,

8 right iha.

The ilium of this species is characterized by

the following combination of features: the sig-

moidal margin of the ventral acetabular fossa;

a moderate to poorly developed, and pointed,

dorsal acetabular expansion; an oval shaped

dorsal protuberance inclined anteriorly ut an

angle of 45° to the superior margin of the ilial

shaft, and a slight longitudinal indentation on

the lateral surface of the distal half of the ilial

shaft (Figs 2f and 3a).

The fossil tlia are rather small in comparison

with modem representatives of adults of this

species. To a certain extent the range of adult

size varies from locality to locality and the ex-

tremes derived from the data of Moore (191 )

and Littlejohn (1963) are: males 31,7-39,5

mm; females 32.2-43.3 mm. Of seven adult

specimens dissected by me the snout to vent

lengths are 31.0-35.8 mm, and so situated

within the lower portion of the adult size range.

The DAE-VAE span for these same seven

specimens are 4.0—4.7 mm (mean 4.4 mm).

The DAE-VAE span in 17 fossils is 2,5-3.8

mm (mean 3,0 mm), indicating that the fossil

frogs were considerably smaller than modern

adults,

Limrodynastes sp. cf, L, dumerili (Peters)

Victoria Cave: P. 16805, 15 left ilia; P. 16806,

22 right ilia,

Uncertainty about the identity of this

material exists because of problems associated

with distinguishing L. dumerili ilia from those

of ZL. pereni, In each species the dorsal pro-

Mminence and dorsal protuberance exhibit a

degree of variation not observed in any other

species of Australian frogs. In its simplest form

the dorsal protuberance can consist of a slightly

raised § projection with an obtusely angled

Superior margin (Fig, 2b), At the other ex-

treme of its development the prominence takes

the form of an anteriorly inclined hook (Fig.

2a). Between these two extremes there exists

every conceivable intermediate stage.

PLEISTOCENE FROGS FROM SOUTH AUSTRALIA 87

Fig. 2. Fossil ilia from Victoria Cave, Naracoorte. a and b Limnodynastes sp. cf. L. dumerili, P. 16805;

c Ranidella signifera, P. 16808; d Geocrinia sp. cf. G. laevis, P. 16809; e Litoria ewingi, P.

16802; f Limnodynastes tasmaniensis, P. 16804.

Fig. 3. Features of ilia. A. Limnodynastes tas-

maniensis showing lateral indentation. B.

Geocrinia sp. cf. G. laevis indicating re-

duced subacetabular zone.

Sub— acetabular.

The highest DAE-VAE distance in the fos-

sils is 7.7 mm and five others exceed 6.5 mm.

The minimal ratio of snout to vent length/

DAE-VAE distance in the extant material exa-

mined is 8.9, placing the minimal extrapolated

snout to vent length of the largest fossil indivi-

dual at about 68 mm.

In the light of numerous differences in other

osteological features, in gross morphology and

external features, the extent of similarity be-

tween the ilia of L. dumerili and L. peroni is

surprising. Until such time that other means

are found for distinguishing the ilia of these

species the identity of the present material will

remain uncertain.

Ranidella signifera Girard

Henschke’s Quarry Cave: P. 18936, 1 left

ilium; P. 18935, 1 right ilium; P. 18851, 2

right ilia; P. 18934, 2 left ilia; P. 18933, 2

right ilia; P. 18852, 1 left ilium; P. 18932, 4

left ilia; P. 18931, 2 right ilia; P. 19501, 3

right ilia; P. 19502, 2 right ilia; P. 19503, 2

left ilia.

Victoria Cave: P. 16807, 15 left ilia; P. 16808,

17 right ilia.

As can be seen in Fig. 2c the ilium of R.

signifera is characterised by a comparatively

broad acetabular fossa and a sigmoidal shaped

anterior margin of the ventral acetabular ex-

pansion. In addition the dorsal acetabular ex-

pansion and dorsal prominence are both quite

pronounced.

The maximum ilial shaft length in the fossil

material is 11.4 mm. In modern representatives

88 M. J. TYLER

this. sizé ts attamed by individuals approxi-

mately 31 mm long, approximating the maxi-

mum body length reached by this species,

Geocrinia sp. cf. G. laevis (Gunther)

Victoria Caves P. 16909, 1 right ilium,

The short and relatively broad jlial shaft and

very poorly developed dorsal prominence are

features cOmmon to Geocritia and Crinia spe-

cies, However, Geeerinia is sel apart by moder-

ale development of the preacetabulur postion of

the ventral acetabilur expansion, and poor

development of the subacetabular portion

(Figs 2d and 3b). Geacrinia Jaevis is the only

species yvailable for compurison and the fossil

agrees With that species in Most respects, Bear-

ing in mind the striking similarity of habitus

between G, faeviy and G. \icreriana, and the

genuinely close relahonship of these species. iL

is unlikely that the ilia will differ substantially

from one another.

The fossil appears to have a complete shaft

and the total length is 4.9 mm, This is consider-

ably Jess than those of the modern ila exa-

mined and indicates a subadult individual

approximately 15 mm in hody length.

Discussion

The age of the Henschke's Cave material is

at least 32,000 BLP. (N. Pledge, pers, comm.),

Preliminary dating of apatite aiid collagen from

the Victoria Cave deposits is equivocal us the

deposit may be beyond the range of C™ dating.

A more accurate age estimation awaits the com-

pletion of detailed Cl and stratigraphic studies

(R. T. Wells, pers. comm, ),

Evidence of a close relationship between ele-

ments of the Frog fauna of southeastern and

southwestern Australia has heen explained as

reflecting migration of existing species or their

ancestral stocks during the optimal Pleistocene

glacial periods (Littlejohn 1967), The fossil

Naracoorte fauna therefore assumes import-

ance in terms of acting as a sample of a popu-

lation of a southern Australian species, con-

ceivably occurring at a time When the lust

glacial migration may not have reached its

maximal geographic limits,

One unexpected deficiency in the Netacoone

fossil fauna is Litoria raniformiy (Keferstein)

or any other species af the 7, aurea (Lesson)

species complex as defined by Tyler & Dayies!,

The deficiency is quite genuine, for the iia and

other bones of L, raniiforntis are large Cilial

shaft length up to 33 mm) and they exhibit a

number of features distinguishing the species

from all other sympatric species in South Aus-

tralia. Thus there is no possibility that its pres-

ence could have evaded detection during the

seruing procedures,

The absence of &, raniformis is an anomaly

heenuse it currently extends from New South

Wales 10 the southeast of South Australia in-

cluding the Naracoorte area. The £, avree spe-

cies complex is now well established with four

species in southeastern and two species in

southwestem Australial, Although the markecl

level of divergence in the now allopatric spe-

cies ean be attributed to migration from the

southeast to the southWest in a penultimate or

earlier glacial migration, there has been no sug-

gestion that representation of the complex in

southeastern Australia has been ofher than con»

Hours.

Certainly £. rani/ormis has existed in at least

part of southeaster Australia for 12,000 years

Decause the species occurs im Tasmania, und

this js the period of the mast recent isolation of

lasmania from the maitland (Littlejohn &

Martin 1974), To what extent the maxinium

duration of colonisation of the southeast can be

gauged ts uncertain, but there is evidence that

the existing boundaries of its distribution are

far from being static, For example, Tyler &

Roberts (1973) report the recent introduction

afd subsequent establishment of the species at

several locations in the Mt Lofty Ranges, and

subsequently it has been found at other “new”

adjacent localities near Adelaide. Vhe ease with

which it ts now colonising this new territory

indicates that it is unlikely thal the species has

existed (here previously and has since been dis-

placed, unless extinction can be ennsidered a

possibility

It would therefore appear that the species

may not have existed in the extreme southeast

32,000 years aga, bul enlered the arca between

that dale and 12,000 years ago. Since 12,000

B.P. dispersal westward within the southeast

has couceivably been inhibited by arid condi-

trans, ‘

In the Jight of the above absence of L, rani-

Jermis in the Naracoorte Tossil fauna, the pre-

sence there of Limnedyviasies tasmnuntenzis is

particularly perplexing. This species occurs fn

Vin & manuscript on species groups im Liverig. The £. aurea complex currently comprises £. aurea, L,

ranifermis, LE, moerei (Copland), L. albegutatra (Gunther), L, cyclorhvaches (Boulencer), and L.

flavipwnerata Courtice & Grigg.

PLEISTOCENE FROGS FROM SOUTH AUSTRALIA 89

southeastern Australia and its western geo-

graphic limit is Eyre Peninsula of South Aus-

tralia. Ti does not occur in southwestern Aus-

tralia nor is there a closely related species there.

Thus it would appear that DL, fasmuniensis

existed in the southeast at a time when the

hypothetical mesic communication to the south-

west existed, but did not traverse the communi-

cation and thus failed to penetgate the south-

west. The presence of Litoria ewingé in the

deposits is equally surprising in vicw of ils

absence from southwestern Australia,

Two of the other species identified m the

Naracoorte fauna exist in the vicinity today:

Literia ewinvi and Limnodynastes cf. dumerili

(Tyler 1966). The third is Geoerinia laevis

which is limited in South Austrulia to the ex-

ireme lower southeast, and does not now ex-

tend as far north as Naracoorte. However,

Beck (1975) postulates that there have heen

substantial withdrawals of the geographic range

of this species following Lrends towards aridity

and also as a result of drainage and clearing of

vegetation,

In the case of other small vertebrates al Vic-

toria Cave, Smith (197), 1972) has suggested

that they were brought mlo the cave by pre-

dators such as owls, Frogs can form a substan-

tial portion of the diet of some owls so that

this interpretation applies equally well to the

frog material, However, three species live in the

caves al the present time! Litoria aurea, Limno-

dynastes dumenill and L, taxmaniensis.

Acknowledgments

I aim most grateful to Dr R. T. Wells (Flin-

ders University of South Australia) for permit-

ting me to examine sievings from Victoria Cuve

and so jnitiating this study. 1 am also indebted

to Mr WN. Pledge (South Australian Museum}

for providing material fram Henschke’s Cave.

My thanks are due to Dr Wells for reading and

criticising this manuscript and to Mr P. Kemp-

ster (Department of Zoology, University of

Adelaide) for ihe photographs used in Figure

References

Beck, R. G, (1975) Faetors affecting the distri-

bution of the Jeptodactylid frog Geocrinia

laevis in the south-east of South Australia,

Trans. R, Soc. §. Aust. 99(3), 143-147

Lirrcesonn, M, J. (1963) Frogs of the Mel-

bourne area. Vict. Nat, 79(10), 296-304,

LiyiLEJOHN, M, J, (1967) Patterns af zoogeo-

graphy and speciation in south-eastern Austra-

lian Amphibia. Jk A. H, Weatherley (Ed,),

“Australian Inland Waters and Their Pauna;

Eleven Studies.” (Avstcalian National Dni-

versity Press: Canberra, )

Lirmiesoun, M, J, & Martin, A. A (1974) The

amphibia of Tasmania, fa W. D, Williams

(Ed.), “Biogeography and Ecology in ‘Tus-

mania, (W. Junk: The Hague.)

Moore, J, A. (1961) The frogs of castern New

South Wales. Bull. Aen, Mus. Nat. fist. 124,

149-386,

SmttoH, M. J. (1971) Small fossil vertebrates Irom

Victoria Cuve, Naracourle. Sowih Avstralia, §.

Portoroinae (Macropodidae), Petauridac and

Burramyidae (Marsupialia) rams, R. Soc. 8.

Aust, 95(4), 185-198,

Smirn. M. J. (1972) Small fossil vertebrates from

Victoria Cave, Naracoorte, South Australia.

0), Perameldae, Thyacinidsae and Dasyuridae

(Marsupialia). [bid 96(3), 125-137,

Tycer, M_ J. (1966) The frogs of South Australia.

(South Aust. Mus.; Adelaide.)

Tycex, M. J, (1974) First frog fossils from Aus-

tralia, Narure 248(5450), 711-712,

fyver, M, J, (1976) Comparative osteology of the

pelvic gitdle of Australian frogs and deserlp-

lion of a new fossil genus. Trans, R. Soe, 8.

Aust. 1W1(1), 3-14,

TrLer, M. J, & Rogers, J, BD. (1973) Noteworthy

range extensions for some South Australian

frogs, 8S. Ausr. Nat. 4812)-

Van Ters, G. F, (1974) A revision of the fossil

Megupodiidae (Aves), including a description

of a new species of Progura de Vis. Trans, &,

Sec, §, Aust, 9804), 213-224,

VAN Tours, G. FP. & Sarma, M, J, (1974) Small

fossil vertebrates from Victoria Cave, Nura-

coorte, South Australia, T1, Birds (Aves),

Ibid. 98(4). 225-227.

Warson, G. F. Lorrus-Hiius, J. J. & LIvTecicHn,

M. J. (3971) The JLitoria ewinei complex

(Agura’ Hylidue) in southeastern Australia.

fA new species from Vietoria, Anse. J, Zoel,

19, 401-406.

We its, R. T. (1975) Reconstructing the past, ex-

cavauions in fossil caves. Aue. Nat. Mist, 13

(6), 208-21 1.

Woookurr, D. S. & Tycek, M, J. (1968) Additions

to the Frog Fauna of South Australin. Rec. &-

Aust. Muy. 18(4), 708-709,

DISTRIBUTION OF INTRODUCED LAND-SNAILS ON YORKE

PENINSULA, SOUTH AUSTRALIA

BY A. J. BUTLER AND C. MURPHY

Summary

In three consecutive years surveys for five species of introduced terrestial pulmonates and one

native species were conducted on a grid of sampling-points approximately 8 km apart covering the

whole of York Peninsula. Sampling in the different years was done by different observers and,

despite some technical difficulties, they agree on broad patterns of distribution. Those patterns

differ greatly from the results of a survey conducted in 1965. The results, especially possible

interactions between species and the spread of Theba pisana, are discussed.

DISTRIBUTION OF INTRODUCED LAND-SNAILS ON YORKE PENINSULA,

SOUTH AUSTRALIA

by A. J. BuTLER* and C, Murpuyt

Sommary

Burner, A. Jo and Murpuy, C. (1977) Distribution of introduced land-snails on Yorke Penin-

sula, South Australia, Trams. R, Soe, §. Aust. 101(4), 91-98, 31 May, 1977.

In three consecutive years surveys for five species of introduced terrestrial pulmonates and

one native species were conducted on a grid of sampling-points approximately & km apart

covering the whole of Yorke Peninsula. Sampling in the different years was done by different

observera and, despite some technical difficulties, they agree on broad patterns of distribution,

Those patterns differ greatly from the results of a survey conducted in 1965, The results,

especially possible interactions between species and the spread of Thea pisana, are discussed.

Introduction

Ten species ol terrestrial snails have been

introduced into South Australia since colonisa-

tion. Pomeroy & Laws (1967) examined the

earliest records and concluded that, although

exact times of arrival and origins were difficult

to ascertain, it was probable that at least four

species came from the Mediterranean region,

This paper concerns the distribution on Yorke

Peninsula of five of these species, Helicella vir-

sata, H, negleetu, Cachlicella acuta, C. ven-

treya and Thela pisetia as well as that of the

native snail, Awstresnecinea australis,

Pomeroy & Laws (1967) summarised avail-

able information about the distributions of

introduced snails in South Australia at that

time. Their information on distributions on

Yorke Peninsula came from two kinds of

source, For C, acnia, C. ventrosa and T. pisana

they present spot. records from South Austra-

lian Museum collections and their own collec-

tions. For A. virgata and H. neglecta they pre-

sent (he results of their own survey in which

roadsides were examined on a grid of points

about 8 km apari, covering the entire Penin-

sula. AL these points ratings were assigned to

the abundance of snails, The survey was con-

ducted during summer.

In their findings are two major points to be

compared with the results of the present sur-

vey: Y pisana was recorded from only two

sites, Corny Point and Edithburgh; the distribu-

tions of H. virgata and H. neglecta were almost

mutually exclusive, H. neglecta occupied the

“foot” and “ankle” of the Peninsula, being

especially abundant along the north coast of

the “foot” from Corny Point to Pt Turton,

whilst Al. virgata occupied the upper “leg”

being most abundant around Wallaroo and

Moonta. A small gap between the two was

noted in the region of Minlaton.

Pomeroy & Laws (1967) record evidence

that H. neglecta was contracting its range

(having formerly occurred as far north as

Moonta and perhaps over much of the Penin-

sula although the records are unreliable), whilst

A, virgata was expanding its range but, , , “one

ean only speculate as to whether the two events

are related”. They also suggest that T. pisana

had reached its 1967 distribution rather rapidly,

Purther, they present observations over two

years on the numbers of T. pisana and A, vir-

rata in two quadrats at Outer Harbour and Pt

Adelaide from which they suggest 7. pisena

may have been increasing whilst A. virgala was

decreasing in numbers.

Lim & Jenkins (1972) carried our a survey

for T. pisana and A, virgara \n the southeast of

South Australia because the former presents an

economic problem there. They also noted the

known distribution of T. pivana throughout the

State. On Yorke Peninsula they found it at two

more siles than did Pomeroy & Laws (1967),

Lim & Jenkins (1972) also noted that in the

* Department of Zoology, University of Adelaide, North Tce, Adelaide, S. Aust. 5000.

* Present address: Department of Human Morphology, Flinders Medical Centre, Flinders University

of Sourch Australia,

92 Ao J. BUTLER & C. MURPHY

southeast 7". pisana and H. virgata tended not

to vccur at high densities together,

Tt was therefore of both theoretical and prac-

tical interest to re-cxamine the distributions of

introduced snails on Yorke Peninsula. Wheoreti-

cally, because Pomeroy & Laws (1967) raise

interesting ecological questions. They suggest

T. pisana may replace H- virgata by conjpeti-

tion for food: and if H. virgata is indeed ex-

panding its range whilst A. neglecta is contract-

ing at about the same rate, the mechanism is

by no Means obvious,

The study was also of practical interest be-

cause T, pisana is a costly pest in other parts

of the world (Pomeroy & Laws 1967; Rimes

1968; Nevo & Bar 1976) and is already a

nuisance in the southeast of South Australia

(Lim & Jenkins 1972). Its expansion i this

State is therefore a cause for concern.

This paper presents the results of surveys

carried out from 1973 to 1975 on Yorke Penin-

sula. The grid of sampling points was similar to

that used by Pomeroy! and Pomeroy & Laws

(1967) but the procedure for assigning ratings

was 2 little different. The surveys were not

designed to explain the changes. in distribution

but we make some comments on them.

Methods

Sampling points were chosen in the manner

of Pomeroy & Laws (1967). Sites were on

roadsides since these provide a favourable habi-

tat for snails and are readily accessible for

observation,

Roads approximately parallel to each other

and & km apart were plotted on large-scale

maps xynd sampling points (“stops”) were

chosen at 8 km intervals along them (Fig. 1}.

The detailed laying-out was done independently

in 1973 and 1974, In 1975 the 1974 layeur was

available in detail and it was followed again

except for the addition of sonie extra stops,

Nevertheless, due to random error in odometer

readings, 1975 stops do not correspond pre-

cisely with 1974 stops. Thus all four surveys

have the same basic plan but do not |p general

incorporale exactly the same sampling siles, A

standard procedure was used for assiyniiy a

rating to the abundance of snails at cach stop,

on a scale from 0 (no snails) to 4 (over 150

snails/m*). A separate rating was assigned for

each of the six species, The data on abundance

ate available fram the authers, but only “pres-

ence-or-absence” records are presented here.

The fieldwork was done by a different pair

of observers in each of the three years (see

Acknowledgments). Each pair of workers exa-

mined muscum specimens ahd literature in

order to learn to identify snails and practised

the technique of estimating abundance so that

the observers were accurately “calibrated”

hefore each survey bevan, These preparations

were inade independently by the three pairs of

observers, In general, specimens were not col-

lected, but a sel of voucher specimens was

taken from selected stops in August 1976 by

A. Butler; these have been lodged in the South

Australian Museum.

Although in 1973 a watch was kept for sitails

between sampling poilits where ratings of O had

been recorded, as donc by Pomeroy & Laws

(1967), this wus subsequently discontinued.

Several species were very small, well-camou-

Haged and usually hidden in vegetation or

under rocks. Also, careful inypection was

needed ta distinguish between the two Helicelle

species. Moreover, numbers at a particular site

will vary greally with the seasons (Pomeroy &

Laws 1967; Pomeroy 1969) because of both

population Machinlions (Pomeroy 1969) ani

movement (Hodson®). Snails also bury them-

selves (Pomeroy), Therefore our results are

presented nut as maps af continuous distribu

Gon bul as y large set of point observations-

The survey of Pomeroy & Laws (1967) was

conducted in summer When snails are mostly

dormant (Pomeroy 1968; Hodson?) and Heli-

1 Pomeroy, D. F, (1966) The ecology of Heficelle virgata and related species of snails im South Autss-

tralia. Ph.D. thesis, University of Adelaide.

2? Hodson, A, C. (1969) Adaptations that permit the terrestrial snail Helicella viredta (Da Costa) to

survive in dry places. Ph.D. thesis, University of Adelaide.

FIGS 1-10

Fig. |, Sampling-points on Yorke Peninsula, Figs 2-10; Records of smalls ln each of the three yeara

fsummer and winter records pooled). Excepting a few sites (with subseript A in Fie. L) added in

1975, all sites shown there were visited each yeur. Thus, the absence of o spot on the mop in Figs

2-10 indicutes 4 negative record for the corresponding site. 2, Helfeethi virgatmw, 1973, 3, HL vir-

gata, 1974; 4, A. virgata, 1975; 5. H. neglecta, 1973, 6. H. neglecta, 1974: 7, MH. meglecia. 1975; #8.

Thebs pisuna, 1973; 9. T, pisane, 1974; 1h T. plana, 1975,

DISTRIBUTION OF LAND-SNAILS ON

105

jo6” 108 109110 28

YORKE PENINSULA

94 A.J, BUTLER & C, MURPHY

cella might be more conspicuous (although this

depends greatly upon the kinds of sites avail-

able for aestivation), Gur 1973 survey was con-

ducted in winter. The 1974 and 1975 surveys

were done i both summer (Februnry) and

winter (uly). Within one year jhe sampling

sites were the same in the two seasons.

At cach site Vegetlation-type was briefly

noted.

Results

The results are shown m Figures 2-19. Bach

spot represents a non-zero rating. Spots on the

1973 maps represent observations in summer;

a spot on a 1974 or 1975 map represents a

recording in summer, winter or both. This has

heen done to facilitate discussion of broad pat-

terns. Separate summer and winter results, and

the actual ratings, are available from the

authors,

Discussion

Several technical points must be made. It is

sometimes diffieult to distinguish A. virgata

from Fl. neviecta on shell morphology alone

and it is valuable that the three pairs of observ-

ers learned, from literature and South Austra-

lian Museum specimens, to distinguish the spe-

vies. Thus although some: errors. of identifica-

Hon may have been made in the field, there is

unlikely to be a systematic bias dine to “cultural

transmission” amongst the reams, The 1973

workers collected some doubtful specimens for

later confirmation by dissection, Voricher speci-

mens collected hy A J.B. in August 1976 from

points 104, 72, 41, 7, 6, 23, 22, 26, 20, 17 and

28 have been deposited m the South Australian

Museum,

Because there are likely to be Wide fluctua-

tions with season in both numbers and con-

sprcuousness of snails. for numerical compari-

sons Pomeroy & Laws’ (1967) summer results

should be compared primarily with our sum-

mee results. Similarly our 1973 survey should

be compared with the Winter results of 1974,

1975, However in this paper we have pooled

nur summer and winter results, ymd we discaiss

only broud patterns.

Pomeroy! surveyed systematically, jh the

way We did, only for the two Helicelly species,

The information presenled hy Pomeroy & Laws

(1967) on other species is based on museum

records and spot records by the authors and

must not be discussed as complete maps of dis-

tribution.

Finally we note that all of the sampling sites

were on roadsides, Tt must be remembered that

these sites aye especially vulnerable to human

influence such as clearing, dumping, spraying

for weed control and ploughing for firebreaks,

Thus we must assune that to the natural fluc-

tuations which occur in snail populations

(Pomeroy & Laws 1967; Pomeroy!) is added

an unknown variability from year to year, This

could inclade the founding of new populstiuns

(Pomeroy & Laws 1967; Pomeroy 1967) as

well as the changes of habitats from favour:

ahle to unfavourable or vice-versa-

Austrosuccinea australis

This native snail seems to be low in numbers

throughout the Peninsula (Figs 17-19). tt was

scarce in 1973 but nevertheless was recorded at

14 sites. In 1974 it was not observed at all and

in 1975 it was found at one site in winter, Per.

haps its range is ¢entracting, but given the

technical polats above, We cannot conclude sa

with any confidence.

Cochlicella spp.

Pomeroy & Laws (1967) tecorded C, veri-

frosa at Corny Point. As voted above this ts not

strong evidence that this snail was absent at

all other points. Our results (Figs 15-17) show

it in more localities, with a suggestion that it

may be increasing (compare Figs 16, 17)-

Almost all our records of C. ventrosa were it

winter, perhaps part of the reason why they

recorded ii at only one place,

C. acute was recorded at nine sites on the

“foot'' and “lower leg” by Pomeroy & Laws

(1967). Our results do not show it outside this

general area, but there is some suggestion that

it is “filling-in” its area of distribution, (Com

pare Figs 11-13), As with C, ventrosa, how-

ever, we must not make too mach of small

differences between years but the independence

uf our three surveys gives us confidence that

€, aeuita is widespread over southern Yorke

Peninsula whereas C, venfresa occurs in iso-

lated packets scattered over the Peninsula.

Meiicetla spp.

The changes in the distributions of the two

Helicella species can be followed from the sum-

FIGS 11-19

Dewals as for Pies 2-10, IL. Cochiicella acuta, 1973; 12, Co vewte, 1974. 13, ©, acuta, 1975; 14. CC.

veetrosa, \973, IS. CL vertrease, 1974; 16. C, veniresa. 1975; 17,

LB. AL anerealis, 1974; 19, A, auerrefis, 1975.

Austrosmerinva australis, 1973:

DISTRIBUTION OF LAND-SNAILS ON YORKE PENINSULA

%6 A, J, BUTLER & © MURPHY

ier of 1954-55, Prof, H, G, Andrewartha

(pers. comm.) made a tour of the Penmsula

during that summer on which he observed Heli.

cella Virgata aecstivating in great numbers on

fenceposts in the vicinity of Wallaroo, Despile

a search of about two days in travelling south-

wards down the Peninsula he found no more

H. virgata beyond a few miles south of Wal-

laroo; in fact, he found no snails at all until

south of Minlaton, Me did find dense popula-

tions of A, reelecta in the southern part of the

Peninsula where it had been known to have

been present for the past 64 years.

In 196) Pomeroy & Laws recorded MW. vir-

gata extending southwards almost as far as

Minlaton (see Introduction),

By 1973 (Figs 2, 5) the distribution of HW.

neglecta was much contracted and that of H,

virgata greatly extended from those reported by

Pomeroy & Laws (1967), Given the independ-

ence of the three surveys this difference ts very

convincing. It is far ton great to be accounted

for by year-to-year fluctuations and different

observers in the three years agree over the

broad pattern; H. virgata js widespread over the

Peninsula including most of the “foot” where

Pomeroy & Laws (1967) recorded it at only

two locations; A velecta has contracted to

small areas in the “foot”, although the pocket

near Moonta, shown by them may still exist

(Fig. 5), On this point, however, note the diffi-

culty in identification discussed above; we must

concern Ourselves in this paper with broad pat-

terns which were observed by three independ-

ent teams of observers, and the northeri

records of H. neglecta m Fig. 5S were not

repeated by the (974 and L975 teams.

We must again he cautious in interpreting

year-to-year differences, H, pepglecta may be

continuing to contract [compare Fig, 4 with

Fig. 6); in 1974 (Fig, 7) more sites were exa-

mined and this accounts for some of Che addi-

tional records, H, virgata may have filled in

some areas within its overall range (Pigs 2-4),

but this has occurred in areas where Pomeroy

& Laws (1967) found it to be Very abwndant

ind so may be regarded as part of a fluctuation

in numbers,

We note in passing that on a brief visit to 4

few stops in August 1976 essentially the same

pattern was found as in 1975 except that no

live specimens of H, neglecta Were taken, This

species was eXpected, at least in low ounahers,

af sites & 28 and 17 bur only dead shells were

found there.

Theba pleana

This snail is scarce on much of the Peninsula

but appears to be spreading, Cotton (1949)

dees nol record il on the Peninsula at all,

According to Pomeroy & Laws (1967) it was

then known from two sites only, Corny Point

and Edithburgh. Lim & Jenkins (1972) gave

four sites on the Peninsula, adding one near

Stenhouse Bay and another pear Pt Vincent,

Such point records do nat convincingly indicate

a spread, but on our surveys 7, pivana was

found at six sites in 1973, eight in 1974 and

13 i) 1975S (Figs 8-10) and its numbers at

some of these new sites were high.

Further, the number of “point records” con-

tinues to Increase, By the end of 1975, the

Depurtment of Agriculture had records of T.

pisana from Ardrossan, Pine Point, Edithburgh,

Warooka, the coastal dunes 4 km due north

of Warooka, Point Turton, Hardwicke Bay,

Brentwood, Port Minlacowie and Port Vic-

tona (P, R. Birks, pers, comm, ),

The expansion of T pisana has occurred

mostly along coastal areas of the lower Penin-

sula, Since the coastal dumes have wnlil recently

heen nearly inaccessible by road much of this

expansion may depend largely on the snails’

own locomotion, However, Pomeroy & Laws

(1967) thought that man was an important dis-

persal agent for A, virgata, and this is almost

certain to be true of Theba now. P. R. Birks

states that Theba is now present at “almost

every crossroad between Minlaton and Brent-

wood", Snails are likely to hecome established

al crossroads because people park cars there,

having previously parked at centres of dense

snail populations. (Many of our sampling sites

Were not at crossroads.)

Interactions berween species

Pomeroy & Laws (1967) noted almost mutu-

ally exclusive distributions for H. virgata and

4, weelecia and suggested that H. neglecta was

contracting and A, virgata expanding its range.

Our results seen tO confirm this, hut there is

ho longer a “confrontation” between the two

distrihutions. A. neglecte now occurs in pockels

within the range of H. virgata and there is one

site (No, 6 on Fig. 1) where both have been

recorded, However, there are places where A.

neclecty has disappeared but A. vireara has

never heen recorded. Thus the hypothesis that

A virgata ts displacing H. xeglecta can, at best,

be only part of the story. We can guess at

various environmental changes which may be

responsible, but have no tests, Moreover, in our

DISTRIBUTION OF LAND-SNAILS ON YORKE PENINSULA 97

thiee surveys and that of Pomeroy & Laws

(L967) no relationship could be found between

Vepelution-type and the snails present.

Aa jfteraction may also be occurring be-

tween T. pixena wod Al. virgata. Numbers of 7,

pisana at sites on jhe southern Peninsula have

mereased from 1973 to 1975 whilst H, virgata

may have contracted there (compare Pigs 9

and 10 with Figs 3 and 4), Again not too

much can be made of year-to-year diflerences

and there are sites where M. virgata has de-

¢reased but 7’ pisana has never heen recorded.

Nevertheless the hypothesis of some direct

interaction bears closer consideration here

(sites 20, 24, 25, 32, 39). Moreover Pomeroy

& Laws (1967) presented dala from Outer

Harbour showing a slight trend for TL pisane to

increase and H, virgata to decrease in numbers

over (Wo years in the same area and Lim &

Jenkins (1972) found that in the southeast of

South Australia the (wo species tend not to

occur together ta high densities,

Pomeroy & Laws (1967) tentatively interpret

their observations at Outer Harbour in terms of

competition far food, However there is some

evidence (Butler 1976) that at least in one

arca, H, virgara may not be short of food jn an

absolute sense (Andrewartha & Browning

1961). On the other hand it is known for cer-

tain aquatic pulmonates that large numbers of

snails can inhibit one another's growth, survival

and reproduction even in the presence of abur-

dant food (Thomas & Benjamin 1974; Thomas

et al, 1974). Such an effect could occur in ter-

restrial snails, possibly through soiling ol the

food and substrate with excreta, or slime-trails,

and may allow T_ piyaria to replace A. virgata.

Critical experiments will however be needed jo

discriminate between possibilities.

Polymorphism in shell banding

During the 1975 survey it was noted that

three inland populations of 7, pisxana were all

monomorphic for unbanded shells whereas the

coastal dune populations were polymorphic far

shell banding. Vegetation inland was lower,

mostly grass; coastal veyetation was more

dense, and mostly low bushes, This difference

in Morph frequencies was not noted in A. vir-

gata or Hf. reglecta, all populations of which

were polymorphic. Roberts! found a similar

Situation in Hf. wirgate af Semaphore Park ex-

cept that inland populations contained the

banded morph in low frequency (less than

5%).

There aré many suggestions in the literature

that genetic changes are important im the dyna-

mics of populations of aninoals including snails

(for review see Williamson 1972), Thus,

especially in view of the suggestion of Baver

stock? that food may be one of the factors

influencing morph-frequencies in H, virgata, it

muy be instructive to consider morph-frequen-

cies in examining both the distribution of snails

on the Peninsula and the hypothesis of com-

petitive displacement.

Ecatomic impartance

The Melicelle species constitute only a mifor

economic problem, although pot wegligible

(Pomeroy & Laws 1967; Birks pers. comm.),

Where their numbers aestivating on stalks are

high enough to foul grain, they can be con-

trolled with a carbamate bait or by using a

xpecial outngger on the harvester (Rimes

L968; Lim & Jenkins 1972), TL pisane, on the

other hand, not only may cause the same prob-

lem but can alsa damage lucerne and has been

reported us doing so in the southeast of this

State.

Both Cochlicella venrrosa and Theba pixana

have been causing significant fouling of grain

on Yorke Peninsula in the last few years, and

in 1976 the Department of Agriculture and

Fishenes received its first report (from near

Port Lincoln on Eyre Peninsula) of Theda

actually eating barley plants and causing con

siderable damage especially to young seedlings

(P, Birks, pers, comm.). Our records suggest

that this snail is spreading, and so it is poter-

tially a serious pest on the Peninsula,

Acknowledgments

The figldwork was carried out by Stewart

Roper, Peter O'Donoghue, lan Fraser, Tony

Smith, Peter Geant and Chris Murphy as part

of their Honours courses in zoology in t973—

75. without theie efforts it would have been

impossible to reeord sq many observations, We

are grateful also to Prof, H. G. Andrewartha

aod Mr P. R. Birks for critically reading the

manuscript and especially for allowing us to

quote their unpublished records,

4 Roberts, J, DB, (1970) Shell pattern polymorphism in the snail, Helicella virgatd, B.Sc, (Hons) thesis,

University of Adelaide.

4 Baverstuck, P. (1968) Polymorphism in bandiag in the snail) Hellcella virgata, B.Sc, (Hons) IIeesis,

University of Adelaide,

98 A. J. BUTLER & C. MURPHY

References

ANDREWARTHA, H. G. & BROWNING, T. O. (1961)

An analysis of the idea of resources in animal

ecology. J. theor. Bial, 1, 83-97.

Butier, A. J. (1976) A shortage of food for the

terrestrial snail Helicella virgata in South Aus-

tralia. Oecologia (Berl.) 25, 349-371.

Cotron, B. C. (1949) These snails may invade

your garden, J. Agric., S. Aust. 52, 561-565.

Lm, Y. P. & Jenkins, R. B. (1972) The Italian

white snail . . . Theba pisana (Miiller) in

South Australia with particular reference to its

distribution in the south-east of South Aus-

tralia. Exp. Rec. Dept Agric. S. Aust. (6),

23-25.

Nevo, E. & Bar, Z. (1976) Natural selection of

genetic polymorphisms along climatic gra-

dients. Jn S. Karlin & E. Nevo (Eds), “Popu-

lation genetics and ecology”. (Academic

Press: New York.)

Pomeroy, D. E. (1967) The influence of environ-

ment on two species of land snails in South

Australia. Trans. R. Soc. S. Aust. 91, 181-186.

Pomeroy, D. E. (1968) Dormancy in the land

snail, Helicella virgata (Pulmonata: Helici-

dae). Aust. J. Zool. 16, 857-869.

Pomeroy, D. E. (1969) Some aspects of the eco-

logy of the land snail, Helicella virgata in

South Australia. hid. 17, 495-514.

Pomeroy, D. E. & Laws, H. M. (1967) The dis-

tribution of introduced snails in South Aus-

tralia, Rec. S. Aust. Mus. 15, 483-494,

Rimes, G. D. (1968) Snail investigations—a pro-

gress report. J. Dept. Agric. W. Aust. 9 (4th

series), 584-587,

Tuomas, J, D, & BENJAMIN, M. (1974) The effects

of population density on growth and repro-

duction of Biomphalaria glabrata (Say) (Gas-

teropoda: Pulmonata). J. anim. Ecol. 43, 31-

50.

Tuomas, J. D., GoLpswortny, G. J. & BENJAMIN,

M. (1974) Chemical conditioning of the en-

vironment by the freshwater pulmonate snails

(Biomphalaria glabrata) and its effect on

growth and natality rates. J. Zool., Lond. 172,

443-467.

Wivuramson, M. (1972) “The analysis of bio-

logical populations.” (Arnold: London.)

NESTING BIOLOGY OF THREE ALLODAPINE BEES IN THE SUBGENUS

EXONEURELLA MICHENER (HYMENOPTERA: ANTHOPHORIDAE)

BY T. F. HOUSTON

Summary

Results of a detailed study of nest populations of Exoneura (Exoneurella) tridentata are presented

with less complete observations of E. (E.) eremophila and E. (E.) setosa. These species rear their

larvae progressively in open burrows in dead plant stems, tridentata in ready-hollowed woody

twigs, eremophila and setosa in pithy herbaceous stems. E. tridentata exhibits small semisocial

colonies with queen and worker castes. The castes are dramatically morphologically dissimilar, the

large-bodied queens appearing flightless and restricted to nests. By contrast, colonies of eremophila

and setosa are basically subsocial. The subsocial condition in these species may be derived from a

semisocial condition and possibly associated with life in rapidly perishable stems.

NESTING BIOLOGY OF THREE ALLODAPINE BEES IN THE SUBGENUS

EXONEURELLA MICHENER (HYMENOPTERA: ANTHOPHORIDAE)

by T. F. Housron*

Summary

Houston, T. F. (1977) Nesting biology of three allodapine bees im the subgenus Exowieurella

Michener (Hymenoptera: Anthophoridae), Trans, R, Sec. S, Aust. 1Ob(4), 99-113, 31

May, 1977.

Results of a detailed study of nest populations of Exenenra (Exoneurella) tridentata are

presented with legs complete observations of EB. (B.) eremephila abd &, (B.) setosa, These

species rear their larvae progressively in open burrows in dead plant stems, tridentata in ready-

hollowed woody twigs, eremophila and serosa in pithy herbaceous stems. BE. tridentata exhibits

small semisocial colonies with queen and worker castes, The castes are dramatically morpho-

logically dissimilar, the large-bodied queens appearing flightless and restricted lo nests. By

contrast, colonies of eremophila and setosa are basically subsocial. The subsocial condition in

these species may be derived from a semisocial condition and possibly associated with lite

in rapidly perishable stems,

An unidentified encyrtid wasp, reared from pupae of mridenraia, was found in association

with the bees,

Introduction

The bulk of this paper presents the results

of a study of the nesting biology of Exoneura

tridentata. The remainder presents less com-

plete data on the biologies of E, eremophild

und £. sefoxa, ‘These three species were

described only recently (Houston 1976) and

nothing hitherto has been recorded of their

bionomics, With &. /awsoni Rayment, they

comprise the subgenus Exoneurella (some-

times accorded gencric status) and belong to

the group of bees termed allodapines (after

Allodape). A general account of the biology of

allodapine bees ix given by Michener (1974),

Most species make their homes in burrows in

pithy stems or twigs or in dead wood, They

construct no cells but raise their immatures

together in the commen nest chamber. Usually

the lurvae ure fed progressively to maturity by

the adult females. Most allodapines display

primitive social behaviour in at least part of

their life cycle, two or more females occupying

one nest and exhibiting division of labour, One

female functions as an egg layer (or queen)

and seldom forages, while one or more other

females with undeveloped ovaries function as

workers, foraging and tending the brood.

Michener (1964) considered A. lawseni to be

essentially ‘solitary’ (really subsocial) although

several other species of Exoneura are semi-

social (Michener 1965). He postulated that the

‘solitary’ habits of &. lawseni were probably

derived from a semisucial condition and not

primitive. The new information on close rela-

tives of E, lawsoni presented below is con-

sidered in relation to this problem.

The study of E. rridentera was prompted by

discovery of the first nests which contained

relalively huge females amongst )ormal-sized

individuals (cf. Figs 1A, 1B). Female size

variation and allomeiry in this species is

described by Houston (1976). Such variation

iy absent from other Exereure!/a,

Exoneura tridentata Houston

Methods

All nest material was collected from north-

eastern Eyre Peninsula, S, Aust, and the

greater part of it from the Lake Gilles National

Park, 110 km SW of Port August. Nest col-

lection was carried out during carly moming,

late evening or during cool rainy periods when

all occupants should have been present-

* South Australian Muselim, North Tce, Adeluide, S. Aust. 5000.

100 T. F. HOUSTON

Fig. 1. Females of Exoneura tridentata (both to same scale). A. Small minor (pinned). B. Large major

(live).

Nest contents were observed alive and then

preserved as soon as practicable after collec-

tion in 70% ethyl alcohol.

Adult females were measured, examined for

wing wear and most were dissected to deter-

mine ovary condition and whether or not mat-

ing had occurred. Wing wear was rated 0 (mar-

gins entire), 1 (one to three nicks), 2 (four to

ten nicks), 3 (over ten nicks), 4 (badly

tattered). Ovary size was rated 1 (ovaries tiny,

slender and borne on long oviducal stalks), 2

(slightly enlarged), 3 (moderately enlarged),

4 (large, at least one ovum near egg size). The

length of the largest ovum of each female was

also measured.

General observations

Nests

Adults of E. tridentata utilise ready made

burrows in dead woody twigs or branches of

standing trees for both shelter and brood rear-

ing. The term ‘nest’ is used below to denote

burrows used for brood rearing. Shelters are

virtually identical except that they are not

always cleared of debris, especially when

occupied solely by males.

Except for one nest in a branch of native

pine (Callitris sp.), all occupied tunnels were

in dead twigs and branches of Bullock Bush

(Heterodendrum oleaefolium Desf.), a com-

mon small tree of semi-arid southern Australia.

NESTING BIOLOGY OF KNONEURELLA

The lower branches of the trees are commonly

dead and prone lo attack by beetle larvae which

bore through the centres of the Lwigs. Searches

of other shrubs and trees revegled very Tew

hollows which the bees might utilise,

Occupied tunnels were O0.6-2.0 m above

ground, 2-4 mm in diameter and 25-628 mm

in lenyth (most were shorter than 100 min),

They were inclined at various angles from hori-

zontal to about 45° from vertical, Their

entrances were either terminal i the ends of

hroken twigs or Taleral but none opened

directly upwards so that rain might enter,

Entrances were unmodified, showing no traces

of the collars of compacted plant material that

characterise nests of most other allodapines.

Some himnels with jateral entrances were

double-ended und the bees occupied just one

or both ends, In one case (nests 26, 27. Table

1) it appeared that two independent nests had

been estublished in opposite ends of a common

burrow,

All nests collected m December and October

contained aw umorphous patch of dry pollen

covering the Wall of one side between the broad

and the enlrance, The patches varied from the

merest smear to thick masses up to 30 nim

long. Some were composed af one kind of

pollen, others of two or more kinds. The quan-

tity of pollen in some nests leaves little doult

that the bees hud been accumulating it as a

tood store and had pot simply deposited it

uecidentally while preparing. provisions for the

larvae.

Nests collected at other times lacked fresh

pollen stores. although a few contamed old

mould-eneristed ones. The ubsence of pollen

stores from these nests cannot be attributed to

lack of flowers for they occurred in abundance

at the time of collection. More probably the

limited fine weather suitable for foraging was

insufficient fo allow aecumulation of surplus

pollen,

framathres

A deseription of the immature stages is given

by Houston (1976), All occurred loosely with-

in the tunnels, Begs usually lay crisscrossed or

jumbled in the closed onds regardless of the

inclination of the tunnels, First and second

instar jarvae Which remain partially enclosed

in their chorions Were found near the closed

ends of the tunnels often amongst eggs. Later

instars and pupwe were arranged m order ol

increasing age towards and facing the nest

ontraness,

195

A few 3rd and 4th instars were found foed-

ing On moist pollen masses adhering to their

venters. However, most larvac were without

food. probably because weather conditions at

and preceding nest collection were not con-

ducive to flight and food. gathering.

The durations of immature stages were not

determined. The uumbers of cach stage found

in nests are listed in Table J.

Eggs exhibited considerable size variation

even within nests. A pronounced seasonal shift

in average egy size was also noted (Fig 2) so

that the size/ frequency distributions of the June

and October samples barely overlap, The cause

of this shift is unknown,

Parasites

The only organism found ussociated with

nest colonies of &, tridentafa was an unidenti-

fied specics of minute encyrfid wasp. Lurvae of

the wasp developed within the bodies of young

bee pupae, completely consuming all tissue

except the cuticle. Porasitised pupae eventually

became filled with numerous wasp larvae which

pupated within the host cuticle, Adult patie

sites emeryed through holes chewed through the

cuticle.

Of 74 hee pupae collected in December, 39

(53%) were killed by eneyrtids, both sexes

being equally affected, As no pupae were found

in other nest samples the period of activity of

the parasite was not determined,

Foad sources

E, tridentata is polylectic and has been found

eallecting poilen from Amyema Eremophila,

Enealyptus and Melaleuca.

Colony compasition

A colony may be defined as the living

inhabitants of a single nest. The composition

of each colony found is shown in Table |, The

smallest of these colonies comprised a lone

female with one egg (nest 36). Mast nests.

however, contained two or more adult females.

a maximum of 20 occurring in nest 42. This

nest also contained gn udule male and 46

immatures (mostly eges) making it one of the

most populous found,

As mentioned above, some trynels with

lateral entrances were double-ended. In one

such Gise, 2 separate colonies (nests 26, 27)

were recognised, separate eroups of adults and

immatures occupying opposite ends of the

tunnel. In a second case (nest 22), adiilts

occupied both ends while immatures oceurred

only at ohe end, This group was treated

arbitrarily a4 a single colony,

102

December

1O-}

104

20-4

6-+--——

August

20+

> 1

E 10+

a 4

fra

) aS mes

October

—— Sr

Overall

50+

14 16 18 2.0

Egg Length (mm)

Fig. 2. Exoneura tridentata. Histograms showing

egg size frequency distributions for dif-

ferent nest samples.

Nest data reveal that brood rearing and

production of both sexes occur year round.

Eggs, larvae and adults of both sexes were

T, F, HOUSTON

present in all samples. As pupae occurred only

in the December sample, one might suspect

that eggs and larvae collected in the relatively

cool inclement months of April-September

were dormant. However, some adults in the

June sample were teneral and had obviously

just emerged from pupae. Additionally, some

adult females in all samples had ova of or near

egg size.

Every nest contained one or more eggs so

that egg production must occur intermittently

throughout the life of a colony. Were it other-

wise, eggs would not be expected in nests with

advanced progeny. With such continuous egg

production, larval hatching should also occur

continuously so that there ought to be no inter-

mediate age gaps in series of immatures. How-

ever, it will be seen from Table | that many

series do have intermediate gaps (e.g. nests

2-4, 8, 26), The rarity of Ist imstars is par-

ticularly obvious but might be explained by

their brief duration (1st instars are non-feed-

ing). The absence of later instars must be

attributed to mortality, Perhaps adults

occasionally eat eggs or young larvae: eating

of eggs has been reported amongst allodapines

by Michener (1974, p. 186).

Adult females greatly outnumbered adult

males in nests. Males from nests showed little

or no wing Wear and presumably leave their

natal nests while young. Many males were

found singly or two or three together in twigs

which did not appear to have served as nests.

Of the pupae found, 56 were females and 18

were males suggesting a sex ratio of about 3:1.

Colony development appeared to be more

or less synchronous. All 10 nests in the Decem-

her sample contained pupae while none did in

other samples. Nests of the April and August

samples contained eggs but most lacked larvae.

Such synchronisation is unexpected in a species

breeding year round and obviously some

environmental factor(s) regulates the breeding

cycle. Winter cold cannot be the factor in

view of the winter breeding. More probably it

is the lack of suitable flowers at one or more

times of the year which halts breeding and

results in synchronisation.

Within single colonies adult females showed

marked variation in size, age (as judged by

wing wear) and ovarian condition; some were

fertilized, others not, The data obtained from

examination of females are too numerous to be

tabled individually and are summarised and

analysed in the following two sections. *

* Copies of the data are obtainable from the author or the Librarian.

NESTING BIOLOGY OF EXONEURELLA

TABLE 1

Exoneura tridentata. Contents of nest twigs collected at different times. Larvae were sorted into stadia (stadium

v = prepupa). Numbers in the column on the extreme right assign each colony to one of the categories in

Table 2. + indicates pupal series affected by encyrtids.

Date of Nest Larvae Pupae Adults Pollen Colony

collection no Eggs i ii iii iv v 2 a 9 d store category

19.iv.1971 1 4 —- - -—- —- = — — +114 1 a Il

2 1 _- — 1 2 3 1 — 6 — + IV

3 5 —_- — 1 1 — 64 — 4 3 + ul

4 6 —_-_ — 1— 2 30S 2 1 + Ill

5 2 —_— 1 1 i — 3 1 5 1 + ul

30-31.xi1.1973 6 2 — 1 2 o— 2 2 203 + II

7 11 — 1 | 4 2 18; St 9 — + IV

8 1 - - -—- =— 1 3 3 3 2 + Vv

9 12 — 2 2 6 2 13 2 13 1 a IV

10 4 _ 1 i — — 6+ 4f 5 2 a cant

11 4 —_ 2 1 1 2 — 1 3 2 + II

12 4 —- -—- - - — 5 — _ Vv

13 4 -—- =| F—- O -_- — 7 — — i

14 4 oa 1 — 2 — — — 15 1 — Ul

15 3 -—- - - -—- = —- — 6 1 _— IV

16 2 - - —- - — —- — 2 — — Vv

11-16.iv.1974 17 2 —- - - -—- = - — 4 — — Tl

18 10 - - - - — 5 2 — Il

19 1 —- -—- -—|- —- = _- — 5 — — Vv

20 10 — 1 — i— — — 10 — — IV

21 8 —_ _—_ =—- — = —- — 4 — _— Til

22 7 —- -—- -—- - = —- — 3 — _ IV

23 23 — 6 —- —- — — — 15 2 a Ill

24 13 3 1- —- — —- —- 10 — = V

25 9 —- — — — —- — 6 — _ Vv

14-17.vi,1974 26 5 - =—- = 4 — —- — 8 3 — Vv

27 27 1 6 6 12 — —_- — 9 7 _— Hl

28 16 a 3 0 — 1 — _- — 6 2 _— ut

29 2 —_ =|- —_ —> — _- — 2 — — VI

30 2 —- —- -—- -—- — —- — 1 — a VI

Rp 20 _ 2 —-—- —- — —- — Ww — _ el

32 2 = 1-—- —- — _- — 2 — — II

33 14 _— 1 —- — — —- — 7 — TH

29.viii- 34 5 —- - -—- _- — 3 — — IL

1.1x.1974 35 4 —- -|- -—- -—- = —- — 1 — — I

36 1 = -—- | S| - — i— — I

37 li -- 1 -—- —- — - — 4 1 Ill

38 8 ——s tt Tbs Se: a = Vv

39 5 -—- -—|- - OC he —_- — 2 — — Vv

40 5 —- -—- -|- Se _-_ — 2 — — Wl

41 & — 2 1 1 — —-_ — 5 2 + IV

27.x,1974 42 32 — 5 3 6 — — — 20 1 + it

43 2 a ee Se + I

44 23 — 4 3 2— — — WwW — + Vv

Evidence of female castes

In many colonies one adult female was con-

spicuously larger than the remainder. In some

other nests the females were more graded in

size but smaller females outnumbered the larger

ones, The size/frequency distribution of twig

inhabiting females is strongly skewed (Fig. 3)

reflecting the relative abundance of the

smaller size classes.

As explained by Houston (1976) females

exhibit allometry where the metasoma is pro-

portionately greater in large females than in

small ones (cf. Figs 1A, 1B). This difference

may be quantified by the ‘metasomal index’: the

tatio of the distance between the lateral projec-

tions of the 6th metasomal tergum to the head

width. The frequency distribution of metasomal

indices (Fig. 4) is distinctly bimodal. Thus two

104 T. F, HOUSTON

Frequency

1.6 a

Head Width (mm)

Frequency

44 45 52 56 40 o4 68

Metatomel! Index

Figs 3,4, Exoneura tridentata, Fig, 3—Histogram

showing strongly skewed frequency dis-

tribution of adull female head widths.

Fig. 4,—Histogram showing bimodal

frequency distribution of metasomal

indices of adult females,

partially distinct morphs may be recognized,

termed here ‘minors’ (metasomal index 57 or

less) and ‘majors’ (metasomal index 58 of

more).

Majors made up 25% of the total sample of

females. In individual samples the percentages

were: December, 27%: April, 23%; June,

31%; August, 23% and October, 22%,

The ovaries of most females taken from

twigs were examined. Those rated | (tiny with

stalk-like oviduets) were considered inactive

while those rated 2-4 (small to large) were

considered active. The relationship between

ovarian activity and metasomal index (Fig. 5)

suggests that ovarian activity is more frequent

amongst majors than minors. However, the dif-

fetences between size classes might be due to

differences in the proportion of newly emerged

females in each sample. Very young females

would tend to have undeveloped ovaries. Thus

females were sorted into minors and majors

for separate analysis. The relationship of

Ovarian activity to age (as judged by wing

wear) in the two groups is shown in Figs 6, ?-

Sample numbers for older (more worn) classes

were low but approximately half the older

minors had inactive ovaries (Fig. 6), whereas

all older majors had active ovaries (Fig, 7).

Thus there is a real and highly significant cor-

relation between size (or metasomal index)

and frequency of ovarian development.

Data obtained on the presence or absence

of sperm in the spermathecae of adult females

were similarly analysed (Figs 8-10) and there

is a distinct correlation between size (or meta-

somal index) and the frequency of mating.

As Figures 5 and 8 both suggest that a fairly

abrupt change in the tendency to possess

developing ovaries and to mate occurs around

a metasomal index of 56-58, majors and

minors may be distinguished on physiological

and behavioural grounds as Well as mor-

phology,

A further inference which could be drawn

from Figs 6, 7, 9 and |O is that females of

higher metasomal index are generally longer-

lived than those of lower index: sample num-

bers above the histograms show there were

more older females (Wing wear 2—-+) amongst

majors than amongst minors, despite the fact

thal there were only about one fifth as many

young females (wing Wear O-1). It could also

be inferred that majors suffer more rapid wing

Wear than minors.

Twenty-two females were collected at flowers

and all Were minors, the largest haVing a head

width of 1,67 mm and a metasomal index of

56. The fact that no major was found outside

a nest could perhaps be explained statistically

(low frequency and small sample size), How

ever, other observations suggest that majors

are sedentary. The long ungainly bodies and

relatively short wings of majors appear

unsuited to prolonged flight and my observa-

tions suggest they may even be incapable of

short flights. While opening nests in a closed

tent | found that minors were quick to take

flight when exposed and would fly to the win-

dows. Majors never attempted to fly but per-

sistently attempted to crawl under cover, When

forced to fall, majors did attempt to fly but

NESTING BIOLOGY OF EXONEURELLA 105

73 43 | 10

-—l_""

Females with active ovaries (per cent)

Wing Wear

Metasomal Index

i Hara

Wing Weor

Figs 5-7. Exoneura tridentata. Histograms showing relationships between frequency of developed ovaries

in females of nest populations and (Fig. 5) metasomal index, (Fig. 6) degree of wing wear in

minors and (Fig. 7) degree of wing wear in majors. Sample numbers (N) for each class inter-

val are recorded above each figure.

managed no more than steep descents to the

floor. They were apparently unable to fly

upwards as did minors under the same con-

ditions.

The relatively high incidence of wing wear

among majors seems inconsistent with seden-

tary habits, for wear is usually attributed to

damage during flight. Evidence that wear must

occur other than in flight was found in majors

whose wings were reduced well beyond the

point where they could have sustained even

the briefest flight (Fig. 11). Perhaps wear

106 T. F. HOUSTON

(per cent)

Fertilized females

Wing Wear

Metasomal Index

Wing Wear

Figs 8-10, Exoneura tridentata, Histograms showing relationships between frequency of mated females

in nest populations and (Fig. 8) metasomal index, (Pig. 9) degree of wing wear in minors and

(Fig. 10) degree of wing wear in majors. Sample numbers (N) for each class interval are

recorded above each figure.

results from females scrambling one over the

other, somersaulting within narrow tunnels or

defending their nests against enemies.

Evidence of the existence of a worker caste

was provided by 5 pollen-carrying minors col-

lected at flowers, all of which were unworn and

had tiny ovaries. All but one was unmated.

To summarise, the available evidence points

to the existence of two female morphs amongst

nest populations. There are relatively few inter-

NESTING BLOLOGY OF EXONEURELLA

Fig. tl, Tattered night forewing of a large major

of Exoneura tridentata (dotted fine indi-

cOtes missing portion).

mediates and the morphs differ in their ten-

deney to mate and produce eggs. Majors,

characterised by a head width of 1.67 mm and

metasomal indices tess than 58, function (at

least in some cases) as infertile workers,

although Up to halt of them muy mate and/or

produce eggs,

Caleny development

Direct evidence of how new colonies are

founded was sought by placing 200 artificial

nest sites in the study area in June. These con.

sisted of slender pieces of wood drilled at one

or both ends and wired horizontally to the

Jower branches of Heterodendrum shrubs.

Unfortunately, when collected six weeks Jater,

none was utilised by the bees although many

had been oceupied by spiders and ants. Con-

sequently, only indirect evidence of the manner

of nest establistiment is available.

New nests are probably established by

solitary fertile minors, Four nests with single

females and immatures were found (nos. 30,

35, 36, 43). All females were mated, had

smiull ovanes and entire to moderately worn

witt targins. Two Were minors but the others

were majors, a fact which appears to conflict

with earlier evidence of sedentary habits in

majors, However, the tunnels containing the

majors were ald and stained with pollen, indi-

cating previous use as nests. The only

immatures present were a few eggs, Con-

sequenuy, the two majors may have remained

from carlier broods now dispersed and larvae

hatching from their eggs would have perished

of starvation. The female of nest 43, a minor,

must have entered w new tunnel, laid eges and

Foraged for food spots of pollen were present

on the walls and one larva had reached the

3rd instar.

If new nests are established hy solitary

females, and af these females survive until

emergence of their adult progeny, we should

expect to find nests containing one very worn

\oT

female (founder) and one or more unworn

adults (progeny), In fact, 22 such nests were

collected, Seven of the presumed founders were

minors but 15 were majors, once again

upparently conflicting with the concept of

flightless majors. All of the presumed founders

were fertilized and all but two had medium to

large ovaries, The exceptions had tiny,

apparently depleted ovaries. The presumed

daughters ranged from cullows to slightly worn,

fully matured pndividuals, Mast Were unmated

and possessed tiny ovaries but a few had

medium to large ovarres and/or sperm in the

spermatheca.

All of the above 22 nests contained eggs

and the ovaries of several presumed founders

contained ova of egg size, Thus, it seems that

mothers continue to Jay in the presence of

their adult offspring.

One nest (0. 23) contained 14 adult

Females with little or no wing wear (some were

callows) and the dry carcase of a very worn

inlfor, presumably the colony founder. Another

eight nests contained groups of 2-10 unworn

or litth: worn females and early immatures.

Evidently these pests contained groups of sis-

ters remaining after their mothers had

perished..

Groups of young sisters usually included

one or more majors and several minors.

Majors (except callows) were usually mated

and had medium to large ovaries but minors

were more variable, Drssectians showed that

Iwo Or more majors and sometimes minors in

a group may Carry egg-sized ova and probably

oviposil together in their natal ests, Since no

female examined had more than four ova near

egg size (few had more than two), large clus-

ters of segs most probably arise only where

two or more females are laying together, For

example, nest 44 which contained 23 eges had

three fertile majors with medium to large

OVATrIES,

Undoubtedly, some females in groups of

young sisters disperse to establish new nests

but there js evidence that others remain

together in small colonies: sever nests cach

conthined two or three very worn females along

with newly emerged adults. In each case, one

female was a major, fertillsed and with

enlarged ovaries, the other ane or two bemg

unmated and (except one) with undeveloped

ovaries, In thesc colonies, the majors must

have functioned as egg layers, {he minors ss

workers, The newly emerged adults im such

nests had presumably teen reared by the worn

108

TASLE 2

Eaniwura tridentita, Classification of mest cahieies (ie.

thase centatiing immatures) based ae iaimbers of

adult fertales, their size and degree of wing wear.

ree of

Category quency total

1, A solitary female a I]

if. One very wart minor and one

or Mare wnworh ar slightly

warm females 7 16

ME, One very wore major and one

or more unworn of slightly

worn Femiles is 4

WV. Twat ar mare very worn females

and one of more unworn ar

slightly worn forrales q 16

¥. Two to several unworn or

slightly worn females only In un

VL Other t 2

individuals. In most cases, the eggs in each of

the nests could only have been lald by the

Worn majors, but ina few nests one or iwo of

the young females had enlarged ovaries and

may have oviposited,

If the belief that majors are sedentary and

cannot forage for pollen and nectar is correct,

the 15 colonies cach containing a single worn

major with one or more unworn females appeat

anomalous. However, the young females (plus

males and immatures) may have been reared

by worker-like females which died when the

young began reaching adulthood and which

were survived by their longer-lived major sis-

ters, Brood care may have become the respon-

sibility of the newly emerged females.

In 14 groups of newly cmerged sisters

there were up te S (mean 2,5) majors per

group, Tf majors cannot disperse by Hight, one

would expect to find groups of old worn majors

in at least some nests. However, amongst the

nests examined, not One contained more than

a single worn major. Either the concept of

sedentary majors & incorrect or there is some

process by which all but one major is even-

tually eliminated from groups of sisters. If this

process took the form of physical contests, it

could explain the existence of female allometry,

As explained by Houston (1976), the meta-

soma of relatively large females is more

heavily chitinised, were muscular und more

scoop-like at the apex than that of smaller

females. Larger females thus appear to be

better equipped for posteriorly directed combat

than smaller ones (these modifications should

also be of benefit in mest defence: if majors

remain in nests while minors forage. they would

tT, F. HOUSTON

hear a greater share of the burden of defence

of the colony against intruding ants and other

depredators and it is Uhe usual mode of defence

of allodapines to block their nest entrances

with the dorseapical surface of the metasoma).

Obviously, direct observation of living colonies

is required to sce if majors really do engage in

physical elimination contests,

The categories to which nests were assigned

according to their female contents are listed

in Table 2 with indications of relative

frequency. In Tyble | each nest is assigned to

one of these categories as indicated by the num:

bers in the last column.

Fmally, iL remains lo mention 14 female-

inhabited twigs which contained no immatures.

Interpretation is difficult, Seven twigs contained

solitary minors which were relatively unworn,

mated, had small to large ovaries and which

may have been about to found new colonies.

The remaining seven twigs contained 2-6 adult

females. Two each contuined 9 pair of very

wiirn individuals that could have funetloned as

ege-layer and worker and may have survived

after departure of their offspring, The other

five twigs contained unworn females (three

included majors) which may have beeu reared

in the twigs, for the walls were dark-stained

and showed traces of pollen,

While many aspects remain uncertain, the

course of colony development envisaged may

he summarised as follows. New nest sites are

sought out and occupied hy relatively young

mated minors with enlarging ovaries. Each

female begins to produce eggs intermittently

wid when larvae emerge she forages and feeds

them progressively. She vontinues to lay even

when most of her offspring have reached adult-

hood, Amorigst the brood are males, and minor

and major fernales, For a time, mother and

adult offspring may cohabit, some or all of the

daugilers mating and eventwally contributing

eggs to the colony. Same minors fail to mate

and to develop enlarged ovaries and function

4s Workers tending the combined brood.

Eventually the mether dies and the males and

most fentales disperse to other twigs. However,

one major and one of more minors remain,

functioning us queen ond workers, respectively.

They do not perish until second generation

adults emerge and assume care of the

immatures, Once agin, the voung adults may

contribute eggs to the colony before mast dis-

perse leaving One major and one or two worker-

like minors to tend and protect the Immiatures,

NESTING BIOLOGY OF EXONEURELLA

To all probability, the allometry of adult

females is trophically controlled as in other

insects, those individuals receiving relatively

larger amounts of food as larvae developing

into relatively larger adults with certain features

exuggerated, This being so, ofe might expect

that generations raised during periods of food

abundance might contain a relatively higher

proportion of major females than generations

raised during periods of food scarcity, How-

ever, the facts do not support this expectation:

pollen accumulation in December and October

nests was taken as a sign of food abundance

preceding their collection but the highest per-

cenlage of majors occurred in the June sample,

the nests of which (and the preceding April

sample) lacked pollen stores. Therefore, some

social mechanism must control the minor/

major ratio,

Exoneura eremophila Houston

Nests of this tiny arid-dland bee were

collected and examined in haste during the

course of other work in southwestern Queens-

land and northeastern South Australia and the

account Lo follow is based on my rather meagre

field notes. Dissection of females was not

possible so that data on mating and ovary con-

dition are not available,

A total of 45 nests and shelters of this

species were found in dead dry hollowed stems

of herbaceous plants. Most occupied stems of

Crotalaria cunninghamii R. Br, (Fabaceae) and

Myriocephaluy siuartii (FVM.) (Compositae),

while a few were in unidentified stems. All

nests were within 30 cm of the ground,

In most cases, the bees had burrowed into

the soft pith exposed at the broken ends of

upright or oblique stems, However, some

individuals had utilised naturally hollowed

stems, and barricades of pith particles formed

the bottoms of the occupied sections of tunnel,

Hollows containing immatures ranged in depth

from 25-190 mm (mean = 81,5, N = 40),

Entrances were circular and of slightly smaller

diameter than the tunnels lower down (e.g. 2.0

mm compared to 2.5 mm), None possessed

a constructed “collar” of the kind so typical of

nests of Exeneura s. str. and Braunsapis but

two entrances Were narrowed by small cres-

cents of compacted pith particles,

immatures

The disposition of immatures in the ttinnels

was generally as described for E, tridentata and

the numbers found in. each nest are listed in

109

TABLE 3

Exoneura eremophila. Numbers of adults and

immatures taken from stems with collection data and

tunnel lengths.

Tunnel

Collection Stem length Lar- Adults

dati no, mm gys vac Purae & a

LoT | 1 74 3 —_ — j =s

13 km NE zZ 190 6 — SS

of Windorah, 3 118 4 4 — i _

Qld, 4 9T 1 6 = I —

14. Viit.1968, 5 62 4 —_ _ L —

Collected at 6 ron 6 — _ L =

midday 1 145 ~' ra =

LOT 2 5 103 7 — — 1 _

Same Loc. 9 90 7 — — 1 1

19, vill 1968. 10 67 1 a — i a

Collected in 1 30 4 1 _— L —

cool early 12 8+7— _— a i —

morning i) a4 3 — —_ L _

LOT 3 a 78 1 t _ =

Skm W of 15 25 — i i — 2

Windorah, 16 67 4 3 _ — _

Old, 7 43 _ 1 — — wd

V7 dv 869. 18 78 — 2 —_— —_ —

Collectedin 19 a5 — 6 — _- —

mid-morning 20 53 2 in —_ _ L

Z1 a —_ ag 1 t

22 61 -_-_ — — 1 —

LOT 4 23 —_-_ = _ — i =

New 24 135 2 \7 z 5 t

Kalamurina 25 116 I 5 4 3 3

H.S., § 26 96 3 = — I —

Aust, 2 68 6 1 - 1 —

10.0, (972, 28 6 § 6 1 1 5

Collected 29 33 8 — - 1 —_

incoolwarly 30 05 i 6 8 3 —

morning 3 x 8620 ' 2 3 =

32 va 4 — —_ 1 —

ca ST 6 _ _ 1 —-

34 1a a 14 4 4 z

35 50 5 4 ! 1 2

36 19 iv s 3 3 2

3’ 44 5 4 _ 1 4

33 120 2 9 6 4 2

39 a 7 7 6 1 z

40 162 13 w 9 1 4

41 35 - =— -_-_ —- I

#2 66 & _ _— L 1

43 69 9 LF _ 1 _—

44 36 6 3 _ 1 4

45 85 & 3 — I 1

Table 3. The immature stages are described

by Houston (1976),

Food seurces

E, eremophila is a polylectic species and has

been observed collecting pollen from Calan-

drinia, Eremophila, | Goodenia, Hakea,

Helichrysum, Myriocephalus, Scaevola and

Wahlenbersid.

Colony compasition and development

Nest collection data and the number of occu-

pants of each aré provided in Vable 3, Nests

were collected in four lots, the Isc and 3rd

when conditions suited adult fight and some

occupants Were oul. However, lols 2 and 4

were collected in cool early mornings so that

all actult occupants should have been present.

The August sample (lots 7 and 2) was com-

posed chiefly of single female nests with eggs.

Four nests also contained larvae of various

ages. some of then defaecating, Two stems

(nos §, 12) contained females but to

immatures and were probably under excava-

tion, The highest number of progeny was 12

in nest 7.

Most nests in the Apeil sample (lot 3) lacked

adult females which must have been foraging

at the time of collection, As the highest num-

ber of progeny in any nest was 13 (nest 17),

it is probable that all nests had been founded

by single females. Three females occupied nest

21 but, as a dark female pupa was also present,

auy or all of them may have been newly

emerged daughters of the founder. Eight nests

contained larvae but only 3 had egus as well,

suggesting that egg prodiction ceases after a

short period of laying, The burrow of stem 22

was obviously under excavation when found.

The origin of the males in the April nests

Temains uncertain,

The 23 nests in the March sample (lot 4)

Were generally more populous than the others

and their occupants more diversified, Fourteen

contained single adult females but cight each

contained 3-7 females! In these nests, the

presence of pupae suggests that al least soine

females may have been newly emerged, How-

ever, the presence of relutively larze dumbers

of immatures jn some mests with several

females and age gups in some series of

immatures suggests that 2 or more fermales had

contnbuted to the brood.

The nests of Jot 4 may be considered in three

groups. The ten nests of the first group (nos

26, 27, 29, 32, 33. 37, 42-45) were much like

those of lots 1-3, each having a single adult

female fand males itt some) with up to 12

ivtmatures which were all either eggs or larvae,

Three nests of the second group (nos 28, 35,

39) also had a single adult female each fond

males) with immaliires, However. the

immatures of cach nest incilnded eges, larvae

and pupac and totalled 20 ja nest 39. All stacin

Were represetiled in two nests, stigegesting cof-

tinuous und prolonged ege production but in

T. F. WOUSTON

nest 35 an age gap existed and all larvae were

mature. In the third proup, cight nests (nos

24, 25, 30, 31, 34, 36, 48, 40) each contained

frony 3=7 adult females (and males in some}

with eggs, larvae (or pre-pupac) and pupae.

In all bul one nest, immatures totalled 21741

und must surely have been derived from more

than one mother (note especially the 20 egas

of nest 3L), A conspicuous age gap (the

absence of 3-4 consecutive larval stadia) was

noted in three nests and 3 moderate gap (two

consecutive larval stadia absent) in two others.

All stadia were present in the remaining three

nests.

Without datn on degree of wing wear, ovary

condition and spermathecal content it is diffi-

cull lo interpret the relationships of females in

the nests found. However, from the observa-

tions made one could speculate that colony

development proceeds along the following

lines: new colonies in some cases may be

founded by solitary females each laying up to

12 eggs then ceasing while they rear the emer-

gent larvae. The immatures of such a nest

would not span all stages (as in lots 1-3), In

other cases, founding females may oviposit

intermittently over jong periods, sometimes

with temporary halts, producing series of

immatures spanning all or most stages, Young

adult females emerging at intervals in these

nests May oviposit, too, so maintaining a more

or less continuous senes of bnmatures. Whether

such colonies are communal with each sister

partaking of foraging and broad care or semi-

social with division of labour can only he

revealed hy further studies,

Mole behaviour

Many dozens of males of this species were

observed in Hight near nests on Kalamurina

Staton, S. Aust., during midmorning of

1.uL1972, Each hee flew erratically about 30

cm above eroywnd amongst the many dead

Myriocephalus stems and frequently hovered

about a nest entrance before moving on.

Wahilenhergia flowers growing in profusion

nearby were visited only occasionally by the

males,

Presumably these males Were awaiting the

emergence of virgin females from nests, but no

encounters hetween the two sexes were

observed.

Exoneura setosa Houston

Observations of the nesting biology of this

species Wel'e made during 14965 in coastal dures

at Nerth Glenelg and West Beach, S. Aust

NESTING BIOLOGY OF EXONEURELLA ti!

Nests were collected primarily as a source of

live stages for morphological studies and many

data now desired were not recorded, However,

the availuble information provides a reasonably

clear picture of the life cycle of the species,

The dates of collection of inhabited stems

and the numbers of inhahitants are provided in

Table 4. Stems 12-24 were collected jn cold

weather unsuitable For adult flight. However,

the remainder were taken at times when some

adults may have been absent on foraging trips.

Nests

All colonies and sheltering adults were

found in dead dry pithy stems of herbaceous

plants and grasses, Most nests were in stems

of Euphorbia, Geranium and Foeniculum. Nest

burrows had apparently been excavated by the

females or, in the casé of naturally hollowed

stems, had been refined by the removal of

irregularities and debris, The bees entered the

stems al broken ends and burrowed down their

lengths. The burrows were 1,5~2,.0 mm in

diameter and 35-95 mm deep. Their entrances

were circular and showed no traces of any

special structures. Occupied stems were

variously inclined from vettical to almost hori-

zontal, All Were less than | m above groynd

and most below 30 cm,

Imumatures

The disposition of the immatures in nest

burrows was penecrally as described for E, tri-

dentata. The immature stages were described

by Houston (1976),

Food sources

This species, like the preceding two, is poly~

fectic. Pollen taken from nests was derived

from Cakile, Geranium, Reichardia and

Walilenbergia.

Colony compoyition and development

As will be seen from Table 4, seven nests

(nos I, 2, 4, 6, 7, 9, 11) each contained a

single adult female with 2-9 immatures, Three

other nests (nos 3, 5, 8) each contained 2-4

adolt females with 3-6 immatures. However,

as each of the latter nests also contained pupae

and two contained adult males, probably all

but one female in each were newly emerged

progeny. Assuming this to be so, the total

progeny in each nest would have been 9, 7 and

9 respectively. Thus a single female may pro-

duce up to 9 offspring,

Females must produce a batch of eggs with-

in a relatively short period and cease laying as

the first larvae eclose for, although immatures

TABLE 4

EBxoneuri setosa. Numbers of adults and nmatures

taken Jram stems of West Beach, Adeluide, §. Aust,

Date of Stem Lar- Pre- Adutts

collection no. Epys vse pupwe Pupue 2 a

5.11965 1 _ fr z 1 L —_

11.41.1965 z gE i _ i —

2 _-_ — i 3 4 2

4 2 i 2 1 i

16.41.1965 5 — 4 1 1 z2 —

6 _— i _ 1 ‘ _

7 _ 3 i — rc —

i ——s ea 3 3 4

10.11.1965 9 — 2 — — i —

w = — - — 2 2

il _ i i _ i —

Ps _ - = 40 —

13 _— 2 =—- — 3 rs]

Siy.1965 gh =| — SS 7 5

15 —_ —_ —_ _— 2 1

16 - =—- =— = L L

17 _ = = 1 to—

bo — — — — tt

19 _ - =—- = 5 8

(2.yti.1965 20 — | _— —_ i 5

21 _— _ — — 1 5

2 - —- > — 4 3

18.yiii,1965 3 _— _— — — 1 —

14 Le. £965 oe ne — a ae

= s = — = 2 |]

312.1965 Yo - © 7}

27 6 = =—|- =

within individual nests were graded m age, n0

nest had the full range of immatures.

No nest provides definite evidence of brood

rearing by more than a single female. Although

nest 26 contained two adult females, a male

and seven eggs, there was nothing to suggest

that these adults were more than siblings re-

maining in their natal nest; one female may

have been founding a new colony and the

other adults may soon have dispersed,

While brood rearing was evident in stems

caflected in February, March and October,

those collected in April and July appeared to

contain overwintering groups of adults of both

sexes and a few residual immatures (the larvac

of nest 13 were shrivelled). All adults from

these overwintering groups had unworn wings

and fresh body pubescence. Seven females

(from nests 16, 18, 19) were dissected and

proved to have empty spermathecae and slen-

der undeveloped ovartes.

Most overwintering adults were taken trom

old nest stems but some occupied rough

112

natural hollows, Because four overwintering

groups (nests 13, 14, 19, 20) each contained

12-L5 individuals and no brood nest contained

more than nine progeny, one would doubt that

they represent farmly groups alone, Evidence

of wecgariousness amongst unrelated over-

wintering adults was obtained using artificial

nests. These consisted of Foeniculum stems into

which glass tubes of 2 mm bore had been

inserted and split stems where a clear cellu-

loid strip formed one side of a burrow, Fifty

such nests were placed in the study area but

only two were uiilized as shelters by solitary

females, However, in Augtst, three groups of

adults found in natiral nests Were transferred

to anilicial observation nests. Two groups (10

9, So and 12, 40) gradually dispersed during

September. However, the third group (4, 9, 3¢)

had increased by two individuals after several

days and by four more after two weeks. The

13 adults femained together through Septem-

ber and yellow pollen stains withm the tunnel

enirance testified to foraging activities by at

least one member, By late November most

individuals had gone bul two five and two dead

bees remained.

Five females found sheltering in stems 23

and 24 in August and Septemher were

dissected. All bad mated but their ovaries were

small and no ova wete near egg size,

A total of 105 adults were collected frorn

stems and comprised 63 females and 42 males.

suggesting a sex ratio of approximately 3:2.

To summarise, nests of E. setose appear to

be founded and mamtained by solitary females

which may each produce up to nine offspring.

Adults of both sexes and occasional immatures

overwinter in old nests or congregate in

naturally hollow stems in groups of up to 15

individuals. Females overwinter as virgins but

mate during the spring and eventually establish

new tests.

Discussion

Details Of the nesting biology of Axencura

lawson? have been provided by Michener

(1964). At the time of his study, &. serova had

not heen recognised as a Separate species and

some lowland nest samples may have belonged

to this species. However, Professor Michener

advises me that the bulk of his material, if not

all, was E. lawsev.

All Exonenreila are typical of allodapines in

rearing their larvae in oper tunnels in plant

stems, However, their nests are characterised

by the consistent absence of constructed

T, F. HOUSTON

entrance collars. Three species (eremaphila,

lawreni and setosa) excavate nest burrows in

dead pithy stems of herbs and normally the

entrances are fiarrower than the remainder of

the tunnels, The tiny hody size of these species

78 perhaps an adaptation to life in slender stems,

&, tridentata, by contrast, utilises ready-made

burrows in dead woody twigs of standing trees

and shrubs. Both these methods are employed

by other Exonevra: most members of the sub-

genus Exoneura which have been studied

burrow in pithy stems (Michener 1965) but

some species will occupy trap nests; members

of the subgenus Brevinedra habitually utilize

existing Woody hollows (personal nbserva-

tions).

Like Exoneura s. stt,, Exoneurelle lay their

eggs freely in the burrows rather than attaching,

them to the walls with adhesive secretion us do

Brevineyra, The duration of oviposition varies

amongst the species, being restricted to the

early phase of colony establishment in seteva

but occurring continuously in lawson! and rri-

dentata. In eremophila oviposition muy occur

continuously or be interrupted.

Middle larvae and older immatures are

moved about by the bees and arranged Imearly

in approximate order of ages as in most other

allodapines. Feeding is progressive and, in

Keeping with the year-round actlvity of adults.

all species are polylectic.

Exoneurella appear to be relatively {ree from

pathogens, the only one known heing ali entyr

tid which Was feared from summer pupae of

tridentata.

Brood rearing in fewseni and yetosa ceases

during winter when the populations consist

largely of adult males and females (unmated).

Mating and nest establishment resume in

middic of late spring. In ¢ridentate, which

occupies 4 rather more temperate habitat than

the preceding two spevies, brood rearing occtirs

year round (possibly with brief halts during

flowerless periods}, The situation in ereriaphily

is unknown,

The four species exhibit pronounced dif-

ferences in their degree of sociality, Both faw-

soni and setosa appear to be almost totaily sub-

social, cach fest being founded and maintained

by a single female, At most, a fernmle may care

for her immature siblings after dernise of her

mother. £. erentophila, too, is basically suh-

social but evidence was found that two or more

females may occasionally oviposit and rear

breed together in a common bunnw, By con-

trast. tridentata develops semisocial colonies

NESTING BIOLOGY OF EXONEURELLA

and has morphologically distinct castes. While

subsocial colonies may be founded by solitary

females, they later become semisocial with two

or three sisters functioning as queen and

workers, The social structure of tridentata is

generally similar to that of members of the

subgenus Exoneura (especially E. variabilis)

studied by Michener (1965) and various other

allodapines (Michener 1974), However, tri-

dentata is outstanding amongst its relatives in

respect of caste differences. In the majority of

social allodapines the castes are indistinguish-

able morphologically, differing only in the

functions they perform and in the states of

their ovaries and spermathecae. Queens of

Exeneura variabilis average larger than their

worker sisters. In tridentata the size dif-

ferences are accentuated and accompanied by

allometry (Houston 1976). In addition, queens

of tridentata appear to be flightless and are con-

fined to nests.

The biological characteristics of the four

species of Exoneurella reflect their relationships

as determined on morphological grounds; tri-

dentata stands out from the other three and

shows some similarities to Exeneura s. str.

Michener (1964) noted a seasonal variation

in sex ratio of E. lawson’ which he felt was

inappropriate to a strictly subsocial species and

possibly a residual characteristic from a more

social ancestor. The discovery of tridentata and

its social habits supports this idea,

113

The loss of semisociality m Exoneurella may

be associated with the use of rapidly perishable

pithy stems of annual or ephemeral herbs,

Since such stems may only be suitable for

occupation for several months, groups of sib-

lings would be forced to disperse to new sites

far more often than if living in durable woody

twigs. Thus the opportunities for formation of

semisocial groups in old nests would be

diminished and _ strictly subsocial behaviour

would be favoured.

Acknowledgements

1 am extremely grateful to my wife, Carol

Houston, for her patient assistance in the

search for and collection of fridertata nests, |

am indebted, also, to the South Australian

National Parks and Wildlife Service for per-

mission to study this species in the Lake Gilles

National Park and to Professor Charles D.

Michener of the Department of Entomology,

University of Kansas, for advice concerning

material used in his study of /Jawsoni, 1 thank

Dr Eric Matthews of the South Australian

Museum for reading the manuscript and mak-

ing useful suggestions for ils improvement.

Fieldwork expenses associated with the #i-

dentata study were met almost wholly by a

grant from the Royal Society of South Aus-

tralia's ‘Scientific Research and Endowment

Fund.

References

Houston, T. F, (1976) New Australian allodapine

bees (subgenus Exeneurella Michener) and

their immatures (Hymenoptera: Antho-

Bees) Trans. R, Soc. S, Aust. 100(1),

15-28.

Micuenrr, C. D. (1964) The bionomics of

Exoneurella, a solitary relative of Exoneura

(Hymenoptera: Apoidea: Ceratinini). Pac.

Insects 6039), 411-426,

Micuener, C. D. (1965) The life cycle und social

organization of bees of the genus Exoneura

and its parasite, Inquilina (Hymenoptera:

me he Univ, Kans. Sci. Bull, 46(9),

335-376.

Micuener, C. D. (1974) “The Social Behaviour

of the Bees. A Comparative Study,” (Harvard

University Press: Cambridge, Mass.).

VOL. 101, PARTS 5 & 6 31 AUGUST, 1977

TRANSACTIONS OF THE

ROYAL SOCIETY

OF SOUTH AUSTRALIA

INCORPORATED

CONTENTS

Angel, L. Madeline and Pearson, J. C. Dicrocoeliid trematodes from Australian

hosts - - - - - - - - - - - 115

Tyler, M. J., Davies, Margaret and Martin, A. A. A new species of large, green

tree frog from northern Western Australia - - 133

Wells, R. T. and Nichol, B. On the manus and pes of fieatiee carnifex Owen

(Marsupialia) - - ~ - - - - - 139

Dulhunty, J. A. Salt crust solution during fillings of Lake Eyre - = = - 147

Cane, R. F. Coorongite, balkashite and related substances—an annotated

bibliography - - ~ - = = = = - 153

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DICROCOELIID TREMATODES FROM AUSTRALIAN HOSTS

BY L. MADELINE ANGEL AND J. C. PEARSON

Summary

Eleven species of dicrocoeliids belonging to five genera are described; nine are new.

Brachylecithum comprises seven species (six of which are named) separately by size, width/length

ratio, acetabulum position and egg size. Skrjabinosomum is represented by three species, two of

which are named. Lutztrema and Pancreatrema have one new species each. Proacetabulorchis is

recorded from Australia.

DICROCOELIID TREMATODES FROM AUSTRALIAN HOSTS

by L. MaDELINE ANGEL* and J. C. PEARSON?

Sumumnary

ANGEL, L. MAbgLINE und Pearson, J. C. (1977) Dicrocoeliid trematodes from Australian hosts.

Trans, R. Sov. S. Aust. 15), 115-132, 31 August, 1977.

Fleven species of dicrocoeliids belonging to five genera are described; nine are new.

Brachylecitthum comprises seven species (six of which are named) separated by size, width/

length ratio, acetabulum position and egg size. Skrjabinosomim is represented by three species,

two of Which are named, Luizrrema and Parnereatrema have one new species each. Proacetabu-

larchis dovleli is recorded from Australia,

Introduction

Sandars (1958) described Platynesemum

australiensé (syn, Zonorchis australiernsiy) from

Australian marsupials, and listed seven other

dicrococliids recorded from Australian reptile

and bird hosts, Of these, Paradistomteun crucifer

(Nicoll), originally described from Delma

fraser!, has since heen recorded from four other

lizards (Angel & Mawson 1968), The only

other dicracoehids recorded from Australia are

Platynorreme biliosum Nicoll from Burhinus

magnirostris (svn. B. grallarius) and Threskior-

nly mluece (ayn. This molucca), P, jecoris

Nicoll from B. magnirosrris, and Atheymia sp

from Rattus norvegicus (Munday 1966),

We now add five new species of Brachy-

lecithum, one unnamed Braechylecithum species,

two new species and an unnamed species of

Skrjahinosomum, a new species each of Lutz-

trema and Panereatrremna and Proacetabulorehis

dovicli Belopolskaja & Bychovskaja-Pavloys-

kaja,

All of the hosts of trematodes recorded in

this paper are qutive to Australia; only Petra-

chelidon nigricans is migratory, and Ardea

novaehollandiae, though it has no regular

migration, sometimes occurs Outside Australia.

The tood of the definitive hosts of Brachy-

lvcithum species is given as indications of the

possible intermediate hosts. Information on

food was obtained from Cleland (1910), Cle-

land, Maiden, Froggatt, Ferguson & Musson

(1918), Leach (1958), and from dissection

* Department of Zoology, University of Adelaide.

‘Tce, Adelaide, S. Aust. 5000.

records of the Department of Zoology, Univer-

sity of Adelaide. Where the life cycles of dicro-

coellids are known, the first intermediate host

is a terrestrial gastropod, and the second, an

insect, arachnid or terrestrial isopod,

All drawings were done with the aid of a

camera lucida. Where measurements are given,

the mean is in brackets after the range.

Holotypes and paratypes are deposited in the

South Australian Museum (SAM). Type

material and slides of other specimens men-

tioned are deposited in the former University

of Adelaide Helminthological Collection (now

located in the South Australian Museum) and

in the second author's collection. Additional

measurements from the type and other hosts

are available from the authors on request.

Measurements of suckers are means,

LIST OF PARASITES, ARRANGED WITH

THEIR HOSTS

Mammals,

Rattus fuseipes (Waterhouse). Brachyleci-

thom insulare wisp. Hydromys chrysogaster

Geoffroy. Brachylecithum hydromyes n.sp.

Birds.

Ciconiiformes: Ardeidac.,

Ardea novaehollandiae Latham. Proacet-

abularchis degieli Belopolskaja & Bychoy-

skuju-Pavlovskaja.

Gruiformes: Turnicidae.

Turnix castanota (Gould).

SOMTIN MaWwNOni n.sp.

Skrjabino-

Present address: South Australian Museum, North

} Department of Parasitology, University of Queensland.

116

Caprimalgitormes; Podargidae,

Podargus strigoides | Latham). Braciyleei-

thuim podargil rsp.

Coractiformes: Alcedinidae.

Dacelo novaeguineae (Hermann), Brachy-

lecithin dacelonis n.sp.

Passeriformes: Hirundinidae.

Petrochelidon nigricans (Vieillot), Brachy-

lecithum parvum (Johnston),

Passeriformes: Timahiae,

Pometostomus supercilliasuy (Vigors &

Horsfield), Skriabindsomum pomatostomi

Tsp.

Possenfornmes: Muscicapidae.

Microecu leucophaca (Latham),

Skrjabinesomun sp.

Passeriformes: Meliphosidae.

Meliphaga ernata (Gould), Pancreatrema

meliphagae onsp. Manorina fluvigula

(Gould). Skejabinosomum mawsoni n.sp,

Passeriformes: Cracticidac,

Cracticus terquatus (Latham), Brael\yleet-

thum latius n.sp. Gynnorhina Apyoleuca

(Gould). Brachylecithim latius n-sp,,

Brachylecithum sp.

Passeriformes: Ptilonorbynchidae.

Alhoedus crassirostris (Puykull).

treme ailurocd: \\,sp.

Passeriformes: Corvidue.

Corvus coronoides Vigors & Horsfield.

Brachkylecithient latius asp.

Cervus mellori Mathews, Brachyleeithum

parviim (Johnston),

Reptiles.

Aniphiboluruy fiona’ Proctor. Brachyleci-

dim fnrulere nsp.

BRACHYLECITHUM Shitrom

Denton (1945) stated that Brachylecithum

spp. were very delicate worms thal showed con-

siderable morphological variation as a result of

host, habitat, age and hereditary Factors [sic]

und because of different methods of handling.

Me wlso noted that essential taxonamic charac-

ters were lacking for many species which were

inadequately described, or described from poor

material, The genus then contained 35 species

and subspecies. The number is now aver eighty

and the observutions of Denton stil] apply, Te

seems certain that when 4 thorough revision is

made muy species will be found to be synony-

mous. In the meantime it is not possible to

compare adequately the Australian species with

all previously recorded species, We have com-

pared our specks with any which occur in birds

of the same or closely felated genera in other

Lutz-

L. MADELINE ANGEL. & J, C. PEARSON

parts of the world, Since only three species

have heen recorded previously from mammals,

we have compared &. inswlare nsp. and B.

Avdromyes n.sp. will all three.

Brachylecithum and Lyperesomum are very

similar genera which have caused taxonomists

confusion, From the diagnoses of Skrjabin &

Evranova (1953) after Shtrom (1940) and of

Yamaguti (1971), it seems that the only con-

sistent character used to separate them is the

vitellaria, consisting of a small number of large

follicles, occupying a small part of the body

length in Brachylecithum, and in Lyperosomum

of numerous small follicles occupying a greater

distance in relation to body length,

Yamaguti separated Brachylectthum into two

subgenera, @ruchylecithum and Brachyleci-

thotdes on the presence or absence of the uterus

hetween the testes and between the testes and

ovary. This is a character which varies in speci-

mens from one host (compare Figs | and 2),

so that we have not used it. Further, the size

and even the shape of the testes and, to a Jesser

extent, of the ovary, vary from one specimen

to another, Both characters may depend on

the maturity of the trematade or on tts stute

of extension when fixed.

Specific characters used are size, width/

length ratio, position of acetahulum, presence

or absence of papillae on surface of body, and

size of eggs. In view of the yvaniability of ace-

tabular shape we have uot considered it

advisable to compare sucker ratios,

Key to Australian species of Brachylecithum

|, Width/leneth ratio of body greater than 1/9 .

RB, latins

Width/length ratio of body Jess than 1/9 2

2. Widthength ratio of body between 1/10 an

TA2S asin ree

Width /lengih ratio of body between 1/24 te

1/30 4. 5

3. Acctabulom between anterior 1/4-1/5 of body;

withoul papillae on body surface; eggs 36 x 20

um BR. hydromyos

Acetabulum within anterior 1/5 of body ,,. 4

4. Acctabulum between anterior | /S-1/6 of body:

puna papillae on body surface; eges 43 x 72

. B, insulure

‘Arcsalyulana between anterior 1/6-1/7 of boxy:

with papillae on body surface; eggs 39 x 23 em

&. parviem

5. Acetabulum between anterior 1/4-1/6 of boxty:

with papillne on body surface; eggs 39 x 24 am

B, dacelonis

Acetabulur between anterior 1/7-1/10 of basty;

withoul papillae on body surface, eges 47 x 24

em B. podargi

Characters from the type host i in all cases,

DICROCOELIIDS FROM AUSTRALIAN HOSTS

ego eo

> 2 0

SeSS

sala

1 «OO-5 mm

Figs 1~6. 1. Brachylecithum insulare Holotype; 2. B. insulare Paratype SAM V77; arrangement of testes

and ovary in specimen with comparatively few eggs: 3. B. hydromyos Holotype; 4. B. parvum

from Petrochelidon nigricans; 5. B. dacelonis Holotype; 6. B. podargi Holotype (Figs 1 and

3 to same scale; Figs 2, 4-6 to same scale), ABBREVIATIONS: a, acetabulum; c, cirrus; e.b,

excretory bladder; i, intestinal caecum; M, Mchlis' gland; 0, ovary; 00, ootype; r.s, receptacu-

lum serminis; t, testis; tl, t2, anterior, posterior testis.

dL8

The eggs measured in this study have been

the largest mature specimens from balsam

mounts. For egys of Brachylecithuny parvam

(Johnston), Johnston's measurements are used.

as the eggs in his slides are now unsuituble for

ecurale measurement,

Generic diagnosis: Dicrecoeliidae, Dicro-

cocliinae. Body slender, filiform or lanceolate,

flat or cylindrical; maximum width/length ratio

not less than 1:45; with pre-oral lobe, Body

surface with or without spines or papillae.

Suckers usually subequal, not far apart. Cacca

not reaching to hind end of body, Testes tar.

dem or sometimes diagonal, contiguous or

separate, immediately or a little posterior to

tteetabulum, occasionally underlying latter. Cir-

fus pouch pre-acetabular, Ovary round or oval,

dlightly behind testes. Vilellaria post-ovarin,

commencing near ovary, limited in extent, con-

sisting of small number of large follicles.

Parasitic in liver, gall bladder, pancreas or

intestine of birds, mammals and reptiles.

Only three species of Brachylecithwm have

been described previously from Australia

(Johnston 1917). Johnston assigned them to

Lyperesomum, but Skriabin’ & Evranova

(1953) and YVamaguti (1971) placed all three

in Brachvlecithum. The species are B. parvum

from Strepera versicolor, B. megaustomum trom

Sterna herell and B, harrisoni fram Ninox

novaescelandiae (syn. N. boohook). We have

examined the types of these species. In addi-

tion, in T, Harvey Johnston's collection is a

slide of B. parvum and two of B. megastomim

with S. J. Johnston’s name, and other particu-

lurs On cach showing them to be from §. J.

Johnston's original material, as well as a

mounted specimen complete, except for oral

sucker and a small part of the anterior end,

which bears 4 Zoological Laboratory, Univer-

sity of Sydnev. label written in T. H. John-

ston’s hand “? Lyperosomum, Boohook owl"-

We have compared measurenients of all of

these, and find it impossible to differentiate

helween the three species. There is some varia-

tion between individuals of cach species, such

as the presence of coils of the uterus between

the gonads, The specimen of B. harriyont desig-

nated as type is incomplete and is in three

picees; the anterior end (oral sucker ta hegin-

ing of vitellaria) ts not a good preparation, the

gonads being largely, and the acetabulum

partly, obscured by eges. S. J, Johnston stated

thal &, harriveni was closely related to B, per-

viain—"indeed, all three species are closely

related to ove another". With the material

L. MADELINE ANGEL & J, C, PEARSON

available to us, We hesitate to synonymise these

three species.

Brachylecithum parvum (Johnston)

iG, 4

Lyperosommen parveni Johnston,

Strepera versicolor, N.S.W.

Brachyleelthim poarven: Skrjhin & Evranova,

1953.

Haw, Petrachelidan nalgricans,

Locatian in fest, Liver and yall blake,

Localire, Mannum, 3 Aust., February 1972.

Jacidence. 1 of Z birds from sare locality at same

time.

Food of hast includes insects.

Slides deposited, SAM V85, V8G6. Identification

based on seven balsien mounts. Other specimens

loo twisted or broken to be of wny use,

Rast. Cervus mellori.

Levdton Jn lest, Bile duct and/or gall bipdder

Locality. Tas. September 1969,

Incidence. 29 specimens. in one oF 48 birds Trom

Tas. (24 pa July 1967, 24 in September 196%),

In none of 9 Corvus melloré fram S. Aust,.

1966-1970.

Food of host includes insects.

Slides deposited. SAM YR7, Y88. Identlficatton

based on 10 balsam mounts

Since 8. parvvm has priority in Jehnstun’s

{1917} paper, we haye anid the present

Species, B. parvum,

At first examination it appears thae aceraibu-

lum shape might distinguish specimens from

Petrochelidan and Corvus, Johnston described

the suckers of B. parvien as “rather longer than

broad". In a slide in T. H, Johnston's collection

labelled "“Lyperosomanm Iparvam, Strepere

versicolar’, thought to be ane of S. J. John-

ston's preparations, the acctabulum measures

200 yam x 282 yom and shows, om one side, a

lateral projection which is 2 feature im some

of the species described here.

In cach of four specimens from Perracheli-

don in which hoth dimensions can be measured

the acetabulum is distinctly wider than lone.

and comes to a point on each side (Pig. 4).

In 6 of 7 specimens from Cervus melori the

ucetabulum is longer than broad; in the seventh,

length and breadth ure equal: in an eighth,

mounted partly laterally, there are indications

of the lateral projections. We conclude that

acetabular shape is variable, @epencing perhaps

on the muscular contraction of the organ, and

also on the mounting of the specimen.

The type of B. parvum shows papillae on

the surface of the bedy, which were not men-

tioned by Johnston. They are pointed or

S17, trom

DICROCOELIIDS FROM AUSTRALIAN HOSTS

(mostly) rounded; about 13 jm across the base

umd 8 ym above the surfuce of the body, They

May be present in the acetabular repion, but

oceur only very occasionally anterior to tt,

Posterior to the acetabulum they are about 25

um apart, in the vitelline region about 53 um

apart, diminishing to the posterior end.

Papiliac similar in size and shape, and also

limited ta the acetabular and post-acetabular

Tegion, are present on the types ef & niegayte-

mum jind B, harrison’, They are present in

some specimens from Petrachelidon nigricans,

but not in the specimens from Cereus mellori,

which {ire not so well preserved.

Petrechelidan nigricans, alhough migratory,

breeds in Australia und ts found in winter only

io New Cluines und the sourhwest Pacific, from

whith areas Braclivlecitiah spp. have not been

recorded, Corvus mello’ is restricted to Aus-

tralia.

Keachylecithum insulare o.sp.

FIGS 1-2

Flosis, Ratius fuscipes (lype host), Amphiboluris

fionnt

Levation in host. Gall bladder and bile duets.

Loaculity, Pearson Island, S. Aust,

incidence, lof 4 rats (January 1969), 2 of 9 rats

(Febrnary 1973), many trematodes in each in-

fected rat 2 of 1] lizards (Junuary 1969)-

flolatvpe. SAM V7&4,

“aratypes SMM V77, V7B, V79.

Other slide deposited. SAM V98.

Other host. Turnix castanota. Cowell, S. Aust.,

26.v.1965. In 1 of 2 birds collected together.

(One complete; one complete except for first

testis and anterior end; and four pieces). From

liver. Food of host includes “heavy” insects.

Description based on whole mounts of 5

complete and 3 almost complete specimens,

with 17 pieces of several other specimens from

the type host.

Description

Body (4.0-5.4 mm) very long, narrow,

cylindrical or sub-cylindrical. Width/length

ratio 1:85-1:115 (1:102). Papillae not seen on

surface of body. Acetabulum (131 x 129 ym),

situated approximately in anterior twelfth of

body, a little larger than oral sucker (103 x 89

pm); not strongly muscular, wider than body

and approximately diamond-shaped; in lateral

mounts margin folded irregularly. Oral sucker

more muscular than acetabulum. Pharynx not

clearly seen in most specimens; oesophagus and

alimentary caeca not seen. (In specimen from

Turnix castanota, pharynx obvious and part of

oesophagus seen).’

Testes elongate oval; situated well posterior

to acetabulum; tandem. Cirrus pouch mostly

anterior to acetabulum; cirrus with bluntly

rounded end; genital pore nearer to acetabulum

than to oral sucker. Ovary oval, smaller than

testes. Ovary and testes more or less well

separated from each other, probably depending

on state of elongation of worm, by uterine coils.

Vitellaria, beginning near posterior border of

ovary, consisting of variously sized groups of

small follicles; largely obscured by eggs; up to

800 um in extent. Uterus, with many eggs,

occupying all of hindbody not filled by gonads

and vitellaria.

S. porrectum (Braun) from Halcyon sauro-

phaga, New Guinea, and S. elongatum Yadav

from Sterna aurantia, India, are the only

species previously described in the genus. S.

mawsoni shows a close resemblance to S. por-

126

rectum in general appearance, though it is

much smaller, Travassos (1944) Who repro-

duced Braun’s (1902) measurements and

figures, referred to the great elongation of the

body of S. parrectum, und said that careful

examination of Braun's figures gave a clear

impression of material preserved in the carly

stages of decomposition, The widening of the

body shown in the anterior third is almost cer-

tainly distorted. The length was 17 mm; maar

mum width was given as 0,57 mm (width/

length ratio 1:30), width at posterior end 0.2

mm (1:85). From Travassos’ fig, 3 we esti-

mute the probable maximum width as 0,28 mm

{ratio 1:60).

S. mawsonl differs from §, porrectum in:

size (4-5 mm, 17 mm respectively), width/

length ratio (1:85-115, 1:60); acetabulum not

so near anterior end (forebody/total length

ratio 1:12, 1:22); ovary (oval, spherical);

(estes further forward (in anterior third, middle

third); eggs smaller (32 x 19 «wm, 37-41 x 23

m),

" We have not seen the description of S. elan-

ert iderl.

Skrjabinosomum pomatostomi n.sp.

FIG. 10

Host, Pamaiosiomus superciliosus,

Location if host, Liver.

Locality. The Bunkers, Flinders Ranges, S. Aust,

22.vii, 1965,

Incidence. | of 35 birds, trom S, Aust. and N.T,,

1938-1969 (10 of which were from the Bunkers,

July. 1965).

Food of host includes beetles and other insects.

Holotype. SAM V99, with six pieces of paratype

on same slide, Two other slides, one with

two twisted specimens and several pieces of no

value (paratype SAM V100) the other with only

anterior end (paratype SAM V101).

One entire trematode and a number of pieces

of at least three others were recovered, All

specimens were rather sticky. SAM VIOL

(anterior end only) was studied first in

glycerine. Thus measurements of the suckers

from the ventral and Jateral aspect were taken.

Description based on all available material.

Deseription

Body elongate (2.4 mm), narrowly cylin

drical or sub-cylindrical, with pre-oral lobe.

Papillae not seen on surface of body. Acetabu,

lum (115 x 181 pm) not strongly muscular,

diamond-shaped and transversely elongate in

ventral view, probably a little larger than oral

sucker (97 x 94 pm), lying between anterior

sixth and seventh of body in holotype. Pharynx

l. MADELINE ANGEL & J. C. PEARSON

present; oesophagus and alimentary caeca nol

seen, Testes oval, situated well back from

acefabulum, in tandem, Testes and ovary

separated from gach other by uterus. Ovary

closer 10 posterior end of Body than to acctabu-

lum, Vitelluria notin well-defined folleles

In §. pomatoston’ the sizes of the organs

and the distance of the acetabulum from the

anterior end of the body, the distance of the

first Jestis from the acetabulum, and the

relationship of testes, ovary and vitellaria to

each other are similar ta these features in 9.

mawsoni, However, the much shorter distance

from the ovary to the hind end af the body in

S. pomatostomi makes the relative positions of

the Organs in body length distinetly different tn

the two species. In the five complete specimens

of S, mawsoni from Manering Hey igula the

rao of lenyth of body postendr to ovary ta

total body length is 10:17. 18, tS, 19 and 19:

in a sixth specimen, the length of which can be

estimated closely (lacking only the oral

sucker), the ratio is 10:20; in the only surm-

plete specinien from Turnix curterita the ratio

is 10:21. In &. penerostoml it is 10:30,

Corresponding with the shorienmg of the

hindbody in S$. poenarestoi the ucetybulurs lies

relatively further back. the vitellaria are shorter

im nearer to the posterior end of Che hoy,

wod the uterus and number af eges are much

reduced,

IL appears thal the acetabulum of 4, peynate-

srommi is larger than that of S. menvson! From

Moanorina flavigula (though the same size as

id S. mawsoni from Turntx cartanota). Howe

ever, the acetabulum in both species is noe

strongly miutscular and rts outline is variable

(probably depending on the position in which

it is mounted). The natural shape of the

acetabulum apoeurs to be angular. with the

margin coming to a point on each side of the

body. When such a specimen cin be measured

in this position its width is maximal. The depth

is probably a better indication of the size of the

acetabulum, The width of the evas (71 x 16

am) is less in §. parrerastomé than in 8. naw

Nout.

It is passible that §. pomatosranl is a young

form of S. muwsent in which the him end of

the body has not reached its Cull length, the

uterus and the vitellaria being mot yet fully

developed. But although the propertiou: of

this part of the body are so markedly dillerent,

the size of the gonads (and presumably their

maturity) is mbeh the same i the worns feo

the two hosts. We regard 9. somarnstami a dis-

DICROCOELIIDS FROM AUSTRALIAN HOSTS 27

finet spevies, characterised by the close

proximity of the ovary to the hind end of the

hody.

§. pomaeastond differs from §, perreetiun in

size (2.4 mm, 17 mm respectively); in the

more backward position of the acetybulum

(forebody/hindbody 1:6—7, 1:22), In having

an oval ovary, situated at 2/3 of body length

(round, at midlength in S. porreeiunr), und in

ege size (31 x 16 wm, 37-41 x 23 pm).

Skrjabinosomum sp,

Hast. Micracca leucaplinea,

Location th host. Liver.

Locality, Port Augusta, S. Aust, September, 1965,

fncidence, 1 of 11 birds from various loculities (4

agen Port Augusta) in S, Aust. and N.T., 1958-

Food of lost includes beetles und other insects,

Slide (with 5 pieces) deposited In University of

Adelaide Helminthological Collection,

Five pieces of a very long, narrow trematode

were stained and mounted on one slide. The

worm Was incomplete, and lacked hind ene ail

oral sucker, The acetabtilim, mounted in

lateral view, is 76 pm high by about 39 jam

deep. The testes are obviously situated well pos-

terior to the acetabulum, Vitellaria show in two

pieces; they consist of small follicles in large

groups, not sharply demarcated, and oecupy 4

leneth of at least 506 am, Ten eges measure

28-33 pm x 13-18 pm (30 14 am),

The worn is similar to the elongate species

S. mawseni and §, pamatastami and may be

conspecific with one of them,

LUTZTREMA Travassos

Generic diagnosis, Dicrocoeliidac, Dicro-

cocliinae. Body elongate or relatively broad,

With pre-oral lobe. Body surface with or with-

out papillae or spines, Acetabulum larger thun

oral sucker or suckers sometimes equal, Sitgle

caecum, or with one of caeca rudimenkiry and

nol extending beyond acetabulum; caecum reli

tively long, median, in form of zigzag, Genital

pore about half-way between suckers. Testes

post-acetubular or almost acetabular, tandem

or diagonal, Ovary post-testicular. Vitelline

follicles large, post-oyarian, Interal, sometimes

converging anteriorly, Uterus occupying most

of hindbody, with coils between ovary und

testes. Parasitic in liver and gall bladder of

birds and mammals.

Lutztrema ailoroedi (1.3p.

FIGS 12-14

Host. Allneoedus crasslrasiele,

Locality. Mt Glorious, Old, 16,1957, 30) 1947

{collected KE. Webber), 29.0, 0962,

“aration in trast. Bile duct and at mouth af gall

bladder.

fnefdenee. Prom 7 to imaty specimens. in all 4

green carbirds examined.

Holatype SAM VIO08,

Puratypes. SAM V109, VI10, VIT1T,

Deseription based on live wornis and on bil-

sam mounts of & Whole worms and 3 anterior

ends (16.1,1957), Measurements of worns

from two collections, 29.x,1962 (2 birds) and

301.1957, with some jdditional information on

these whole mounts.

Description

Body elongate. almost cylindrical; forebody

much narrower, bent back dorsally from pro-

truding acetabulum; in whole mounts width

generally slightly greater at level of vitellaria,

Length 3.30-4.28 mm (3.80), width 0,221-

0.272 mm (0.238) for 5 worms, depth 0.214

mm tor 3 worms. Ratio of width to length

bi) 14-1218. (In 2 hosts collected 29x, 1962, &

imiture trematodes (neasure 2,89-4.18 mm

(3,30), and tm one host collected 30.1.1957, 7

worms aré 2.10-292 mm (2,60), Papillac

scattered over length of body; Most Numerous

ar preodral lobe,

Suckers forming deep cups. Acetabular

larger than oral sucker, in anterior seventh to

anterior fifth of bowys 141-200 um (176) long

® 153-188 pm (172) wide, und (in 4 speci-

mens) 165-200 ym (188) deep. Oral sucker

106-129 pm (114) long x 94-100 ym (96)

wide, and (in 3 specimens) 94 pm deep. Sucker

ratio (using mean diameters of length and

width, in five specimens) 1:1.5-3} 1.7},

Pharynx 39-50 jan (47) © 53-63 jm (57)

and (4 specimens) S5-Al) pm (59) deep.

Caecum dorsal to cirrus sac and to middle or

side Of acetabulum, sinuous dorsal to and

hetween lestes, dorsal to ovary and exerctoty

bladder, ending less than hall way from ovary

to posterior end,

Exeretory pore termipal. Excretory bladder

I- shaped, iv youn specimens reachiig nearty

to oolype; common collecting tubules arising

from anterior end of bladder: at level between

testes grving off antenor and posterior caflect-

jog tubules, Exerctory formula 2[(2 + 2+ 2)

+(2+2+ 2)1.

Testes tandem, round to elongate oval, |53-

235 pm (I9L) long « 129-188 pm (162) wide,

und (in 3 specimens) 141-153 ym (149) deep

Anterior testis separated from acetabulum by

119-306 pm, Testes contiguous or up to 85 ym

ipart, Seminal vesicle coiled, Cirrus sac

128 L, MADELINE ANGEL & J. C. PEARSON

2 5 ;

rt eure

So,

Sos

= O

Figs 11-16. 11, Brachylecithum latins Holotype, dorsal view; 12. Lutztrema ailuroedi Holotype, 13. L.

ailuroedi Sketch, of young specimen; 14. L. ailuroedi Sketch, anterior end showing papillae;

15. Pancreatrema meliphagae Holotype; 16, P. meliphagae Ege. (Pigs 11, 15 to same scale.)

immediately anterior to acetabulum, about 158

x 53 pm. Genital pore about midway between

suckers.

Ovary in anterior third to two-fifths of bady,

submedian, round or transversely oval, 88-118

pm (102) x 112-141 pm (129) and 106-123

nm (118) deep; separated from posterior testis

by 34-136 ;.m. Receptaculum seminis up to 79

x 68 pm, Vitellaria composed of 8-10 large

(up to 121 x 60 ,»m) follicles on each side of

body. Vitelline fields 318-447 wm (376) long.

from 34-136 ym posterior to ovary. Uterus

occupying all of hindbody posterior to vitel-

laria, then passing forwards dorsal ta and

between ovary and posterior testis, posterior

and anterior testes, anterior testis and acetabu-

lum: travelling dorsal to cirrus sac and open-

ing immediately anterior to cirrus at genital

pore. Eggs 32-39 um (35) by 21-23 pm (22);

when new laid hatching in water; miracidium

swimming actively,

The character which distinguishes Lutztrema

trom the closely related genera Brachylecithum

and Lyperesomum is the alimentary system, in

which there is a single caecum, or with one of

the two caeca rudimentary; the main caecum

is relatively Jong, median and in the form of a

zigzag. Probably because the alimentary system

was not described in the original descriptions,

a number of species ascnbed to Lutztrema

have subsequently been referred to other genera

(generally to Brachylecithuin), Yamaguli

(1971) listed 11 species from birds and one

from a mammal; he did not mention the species

£. heterocaruxi Bisseru, L. singhi Bano, EL,

DICROCOELIIDS FROM AUSTRALIAN HOSTS

stinkaral Dane or L, vitellacanfluentin Ali,

Deshmukh & Karyakatte, No further species

have been described to date. Most of the ahove

16 species have relatively broad bodies, short

behind the vilellarta, and L, afluveedi differs

from them in this and in other features. tts

hody width/length ratio is closest to that of f,

microstonnun Denton & Byrd [from Cyene-

eine erigrata Linn.), L. sinehi Bune (from Car-

wus splendens (Kereillot)) and L, hererpeeraxt

Hisseru (irom Helervcerax capensis capenits

Sharpe). From the ranges of width and length

wiven for the first two species, we estimate the

with’ length ratia of LL. ynicrestommun to vary

from I> Li-l:16, and of L. singii from 1°10-

1:13, ‘The measurements given of one speci-

mew of L, heterecorax’ (presumably the hole-

type) show a ratio of 1:16,

The most notable difference between L.

ailuroed? and L., thicrostonmun is in the posimon

ef the aeectabulum {situucd “about one-tenth of

body length from anterior end” in L, iniera-

‘Tomi, and between the anterior seventh anal

Hfih in &. allveoedt). Other differences are: the

body behind the vitellariy fas shown in the

figure) ig relatively longer in Ly wiierusromun)

in 4. ailuveedi, the testes are round or elongate

oval anid jie in tandem, while m L. niterayre-

iin they ate rounded to transversely oval and

situated shghtly obliquely; in L eierasteniem

the rauia of the diameter of oral sueker to

acetabulum is 1)1.75-1:2.80, but in LZ,

ailureedi, 171,5-1:1,7, The eges of L, micro-

stoma may be slightly shorter (29-35 jum

long, 19-24 ,»m wide). Laurer's canal was not

observed in L, aluroedi, but with these excep-

tions, this species agrees very closely with the

description of £. jilerastonuin,

Apart trom tts relatively narrower body. £.

ullureedé iffers from L, singhi in having

scattered papillae on body surface (L. singhi

is covered with minute spines); egay (32-39

wm) are slightly longer (30-36 »pm in Lb.

simeh’) > vitcllana ending <omewhat nearer hind

end of body; tesies in tundem fsomewhwt

oblique in £. simi), Bane gives the position

of the acetabulum as “posteriar region of

uoterior third of bowy”. but from the figare of

the type, it is im the anterior fifth (in be

wilvrecdi, antenar seventh to Atth),

£, adiluroedi appeats to be close to ZL,

heterevorisi, bul dilfers chiefly in the sucker

rateo [be 05-11 6.7 an former, 1:2 in latter) and

ity eee sive (35 % J2 pm in former, 39 x 26 wm

in latter), The absolute sizes of the suckers

appear tu be greater in L, frelerecerani, the

128

acetabulum is slightly further back (in anterior

fifth of body in &. heterocoraxi, between

anterior seventh and fifth in L. alureed!) and

the maximum width of the body is at the

acetabulum in &. heteraceraxi; at the vitellaria

in L. wlarvee.

PANCREATREMA Oshmarin

Generic diagnosis. Dicrocoeliidge, Body

flattened fusiform, with pre-oral lobe. Oral

sucker moderately Jarge, acetabulum lacking,

Pharynx, short oesophagus, two cueca, Testes

small, rounded, symmetrical, pre-equatorial;

venital pore post-pharyngeal or post-bifurcal,

Ovary round, submedian, in equatorial region.

Vitellaria forming rather compact clusters on

each side. close to posterior border of ovary,

Uterus extensive Hut not in the extremities of

body-

Parasite in bile or pancreatic ducts of birds,

Pancreatrema meliphagae n.sp.

FIGS 15-16

Hlosl. Meliphaga ornata,

Locality. Blanchetown, S. Aust. April, 1965.

Location ti hast, “Probably from bile duct",

Jacidence. | specimen trom | of & birds.

Holotype, SAM V 112.

Deseription af holarnipe.

Flatlened fusiform worm, 4.35 x 0.85 mm,

Greatest width just anterior to testes. Oral

sucker 400 x 388 wm; pharynx 118 x 118 am;

oesophagus short; two caeca, apparently ter

ininating near posterior end of body. Acetabu-

lum not seen, Testes entire, pre-equatorial, sym-

metrical, partly obscured by uterus; right testis

about 176 x 200 jm, left testis about 165 x 188

pm, Ovary round, entire, 235 pm in diameter,

submedian, just pre-equatorial. Genital pore

apparently nearer to pharynx than to caecal

bifurcation, Vilellaria in compact clusters; on

left side reaching mid-ovarian level, contiguous

with ovary, on right side post-ovarian; left

cluster 376 x 282 ,»m; right cluster 470 x 400

jon. Uterus extensive, but not obscuring most

of organs. Eges (Fig. 16) plump, somewhat

flattened at opercular end with slight shoulders

at rim of opercultim, shghtly pointed at other

end; 44-46 pm» 19-22 pm (average of ten, 34

x21 yim).

In another specimen of Meliphagu arnuta

(from Cowell, S, Aust, May, 1965), about 10

trematodes im very poor condition were

recovered fram the bile duct of liver, It is not

possihle ta identify these specimens definitely,

but i is thought likely thut they belong to Pun-

creatremed. They appear to be similar to P.

130

meliphayae in size and shape, and ihe eggs

have the characteristic appearance of those of

P. weliphagae, though they are slightly larger

(35-42 ym long x 22-26 wm wide (40 x 24

ym) >. Six oral suckers, taking stain only

faintly, measure |76-2/2 pm long x 176-235

pm wide (193 x 207 pm), and one pharynx

measures 59 x 59 ,m.

Pancreatrema disacetabuluum Oshimarin from

Eurystomus orientalis from Russia, including

the Primorskii Krai, is the only previously

described species, but Lalitha & Alwar (1980)

recofded Pancreatrema sp. from Anas boschas

dorriesticus in Madras.

P, meliphagae differs [ram P- dixacetabulunt

in ats more slender shape (width/length ratio

1:5 in former, 1-3 in latter) in the slightly

more forward position of the ovary, in the

apparently more forward position of the genital

pore, and in the size of the eggs (34 % 21 jit

in PL neliphagae, 62% 42 pm in P, disacerahu-

lw). The ezes, although more pointed at one

end in P, meliphagae, ave the same general

appearance in the two species. Measurements

of the organs are much the same, except thal

the largest vitelline field in P. disceerabuluir is

almost twice the length of that in P. meli-

phagae, P. disacetabulum was found in the pan-

creas of Eurvstomuy orientalisy P. meliphagac

was thought to be from the bile dliict of its

host

Proacetahulorchis dogieli Belopulskaja &

Bychovskaja-Pavlovskaja

Host, Ardea novuchollandiae,

Localities. Brisbane, Qld, tix.L965 (21 speci

mens). Deception Bay, Qld, VS5.vili,19h1 et

specimen; colletled G. Monteiiir),

Eocatlen Ix fiest. Bile ducts.

fncidence. 2 of 2 birds from Qld, none of 14 birds

from S. Aust.

Slides deposited. SAM V113, VI14.

Description based on 9 stained mounts and

% specimens in glycerine. (Measurements fram

stained mounts). The specimen from Decep-

tion Bay is much farger than the others, and

measurements for it ure given separately,

Deseription.

Bedy flat, elongate; width generally uniform,

slightly greater behind acctabylum, gencrally

behind mid-body, Length 3,4-4.2 mm (3.8),

width 544-816 am (646), Width/length ratic

1:4.8-1;6.9 On body surface, papillac, 18 ,,m

across base, as chose as 26 mM apart in preace-

tabular region, much sparser posterioely (noe

Wpparent in all specimens). Suckers rounded,

approximately equal, Oral sucker 329-435 am

L, MADELINE ANGEL & J, C. PEARSON

(366) long & 294-388 gm (341) wide,

Acetabulum, in anterior third of body, 318—-

412 um (355) in diameter, Pharynx rounded,

94-112 wm (99) long x 88-118 pm (101)

wide. Ocsophagus, slightly contracted, 106-223

wm (141) long, Caeca extending to distanee

S10+816 jam (595) from end of body.

Excretory pore terminal; main stem of

bladder bifurcating just posterior to ovary;

excretory arms extending to leVel of pharynx.

Testes 94-212 pm (169) long x 118-247

um (147) wide: contiguous, generully diagonal,

(In 18 specimens, testes of 11 diagonal, 2

approaching tandem, 4 approaching symmetri-

cal, 1 symmetrical), Anterior testis on right or

left side; posterior testis slightly dorsal to

antenor rim of acetabulum or slightly anterior

to latter, Seminal vesicle internal: cirrus in one

specimen 176 x 59 wm, with flattened end.

Genilal pore mn region of caecal hifurcation,

generally very slightly posterior to latter.

Ovary rounded, 71-153 wm (98) x 82-165

pm (109), median, near mid-length of body.

Mchlis’ gland somewhat diffuse, posterior and

partly lateral to ovary. Receptaculum semines

large, rolinded (in four specimens, 165-176 pm

(174) x 141-164 (154)), abutting posterior

lateral border of ovary,

Vitelline fields mostly extracaecal, from 680—-

1426 wm (1075) long, sometimes reaching

acetabular level anteriorly; posteriorly reaching,

approximately, posterior third of bady. Follicles

up to 82 x 33 ym, mostly in single or double

row on each side.

Uierus intercaccal; posterior to caeca spread-

ity oul to sides and end of body. Ascending

wlerus passing dorsal to acetabulum and to

testes, to genital pore. Eggs 29-32 ,an (30) »

18-22 wim (20),

Specimen frorr Deception Bay: Length 7,2

mm, width 0.95 mm, width/length ratio 1:7-5.

No papillac on surface. Oral sucker 447 x 494

pm, acetabulum 329 x 529 ,,m_ Pharyms 141 x

12% mm, oesophagus 318 pm; cacea ending 1.19

ma from posterior end. Testes diagonal;

anterior testis (on left), 517 x 270 pm: pos

terior, 435 x 282 jam, Cirrus 247 x 59 pm,

Ovary 188 x L76 pm. Vitelline folds 2.18 and

2.07 mm, reaching ta 221 mm from posterior

end of body. Eggs 28-31 am (30) x L7-20 pm

(18).

Discussion

Three species bave been assigned to the

penus: PF prashkadi Gogate, P. dogicli Bolopol-

skaja & Bychovskaja-Paviovskaja and Jt

strigosts Sudarikoy & Pavlov but only dogiell

DICROCOELUDS FROM AUSTRALIAN HOSTS wt

aprees with Australian specimens in a sucker

ratio approximately 1) 1.

We have been unable to consult Belapolskaja

and Hychovskaja-Paviovskaja’s description of

Proacetabloreiis dogieli. Skrjabin (1970)

{who followed Bychovskaja-Pavlovskaja

(1962) in including the species in Platyno-

trema Niooll) gave a figure, presumably from

Helopolskaja and Bychovskaja-Pavlovskaja

(1954). and a description of the species

according to Belopolskaja (1954). We have

corpared our specimens with a further descrip-

tion by Bychovskaja-Paviovskaja (1954) and

with the figure and description in Skrjabin

(1970), and conclude that they are conspecific,

(Alfhough m the figures given by these

suthors the vitelline ficlds appear to he shorter

than in Our specimens, io Lhe measurements

given in Skrjabin (1970) they are comparable

with ours). Belopolskaja and Bychovskaja-Pay-

lovskaja’s original specimens were from

Numeniva avqidties and N. madagetvarienvis

from Siberia, Fischthal & Kuntz (1974) gave a

description of P. dagieli from the mangrove

heron, Butorides striatus, from Sabah, with a

discussion of the species. They noted variation

in the relative positions of the testes, which

Were somewhat diagonal in most of their speci-

mens bul were frequently symmetrical. As metfi-

tioned in our description, the testes in worms

from one individual host varied from symmetri-

cal to ulmost tandem,

Acknowledgments

We wish to thank our colleague Patricia M.-

Mawson for providing us with the first speci-

mens of Brachylecithum insulare, Me P, E

Hornsby (Department of Psycholagy, Univer-

sity of Adelaide) for bringing us rats from

Pearson Island, Mr, C. M, Weaver (National

Parks & Wildlife Service, Queensland) for trap-

pug Water rats in Cairns, and Dr P. Hutchings

(Australian Museum) for sending us type shides

of Brachylecithum spp.

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A NEW SPECIES OF LARGE, GREEN TREE FROG FROM NORTHERN

WESTERN AUSTRALIA

BY M. J. TYLER, MARGARET DAVIES AND A. A. MARTIN

Summary

A new species of large, green tree frog of the genus Litoria is described from the vicinity of

Kununurra in northern Western Australia. Details of external morphology are supplemented by a

study of radiographic plates. The new species is most closely related to L. caerulea and lives

sympatrically with it.

A NEW SPECIES OF LARGE, GREEN TREE FROG FROM NORTHERN

WESTERN AUSTRALIA

by M. J. Tyter,* Marcarer Davins* and A, A. Martint

Summary

Vyter, M. J., Davins, MArGarey & Martin, A. A, (1977) A new species of large, green tree

frog from northern Western Australia. Prats. R. Soc. S$. Aust. WO1(S), 133-138, 31 August

1977,

A new species of lurge, green tree frog of the genus Literia is described from the vicitity

of Kufuourrs in northern Western Australia, Details of external morphology are supple-

mented by a study of radiographic plates. The new species is most closely related to L. caerulea

und lives sympatrically with it,

Introduction

The green hylid tree frogs of Australia range

in size from the predominantly Papuan Literia

infrafrenata (Giinther). attaining a maximum

size of 135 mm, to small and delicate species

such as L, fallax (Pelers) and ZL. hiceler

(Gray), adult as small as 23 mm. The con-

spicuous nature of these green animals renders

them particularly highly disposed to being col-

lected by herpetologists, leading to the reuson-

able assumption that this group of Australian

frogs should be well established and txono-

mically stable,

In the course of a visit to northern Western

Australia in February 1977, we collected rep-

resentutives of an hitherto undescribed green

hylid, readily distinguished by several features

of its gross morphology, but with distinet

affinities with L. ederulea. Subsequently addi-

tional preserved specimens were provided by

the Western Australian Museum (WAM).

Here we describe this new species, compare it

with L. caertled and discuss the significance

of this discovery,

Methods of measurements follow Tyler

(1968). X-ray illustrations were obtained by

the Rank Xerography process employing posi-

live mode, Direct comparisons were made

between X-ray prints of the holotype of the

new species and X-ray prints and dried osteo-

logical material of L, caerulea.

Litoria splendida sp. nov,

FIGS 1, 3, 4

Holotype: WAM R56840, An adult female col-

lected at Lake Argyle Tourist Villaze, Kimber-

ley Division, northern Western Australia, by a

joint University of Adelaide and University of

Melbourne ficld party on 22.ii,1977,

Definition: The characteristic features of this

Species are its very large size (males and

females ca 100 mm S-—V length); possession

of a vast and bulbous gland on the entire dor-

sal surface of the head; relatively short and

sightly webbed fingers with prominent ter-

minal discs; brilliant green dorsal colouration

bearing sulphur spots, and its orange flanks

(Fig. 1).

Description of Helotype: Head considerably

hroader than long (HL/HW 0,85), its length

equivalent to one-third of body length (HL/S—

V 0.33). Entire dorsal cranial surface obscured

by a dermal gland raised approximately 5 mm

ubove surface, Snout not prominent, truncated

when viewed from above und in profile, Nos-

trils more lateral than superior; their distance

from end of snout considerably less than that

from eye. Distance between eye and naris

greater than internarial span (E—-N/IN 1.18).

Canthus rostralis slightly defined and gently

rounded, Eye relatively inconspicuous, its

diameter less than eye to naris distance. Tym-

* Department of Zoology, University of Adelaide, North Tce. Adelaide, S.A. 5000.

* Department of Zoology, University of Melbourne,

134

panum visible and inclined medially at 45°;

diameter equivalent to eye diameter.

Vomerine teeth on short, posteriorly

directed, triangular elevations between and pos-

terior to chonnae, Tongue extremely broad.

Fingers short and equipped with broad

Jateral fringes: in decreasing order of length

3>4>2>1, Webbing between fingers only

basal. Terminal discs extremely broad and

extending considerably beyond margin of

lateral fringes, Subarticular and palmar

tubercles prominent,

Hind Jimbs rather short (TL/S-V 0.48),

Toes in decreasing order of length 4>5 =

3>2>1, Webbing reaches half way up penulti-

mate phalanx on 5 and base of penultimate

phalanx of 4, Subarticular tubercles prominest.

Very large oval inner and small rounded outer

metatarsal tubercles.

In addition to cranial gland, skin modified

in following ways: (1) poorly developed der-

mal ridge ou posterior margin of forearm; (2)

ventral surfaces of body snd femora unifornily

granular,

In preservative dorsal surface of head, body

and portions of limbs hidden at rest in a living

frog are dull slate blue. Dorsum of body

liberally spotted with very small white areas,

Dorsal surface of third and fourth fingers, and

third, fourth and fifth toes, anterior and pos-

terior surfaces of thighs and almost entire ven-

tral surface very pale cream,

In fife dorsum dark lime green bearing

numerous small sulphur yellow spots, cach of

which has a dark border. First two fingers, first

three toes, groin and back of thighs brilliant

orange. Ventrally white, with pale green mandi-

bular border.

This female specimen has a few very small

plemented ova and the oviducts are convoluted

only posteriorly,

Dimensions of Holotype: S-V 100.7 mm: TL

48.0 mm; HL 33.4 mm; HW 39.2 mm; E-N

8.6 mm: IN 7,3 mm; EB 7.7 mm; T 7.8 mm.

Variation

There are six paratypes all taken in northern

W.A.. WAM 26818, Old Napier Downs

Cave, Napier Downs, Napier Range, A. M.

Douglas and G, W. Kendrick 8,vil.66; Ra4601—

O02, spillway at northern boundary of Lake

Arityle, Western Australian Museum survey

LLii72 and 203,72; R47231, Prince Regent

River National Park, Western Australian

Museum survey 17.vn.74; R56779, Kimbolton

M, I. TYLER, MARGARET DAVIES & A. A. MARTIN

Spring, Kimbolton Sin, W, H- Butler 26-vi.76;

R56780 Drysdale River National Park,

Westeru Australian Museum survey | 3.vili.76,

The snout to vent fenygth ranpe is 9YO.2—

106.3 mm; the largest of these is a fernale con-

taining numerous small pigmented eva.

Ih their gross morphology the paratypes dit-

fer from the holotype only slightly, All extibit

an extremely prominent supracranial gland.

sOmetimes extending anteriorly beyond the

anterior tip of the snout, and also overlapping

the tympanum. The head is consistently

broader than long (HL/HW 0.85-0,95), whilst

the HL/S-V ratio is 0,32-0.37. Similurly the

E-N/IN range is comparable to that of the

fiolotype (€1.00-1,23). The limbs are eon

sistently rather short (TL/S-V 0.46-0.50).

All specimens exhibit qumerons, small, white

spots on the dorsum,

Comparison with other species

(a) External murpholegy

Within the Australo-Papuan region there are

several large tree frogs that are predominantly

greew in life. Of the species restricted to New

Guinea, f. splendida may be distinguished

from £. graminea by its slightly broader head

(HL) HW 0.85-0.95 in L. splendida; 0.9061 At

in L. graminea), shorter hind limbs (TL/SV

0.46-0,50 in L. splerdida: 0.56-0,60 in b,

graminea), less cxtensively webbed fingers

(approximately one-half webbed in L. splen-

dida but reaching the discs of all fingers in £,

graminga); L. geaminea lacks the supracranial

gland of L. splendida and is probably smaller

(males of L. gremittea are only up to 65 mm

in length).

Litoria infrafreneta is a predominandy

Papuan species known in Australia only from

the Cape York Peninsula in Queensland, Its

size overlaps that of L. spletadidy, but it is a

shallower bodied creature, always exhibiting a

brilliant white mandibular stripe. The head is

structurally quite different. tending to be

flattened with a prominent and concave [jcil

shelf, In contrast, £. cplendide has a high heal

and the premipent supracranial gland is totally

lacking in the former species, The hind limbs

of L, splendida are sthrhtly shorter, TL/S—V

0,460.50 as opposed to 0,57-1L63 im L. infra

frenata.

Litoria splendida is most closely related to

L. caerulea (Pig. 2) and occurs sympatrically

with it, The size and general habitus of the two

species are similar, They differ principally in

NEW LARGE, GREEN TREE FROG

Fig. 1. Adult individual of Litoria splendida.

Fig. 2. Adult individual of Litoria caerulea.

136

Fig. 3. Distribution of Litoria splendida (solid

circles) and the Australian portion of the

geographic range of JL. caerulea

(stippled).

colouration, dermal gland development and in

behaviour (see discussion). L. caerulea occu-

pies a large area of the Australian continent

(Fig. 3) and is somewhat variable in its

colouration throughout its range. Central Aus-

tralian individuals commonly have large white

spots and patches on their bodies and are

referred to L. gilleni by some authors (e.g.

Cogger 1975). In the northern part of the

Northern Territory and Western Australia (in-

cluding the type locality of L. splendida), indi-

viduals are usually a pale yellowish green; else-

where the dorsum ranges from dark green to

olive. However, the distinctions between L.

splendida and L, caerulea remain conspicuous.

L. caerulea lacks the brilliant yellow markings

found on the thigh, groin and hand of L. splen-

dida and lacks the small sulphur spots com-

monly occurring in the new species, L. caerulea

is otherwise unique amongst Australian hylids

in exhibiting a parotoid gland but it lacks any

comparable glandular tissue on the dorsum of

the skull.

Osteological features of the two species are

listed below.

(b) Comparative osteology of L. splendida

and L. caerulea

Recognising that a close relationship exists

between L. splendida and L. caerulea we under-

took a detailed comparison of a number of

osteological features: L. splendida is shown in

Figure 4.

M. J. TYLER, MARGARET DAVIES & A. A. MARTIN

1. Skull

(a) Shape: the shape of the skull of L. splen-

dida differs from that of of L, caerulea in that

it is narrower and less blunt.

(b) Nasals: the shape of the nasals and their

relationship with the sphenethmoid are difficult

to determine from X-ray photography. It is

known that in L. caerulea the well-developed

nasals are narrowly separated medially by the

sphenethmoid with which they articulate and

overlap extensively (Tyler & Davies!).

parietal flange is present in both species, simi-

lar to that found in L. infrafrenata (Davies in

press) but in L. caerulea this is developed to a

lesser extent than in L. splendida and L. infra-

frenata.

(d) Frontoparietal fontanelle: this is

moderately sized and ovoid in both L. splen-

dida and L. caerulea.

(e) Squamosals: the otic rami of the squa-

mosals do not overlap the crista parotica in

either species.

(f) Otoccipital region: the lateral extremities

of this region are cartilaginous in both species.

(g) Pterygoid: the well developed pterygoids

do not articulate with the otic capsule in either

species.

(h) Quadratojugal: the quadratojugal is well

developed in both L. splendida and L, caerulea.

(i) Prevomers: the alae of the prevomers are

developed to differing degrees in L. splendida

and L. caerulea. Those of L. splendida are

elongate and extend almost to the level of the

palatal shelf of the maxillaries, whereas those

of L. caerulea are short and barely reach the

level of the extremities of the supraorbital

frontoparietal flange.

(j) Palatine processes of premaxillaries: in L.

caerulea, these processes abut along their

medial extremities, whilst in L. splendida there

is no medial articulation.

2. Vertebral column

(a) Sacral diapophyses: the sacral diapophyses

are poorly expanded in L. splendida and

moderately expanded in L. caerulea.

(b) Mia: the ilia extend anteriorly to the sacral

diapophyses in both species.

vertebrae: in L. splendida these are equal and

1 Tyler, M. J. & Davies, M. (in manuscript) Species group within the Australopapuan hylid frog genus

Litoria Tschudi.

NEW LARGE, GREEN TREE FROG 137

alary process

of prevomer

medial ramus

of pterygoid

presacral

vertebra Ill

palatine process

of premaxilla

supraorbital

S frontoparietal flange

> ast

AVS

Ve/)\ otoccipital

td region _ lateral

TA ee extremity

posterior

presacral vertebra

Fig. 4. X-ray print of Litoria splendida (Holotype: WAM R56640).

directed slightly anteriorly, whereas in L.

caerulea the processes are subequal and are

directed anteriorly at a slightly more acute

angle.

(d) Transverse processes of presacral verte-

brae III: these are equal in width to the sacral

diapophyses in L. caerulea, but narrower than

the width of the sacral diapophyses in L. splen-

dida.

Habitat

The specimens collected by us were taken

in artificial situations such as upon bitumen

roads at night following torrential rains. How-

ever it is clear that such a large frog would

require a summer retreat to avoid very high

temperatures. Two of the paratypes were col-

lected within caves: R26818 at Old Napier

Downs Cave, and R44600 at a site 8 km N of

Lake Argyle. Similarly R27231 was taken in

a ledge in a sandstone gorge.

Litoria splendida is evidently adept at sur-

viving in areas of low rainfall, so paralleling

the habits of L. caerulea. In the vicinity of the

type locality the vegetation is predominantly

scattered scrub of sparse, low-growing

eucalypts with pockets of Pandanus. Soils

range there from sand to clay. The area

receives an average rainfall of 50-75 mm from

138 M. J. TYLER, MARGARET DAVIES & A. A. MARTIN

monsoonal rains between December and

March.

Litoria splendida is commensal with man

and the species is well known to inhabitants at

Lake Argyle and at Kununurra. In common

with L. caerulea it inhabits bathrooms, toilets

and other such sites where there is water. The

holotype was taken at night on the floor of an

ablution block.

Distribution

As demonstrated in Figure 3 all of the locali-

ties at which this species has been taken occur

in the extreme northwestern and peripheral

portion of Australia,

Discussion

As indicated above, the new species is phylo-

genetically very closely related to L. caerulea.

The supracranial gland seems to represent

hypertrophied parotoid glands that have in-

creased in extent as well as mass, thus becom-

ing a single unified gland.

In captivity, behavioural differences between

the species have been noted, Whereas L. cue-

rulea tends to become a rather sedentary

creature in captivity, L. splendida after three

months is exceptionally active and avoids

attempts to handle it.

In finding such a large and spectacular

creature in 1977, we are led to the conclusion

that the extent of sampling of the Australian

lower vertebrate fauna is highly deficient.

Acknowledgments

We are greatly indebted to Fish Farms Inter-

national, and in particular to Mr K. Light-

burn and Mr J. Caratty of that company, for

their generous assistance with our visit to

Kununurra. Dr J. E, Bishop of the Department

of Zoology, University of Adelaide, helped us

greatly in the field. Ansett Airlines of Aus-

tralia provided in-transit facilities and trans-

ported live animals back to Adelaide.

The colour photographic plates are the work

of Mr Bohdan Stankovich-Janusch; prepara-

tion and the cost of preparation was borne by

Mercury Walch, Hobart. For the X-rays we

are indebted to Dr R. MacDonald and Mrs A.

Lucas.

A. A. Martin was supported by the Ian

Potter Foundation, and M. Davies and M. J,

Tyler by the Australian Research Grants Com-

mittee.

References

Coccer, H. G, (1975) “Reptiles and Amphibians

of Australia” (Reed: Sydney).

Davies, M. (in press) Cranial osteology of the

Australopapuan hylid frog Litoria infra-

frenata. Ree. 8S, Aust. Mus. 17(22).

Tycer, M. J. (1968) Papuan hylid frogs of the

genus Hyla. Zool. Verh, Leiden 96, 1-203.

ON THE MANUS AND PES OF THYLACOLEO CARNIFEX OWEN

(MARSUPIALIA)

BY R. T. WELLS AND B. NICHOL

Summary

An articulated left and right manus and a partially disarticulated left pes of the cave lion Thylacoleo

carnifex were recovered recently from a Pleistocene cave deposit at Naracoorte, South Australia.

The structure of the manus and pes is described. The manus is digitigrade with limited flexion in

digits IJ, III. IV and V. Digit I is extremely robust, bears a large hooded ungual crest and is

pseudopposable to a spatulate pisiform. The pes is plantigrade with major part of the weight

transferred through the astragalus to the calcaneum. Analysis of the structure of the pes indicates

syndactyly and the presence of a divergent hallux. The manus and pes show structural affinities

with the arboreal phalangerids.

ON THE MANUS AND PES OF THYLACOLEQ CARNIFEX OWEN

{MARSUPIALIA)

by R. T. Wects* and B. NicHoL*

Summary

Wrins, Ro T, & NicHot, B., (1977) On the munus and pes of Thylucolen carnifex Owen (Mar-

supialia). Truas, R. Soc, 8S, Aust, 101(§), 139-146, 31 August, 1977,

An articulated left and right manus and a partially disarticulated left pes of (he cave lion

Thylacolea carnifex were recovered recently from a Pleistocene cave deposit at Naracoorte,

South Austrulia. The structure of the manus and pes is Wescribed. The manus is digitigrade

with limited flexion in digits 1, (11, 1V and V. Digit I is extremely robust, bears o large hooded

wngual crest and is pseudopposable to a spatulate pisiform, The pes is plantigrade with major

part of the weight transferred through the astragalus to the calcuneum. Analysis of the structure

of the pes indicates syndactyly and the presence of a divergent hallux. The manus and pes

show structural affinities with the arboreal phalangerids.

Introduction

Thylacoleo carnifex Owen (1859), the

“Nesh eating marsupial lion", was described

from skull fragments from a late Pleistocene

deposit at Lake Colangulac, Victoria, Although

the tooth formula of such thylacoleonids is

typically phalangeroid, the animals are charac-

terised by development of exceedingly large

sectorial upper and lower third premolars,

large conical upper and lower first incisors and

u marked reduction in the remaining incisors,

canines and molars. The niche occupied by T,

carnifex therefore has heen the subject of con-

siderable speculation (vide Gill 1954; Finch

1971). However recent studies of jaw

mechanics (Finch 1971) tend to support

Owen's conjecture that it was a large carni-

vore.

Finch (1971) made a preliminary analysis

of the skeleton and suggested that the relatively

long fore-limb muy be used to strike al prey

io a fashion similar to that of Sarcephilius, This

suggestion relied partly upon her interpretation

of the paw as a strong and heayy structure in

which the digits could not be widely separated,

However the structure of the pes was unknown

to her, as the major portions of the hind feet

ure missing from all the skeletal material pre-

viously reported. The following is a pre-

liminary description of the general morphology

and arrangement of the bones of the manus

and pes of this enigmatic animal.

Materials and methods

During excavations at Victoria Cave, Nara-

coorte, numerous skeletal elements of Thy/a-

coleo were collected including the articulated

right and left manus with portions of the fore-

limb in association. The specimens were held

together by a patina of calcite, In the labora-

tory the left manus Was partially cleaned, bul

left in its fused state; the right manus was cure-

fully disassembled, cach element cleaned and

then reassembled (Fig. 1). In the course of

preparation of the right manus the cuneiform

and pisiform of the left manus were recovered

from the sediment.

All specimens removed from the Victoria

Cave deposit are allotted numbers indicating

their positions in a three-dimensional grid

(Wells 1975). During the sorting of bone

material, portions of a disarticulated left pes

bearing similar grid reference numbers were

recovered. This partial pes (Fig. 3) is not

referable to any extant marsupial, nor could it

be attributed to any extinct form for which. the

foot structure is known. The caleaneum is con-

sistent with one recovered with skeletons from

James Quarry, Naracoorte (M, Plane, pers.

comm.). The close proximity of Thylaceleo

‘ School of Biological Sciences. Flinders University of South Australia, Bedford Park, S. Aust. 5042.

140)

P 16679

Fig. 1, Assembled right manus (PL6679) of T-

carnifex, The Sth medial phalange, cune-

iform and pisiform are missing from this

specimen.

tibia and fibula, and similarity in proportions to

the manus, suggest that it is indeed the pes of

Thylacoleo.

The following specimens and casts of the

assembled right manus and left pes are lodged

at the South Australian Museum:

P16678:; left manus including fused carpus

P16678a, cuneiform P16G678b, pisiform

P16678c, fused metacarpus P16678d, fused

proximal and medial phalanges. P16678e,

distal phalanges P16678f,

P16679: right manus including scapholunar

P16679a, trapezium P16679b, trapezoides

P16679c, magnum P16679d, unciform

P16679c, metacarpals I to V P16679f, g, h,

i, } respectively, proximal phalanges I to V

P16679k, |, m, n, p respectively, medial

phalanges If to TV P16679q, r, s respectively

and distal phalanges I to V P16679u, v, w,

x, y respectively and sesamoids P16679z.

P16680: left pes including astragalus P16680a,

caleancum P16680b, cuboid PI6GROc, mavie

cular P16680d, metatarsals M, TT, TV V

P16680e, f. g, h respectively

kK, f, WELLS & B. NICHOL

Genera) description of the manus

The manus of Thylaceleo includes the car-

pus. metacarpus, phalanges and certain sesa-

molds associated with them (Figs 2A, B). The

carpus is composed of seven bones arranged in

two transverse rows, Articulating with the dis-

tal row of carpals are five metacarpals. Four of

the five metacarpals are closely apposed, and

each bears three phalanges. Metacarpals II-V

show little lateral mobility, whereas the first

(which is considerably shorter and more

robust) bears only two stout phalanges. The

distal phalange of the first digit bears am

extremely large ungual crest and ungual pro-

cess, suggesting the presence of large

recurved claw or unguis. Unlike digits II to V,

the first is capable of considerable divergence

us well as flexion. Ij appears to be opposable

to a broad spatulate pisitorm rather than to

the remaining digits.

Carpus: In Thylacoleo there has been a loss

and fusion of carpal elements to produce two

rows. which have # transverse convex cranial

outline and a concave caudal one,

The proximal row is composed of three

elements, the scapholunar (a fusion of

scaphoid and the lunar) the cuneiform and a

sesamoid, the pisiform, The scapholunar articu-

fates with the distal end of the radius and bears

on the trapezoides, trapeziuni and portion of

the medial face of the magnum. The cunei-

form, or ulnar carpal is greatly reduced, In

comparison with most mammals it is a small,

wedge-shaped hone concave on the proximal

surface for receipt of the styloid process of the

ulnar, and convex on the distal surface where

it inserts in a basin in the unciform, It

broadens caudally and bears a facet on the

proximal surface at the point of attachment of

the pisiform, The pisiform is short, broad ane

dorso-ventrally flattened and slightly expanded

distally,

The distal rows of carpals is composed of:

trapezium, trapezoides, magnum and unciform.

The trapezium (carpale 1) is a small flattened

bone in the form of a partial helix. It articu-

lates laterally with the trapezoides and distally

wilh the first metacarpal; the proximal end in-

serts in a notch in the palmar face of the

scapholunar. Small rotations of the trapezium

pbout its articulation with adjacent carpal

bones result in considerable lateral-medial dis-

placement of the first digit.

The trapezoides (carpale 2.) is a small wedge

shaped, proximo-distally compressed bone the

141

MANUS AND PES OF THYLACOLEO CARNIFEX OWEN

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Pi66e0

Fig. 3. Dorsal view of the partial left pes

(P16680) of T. carnifex, The cuneciforms,

metatarsal 1, portion of metutarsal TV and

the phalanges ure missing tram this speci-

men,

ventral face of which ts approximately triangu-

lar in outline. It articulates proximally with

seapholunar. medially with the trapezium, is-

tally with metacarpal [1 and Jaterally with the

magnum. The magnum (carpale 3) forms a

triangular wedge between the scapholunar and

the unciform which bear ou it medially and

laterally respectively, It has a narrow medially-

jaterally flattened palmar projection. lt bears

distally mainly on the third metacarpal with a

small portion bearing on the oblique lateral

projection of metacarpal I.

The unciform (carpale 4) is the largest bone

in the carpus, its lateral aspect is convex and

extends down if a contitiuous curve behind the

fifth metacarpal, A depression in the middle

of the proximal fate receives the distal portion

of the cuneiform. The convex palmar face of

this bone hears facets which articulate with

metacarpals IV to V and the lateral face of

the magntim,

Metacarpus: There are five metacarpals here

numbered | te V from the medial to the lateral

side. Mctacarpal | is exceedingly rohust, being

dlmost tWiee the cross-sectional area of the

remaining four metacarpals, yet only one-third

as long. The expanded head of metacarpal 1

articulates proximally with the distal end of

the trapezium and at its furthermost lateral

displacement contacts the medial face of meta-

carpal H close ro its proximal end.

Distally the axis of articulation of meticur-

pal T with the proximal phalanx is rotated

medially approximately 30° to the axis of

urticulation with the trapezium, Metyearpals 1,

TH, TV are closely apposed slender bones,

medially-laterally fluttened and rectangular in

cross section. Metacarpal Lf is slightly shorter

than TH ang TY which are of almost ecu

lenuth. The proximal ends of these bones ape

expanded and displaced laterally; the shape of

the urticulating facets restricting movement to

the dorso-ventral plane. Metacyrpal UW bears

proximo-medilly on the trapezoides and to a

lesser degree proximo-laterally on the magnum,

Metacarpal IL bears proximo-luterally on the

lateral face of the magnum while metacarpal

[V inserts proximally in a notch in the unci

form..

Metacarpal V differs from TH, If and 1V in

the possession of » pronounced lateral flange,

and Jacks the expansion and lateral displace-

ment of the proximal end; the articulating facet

suggesting a small degree of Jateral-meujl

rotation in addition to the dorso-ventral,

Phalanges: Digit | compused of ai proximal and

distal phalany is extremely robust, the proximi!

portion being approximately 3X the cross-see-

tional arca and 1,5X the length of the adjaceny

proximal phalanges. The distal phalanx bears

u large ungual crest and ungual process. Digits

Thro V, although contuining a medial as well

as a proximal and distal segment, are overall

shghtly shorter thin those of digit [, All the

distal segments bear the sheathing hase of small

hooded claws, The ungual process of these seg-

ments iy relatively long and slender and js less

recurved than that of digit 1,

Sesaiold bones: Qn the palmar surface of the

metacarpophalangeal joint of digits IIT and IV

are Iwo sesamoid bones articulating prioarily

with the head of each metacarpal, and

sccondarily with the palmar tubercles of each

proximal phalanx. A single sesamoid was found

associated with the meltucarpophalangeal joint

of digit 1; the small sesamoid found adhering

to the proximo-distal phalangeal joint of digit

1 is probably the other clement in this puir.

The mechanism of the grasp in tte minus

The relationship between articulating facets

was used to determine the range of movement

between adjacenc bones. A lateral view of the

range of movements in digits 11, HI, IV and ¥

is depicted in Fig. 4 —in the extended position

{A} and flexed (B), The distal phalanges ure

prevented from opposing the palm by restricted

movement al the metucarpo-phalaligeal joint

and to lesser extent at the proxrmeo-medial

phulanwenl joint.

In dorsal view, with digits extenued (Pig. 5).

the extreme divergence of digit F find the slight

divergence of digit V contrasts markedly with

the close apposition of digtts 1, LIT and TY

The range of Latero-medin) movement in

digit I, and its relationship tu the pisiform. is

MANUS AND PES OF THYLACOLEQ CARNIFEX OWEN 143

Hig. 4. (A) Lateral view of the right manus of

T, carnifex with digits fully extended;

(B) Medial view of the right manus of

T. carnifex with digits fully flexed,

also shown in Fig. 5. Flexion of digit I and the

pisiform would bring them into close opposi-

tion and it is suggested that this is the primary

grasping movement. With the digits in their

most divergent position it can be seen that the

plane of the grasp is almost at 90° to the

remaining digits. A similar grasp is found in

the brushtailed possum Trichosurus vilpectle

(Breeden & Breeden 1970),

Description of the pes

The skeleton of the pes in mammals is com-

posed of the tarsus, metatarsus, phalanges and.

ussociated sesamoids, The following bones are

missing, From the partial pes recovered from

the Victoria Cave; the cuneiform bones of the

distal row of the tarsus, the first metatarsal, all

the phalanges ond any associated sesamoids,

Fig. OA, B.

Tarsus: The tarsus normally consists of seven

tarsal bones arranged in two rows with a cen-

tral bone, the navicular, between. The proxi-

mal row consists of the astragalus and cal-

caneum, In Thylacoleo the tibia and fibula

articulate only with the astragalus which is the

second largest bone in the tarsus, It articulates

proximally with the tibia and fibula, distally

with the navicular and ventrally with the cal-

caneum and the proximal dorsal surface of the

cuboid. The proximal surface of the main body

of the astragalus bears a shallow trochlea

groove which articulates with the distal process

of the tibia, an oblique lateral facet articulates

with the lateral malleolus of the fibula, The

astragalus articulates with the calcaneum by

two distinct facets, a lateral concave one and a

medial convex one. The distal portion, (head

of the astragalus) bears a large convex,

rounded facet which articulates with the navi-

cular.

The calcaneum is the largest and longest

bone of the tarsus. It bears two facets on the

Fig. 5. Dorsal view of the right manus of T. car-

nifex showing digits Il to V fully extended

with digit 1 flexed. The outline of the

cuneiform and pisiform was produced

from a mirror image of those from the

Tight manus.

Inset—depicts dorsal view of digit I in

extended position,

tad R. T, WELLS & B. NICHOL

P (6680

Fig, 6. Elements of the left pes of T, cariifex

upper = dorsal view; lower = ventral

view. a, astragalus; b, calcaneum; c,

cuboid; ud, mavicular; ¢, f, g, h, metatar-

sals If, 1, 1V, V.

proximal surface which articulate with those

on the distal surface of the astragalus to form

a stable interlocking joint, The calcancal

tuberosity is short, robust and slightly flared

caudally. The distal surface of the caleaneum

is deeply grooved dorso-ventrally where it

articulates with the proximal surface of the

cuboid. The navicular is the central bone in

the tarsus, it articulates proximally by way of

a large concave latero-medial facet with the

astragalus, ventrally by way of a small concave

facet with the cuboid and distally by way of a

large latero-medial convex facet with the miss-

ing cuneiforms. A pronounced tuberosity on

the distal medial surface may articulate with

the proximal portion of the missing medial

cunciform. The temaining tarsal bone, the

cuboid, appears to be ag long as the combined

length of the navicular and the missing cunei-

forms. Tt articulates proximally by way of a

long narrow facet with the dorso-ventral

groove in the distal face of the caleaneum. Two

smal] facets occur om the dorsal surface; the

Jarger proximal one articulates with the astra-

galus while the smaller anterior one presum-

ably articulates with the missing lateral cunet-

form. [t bears a small plantar process. The

distal end forms a large slightly recurved oval

sutface which for 4/5th of its length articu-

lates with the proxintal surface of metatarsal V

and for 1/5th with the lateral portion of the

proximal surface of metatarsal IV.

Metatarsus: Metatarsals UW, IL and the proxi-

mal portion of [IV were found fused together

and in the vicinity of metatarsal V. A latere-

medial section through the assembled metatar-

sals has a convex dorsal outline and concave

ventral one (Fig. 7),

The structure of metatarsal I is unknown,

but the presence of a medial distal tuberosity

on the navicular is suggestive of a divergent

hallux comparable to Trichosurus. Metatarsals

Hsoand Wil are relatively short and slender

bones, their close apposition and general form

is similar to thal of the svndactylous toes in

Trichosurus, The size of the proximal portion

remaining of metatarsal IV and its relationship

to metatarsals II} and V indicate that it may

have been the longest of the three. The lateral

half of the proximal facet on metatarsal IV

articulates with the dorsal 1/ Sth of the distal

facet of the cuboid. A small proximo-lateral

facet articulates with the proximo-medial face

of metatarsal V. Metatarsal V is the most

robust of all four, It bears pronounced

tuberosities on the proximal and distal latero-

ventral surfaces. A large proximal facet articu-

lates with the cuboid, while a small ventro-

medial facet articulates with metatarsal IV.

In Thylacoleo the greater part of the body

Weight supported by the hind limbs is trans-

ferred from the astragalus onto the calcaneum

and portion of the cuboid. The surface area

of articulation of the caleaneum with the distal

tarsals is 20% larger than that of the astragalus

suggesting a distribution of weight through

these elements to the metatarsals in the ratio

3:2. The larger portion of the weight is borne

Fig, 7. Distal view of proximal sur-

face of metatarsals 2, 3, 4

and 5.

MANUS AND PES OF THYLACOLEO CARNIFEA OWEN 148

by metatarsal Y and a small portion of (V,

while the renmoining 40% is barne by digits |,

i, Ut and a portion of IV. However the

slender nature of metatarsals (and IM, and

ihe presence of a pronounced niedial Luberasity

on the nmavicular, suggests that much of this

load is borne by # robust divergent hallux.

The shallow trochlea groove, the plantigrade

tature of the stance (Pig, 3), the apparent sya-

duectyly of digits It and LL and the inferred

presence of a large divergent pollex are charac-

teristics which in Australian marsupials

resemble mast closely the pes of phalangerids,

Discussion

Che digits of the hands of most mammals

diverge when extended, and canverge when

flexed (Haines 1958), Such hands are ¢an-

sidered suitable for scrambling over rough

round and, if clawed, to climbing; however

they appear ulisultable for gripping branches of

irees unless used in a clasping manner as is the

case with Didelphiy virginiana (Haines 1958),

Cartmill (1974) examined the nun-primute

arboreal sciurids, and coneluded that in the

structure of the hand the arboreal genera were

distinguishable from the terrestrial forms only

hy their longer 4th digits and carpal pads and

sharper, more recurved claws, Bishop (1964)

found {lttle anatomical difference in the hanes

of terrestrial and arboreal carnivores.

Animals which climb by grasping branches

é.g many primate species, usually have prehen-

sile bands, If carpa-metacarpal movement ts

restricted to one plane, the divergent thamb is

classified by Napier (1961) as pseimdto-oppas-

able bul opposable if the thumb cai rotate

about the carpo-metacarpal joint so as to

oppose the remaining digits,

In primates the degree of curvature of the

carpal arch is related to the exten! of opposa-

bility of the thumb and to the size and fune-

uvonal nature of the long flexors of the digits.

These lie ina tunnel formed by the carpal arch

und the flexor retinaculum which binds the two

sides of the arch together (Napier 1961), A

deeply curved carpal urch is characteristic ot

animals in which flexing of the digits pluys a

mayor cole in locomotion and is likely to be

similarly deepened and strengthened to match

the exerted stresses of clasping during climbing.

The manus of Thylacaleo exhibits several

paradoxical features. (a) Digit | 1s capable of

wide divergence, and is pseudo-opposable yet

digits HW, TW and TY show little divergence ov

convergence, (b) Digit V could diverge

TABLE |

Hebe and digital formala af the memes vf somte mar-

niypehatdn

Digital formula

of mans

a>3>l1>s>t

Habit

Arboreal

Arboreul

Arboreal

Artureual

Specics

Phascolurcios cinereus

Peeudocheirtis pw regres

Acrentastey pv girenens

Cercattets cancinnus

8>S>i>i>l

t>4SiS>sS

Pelayriy brevicepy

i) Thvlacaleo carnlles

Arboreal = Trichesurity valpecila

Arboyeal/

Terrestrial

Arhoreal/

Terrestrial

Arboreal/

Terresirial

Arboreal/

Terrestrial

Terrestriul

Terrestrial

Desycerc Neerls eanide

Dasyurus Vivertians

Aitecinirus flavipes

Sminthopsis erassiounditad

Wombats ursihis

Macropus ewgenit

At gigas

Thy lainey cynoceptalus

Saerophilty haveisit

lsaodun abesulas I>l>4>s>i

Macrous lagatis

‘Terrestrial

Terrestrial

slightly; (c) AJL digits are clawed. the amgual

crest on digit | being extremely robust and

recurved while those on the remaining digits

are amall und slender with little recurvalure;

(d) the carpal arch is deep and robust yer the

digits with the exception of I, are capable of

only Limited palmar flexion.

Finch (1971) suggests that compated with

other marsupials Thylacelco has relatively long

limbs, and that the almost equal length of fore

and hind limbs implies 4 cursorial mode of

locomotion, However she also notes that the

scansorial (Trichesurns) and koala (Phaxce-

laretos) have “lairly long fore limbs” bur have,

ii contrast to Thylacoleo, mobile digits on the

forepaws which enables the animal to maintain

a firm vrasp of the branch of a tree. Prom our

analysis of the manus of Thyluceleo we con-

clude rhut, in spite of the limited convergence

in digits HI, TH and IV the animal does indeed

have an efficient and powerful grasping

mechanism in the opposition of digit [ to the

pisiform. Furthermore although the manus

would adopt u digitiyrade stunee on a horizan-

tal surface it would be ideally adupted to a

climbing grasp, Tree frogs of the family

Hylidae illustrate this point very Well. How-

eVer as pointed out by Bishop (1964) there is

little anatomical difference in the hands of ter-

restrial and arboreal camivores, o hand well

146

adapted to a climbing grasp could equally well

be adapted to holding prey or both. The struc-

ture of the hind foot of Thylacoleo suggests «

plantigrade stance when on a horizontal sur-

face, Due to the absence of the cuneiforms and

digit I of the pes, speculations about its func-

tions are less meaningful, nonetheless it is

remarkably similar in form and structure to the

pes of the brushtailed possum (Trichosurus)

which is used mainly in a clasping manner. The

slender nature of metatarsals If and ITI and

their close apposition is strongly indicative of

syndactyly and is consistent with its proposed

phalangeroid ancestry,

Table I lists the digital formulae of a range

of marsupials. The truly arboreal forms among

the phalangeridae all have the typically long

4th digit similar to Cartmill’s (1974) arboreal

sciurids. The digital formula for the manus of

Thylacoleo is similar to a large tange of semi-

R. T. WELLS & B. NICHOL

Acknowledgments

The field work in Victoria Cave is carried

out with the help of voluntary workers includ-

ing students of Flinders University and mem-

bers of the Cave Exploration Group of South

Australia, To all these people we express oul

thanks. We would especially like to thank Mr

D. Williams and Mr E. Bailey for their untiring

help with the excavations and their helpful dis-

cussions, and our thanks are also extended to

Mr P, Aitken and Mr N, Pledge of the South

Australian Museum for the loan of compara-

tive material,

Photographs were taken by Mr R. Brook of

Flinders University,

This work is supported financially by grants

from the South Australian Government

Department of Environment and Conserva-

tion, and the Australian Research Grants Com-

arboreal marsupicarnivores and terrestrial mittee. Dr M. Smith and Mr M. Tyler kindly

herbivores, criticized the manuscript.

References

Bisnop, A. (1964) Use of the hand in lower pri-

mates. Jn Buettner-Janusch, J. (Ed.) “Evolv-

tionary and Genetic Biology of Primates",

Academic Press: London.)

Breepen, S. & BREEDEN, K. (1970) A Natural

History of Australia: I Tropical “Queens-

land”. (Collins: Sydney.)

CaRTMILL, M. (1974) Rethinking primate origins,

Science 184, 436-443,

CLEMENS, W, A. & PLane, M. (1974) Mid-Ter-

tiary Thylacoleonidae (Marsupialia, Mam-

malia). J. Palaeontolagy 48, 652-660,

Daity, B. (1960) Thylacoleo, the extinct mar-

supial lion, Aust. Mus. Mag. 1960, 163-166.

Fincu, E, (1971) Thylacolee, marsupial lion or

marsupial sloth? dust, Nat. Hist. 17(1), 7-11.

Git_, E. D. (1954) Ecology and distribution of

the extinct giant marsupial “T/rylacolee".

Vict. Nat. 71, 18-35.

Haines, R. W. (1958) Arboreal of terrestrial

ancesiry of the placental mammals. Quart,

Rev, Biol, 33, 1-23.

Napier, J. R. (1961) Prehensility and opposability

in the hands of primates. Symp. zool, Soc,

Land, (5), 115-132.

Owen, R, (1859) On the fossil mammals of Aus-

tralia. 1. Description of a mutilated skull of a

large marsupial carnivore (Thylacoleo carni-

fex Owen) from a calcareous conglomerate

stratum, eighty miles SW of Melbourne, Vic.

Phil. Trans. R. Soc. Lond., 149, 309-322,

Weis, R. T. (1975) Reconstructing the Past—

Excavations in fossil caves, Aust. Nat. Hist.

13(6), 208-211.

SALT CRUST SOLUTION DURING FILLINGS OF LAKE EYRE

BY J. A. DULHUNTY

Summary

Investigations of solution of salt crust during the minor (1973) and major (1974) fillings of Lake

Eyre were carried out in relation to water depth, salinity layering and wave action. When water

depth exceeded half the maximum wavelength of surface waves, a saturated brine layer formed over

residual salt crusts and retarded solution. Following the filling in February 1974, a large area of the

thickest crust in central Madigan Gulf survived to the end of 1975 or early 1976, when water level

fell to less than half maximum wavelength, and the residual crust dissolved; the thickest part of the

Belt Bay crust survived until after September 1974 but dissolved by August 1975; in Jackboot Bay

the thickest crust dissolved by September 1974.

SALT CRUST SOLUTION DURING FILLINGS OF LAKE EYRE

by J. A. Dutnunty®

Summary:

Duntunty, J. A. (1977) Salt crust solution during fillings of Lake Eyre. Trans, R, Soc, S.

Ast. WICH), 147-151, 31 August, 1977,

investigations of solution of salt crust during the minor (1973) and major (1974) fillings

of Lake Eyre were carried out in relation to water depth, salinity layering and wave action,

When water depth excecded half the maximum wavelength of susface Waves, a saturated brine

Juyer formed over residual salt crusts and retarded solution, Following the filling in February

1974, a large area of the thickest crust in central Madigan Gulf survived to the end of 1975

or early 1976, when water level fell to less than half maximum wavelength, and the residual

crust dissolved; the thickest part of the Belt Bay crust survived until after September 1974 but

dissolved by August 1975; in Jackboot Buy the thickest crust dissolved by September 1974,

Introduction

Salt crusts, up to 46 ¢m thick, rest on

Quaternary sediments in the southern bays of

Lake Eyre North (Bonython 1956; Dulhunty

1974), Small inflows of river water covering

parts of the lake bed, termed minor fillings,

frequently dissolve some of the salt, but it is

soon redeposited by rapid evaporation, On

widely spaced occasions, as in 1949 and 1974,

sufficient water enters the lake to cover the

whole of its bed and dissolve all the salt crusts.

On such occasions, referred to as major fillings,

it has been assumed that the salt crusts rapidly

dissolve; newly introduced sediments are

deposited, and then new salt crusts reform on

top of the sediments as brines evaporate. How-

ever, nO precise or quantitative investigations

have previously beet made on the solution of

salt during fillings of the Jake, although

redeposition of salt after the 1949 filling was

described by Bonython & Mason (1953), and

Bonython (1956),

The distribution and thickness of salt crusts

in the southern bays of Lake Eyre North, were

surveyed under dry lake conditions in 1972

(Dulhunty 1974). In 1973 a minor filling of

the lake occurred when a limited quantity of

waler entered trom the Warburton River dur-

ing May, June and July. Tt flawed to the south-

western corner of the lake filling Jackboot and

Belt Bays, where solution of salt Was examined

in July 1973, By late December 1973 the whole

of the water had evaporated and the dissolved

salt was redeposited, and the Jake bad returned

to a dry condition, 45 reported by Mr M, Q,

Hughes of Muloorina Station.

Early in 1974 Lake Eyre filled to the greatest

known depth since European settlement, and

possibly for 500 years (Dulhunty 1975), The

lake commenced to fill early in February 1974

and remained full throughout 1975 and 1976

during which time the solution of salt was

investivated in relation to water depth, salinity

layering and wave action: results are recorded

in this paper.

Methods of investigation

Areas of undissolved sult crust were found

by probing the lake bottom with a long pole

from u boat. Where salt crust was present the

pole hit its surface with a resounding impact

and tended to bounce. Where it was absent,

the pole penetrated the soft silt which had

underlain the crust before it dissolved. This

provided a simple, reliable and positive test for

presence or absence of salt crust beneath lake

waters, Areas of residual crust were mapped

by running pole tests at intervals along pre-

determined lines, and plotting results,

The thickness of a residual salt crust was

measured by boring from a boat anchored by

three radially disposed concrete blocks,

A 15 mm wood-boring auger was attached

to lengths of 12,5 mm water pipe, screwed

* Department of Geology and Geophysics, University of Sydney, Sydney, N.S.W. 2006.

146 J. A. DULHUNTY

together so as (tO extend about | m above the

surface of the water when the suger rested on

the salt crust. A wood-boring brace was

attached to the upper end of the piping, A long

pole was then lowered vertically ta rest on the

salt beside the auger, und the position of a

mark near the top of the boring pipe was noted

on ja scale allached to the pole. The auger Was

then rotuled by means of the brace, und the

depth penetrated into the salt by cach rottion

was measured on the pole scale as the pipe

moved down, When the auger reached the hase

of the salt crust and broke through into the

soft imdertying silt, the depth of penetration

measured gave the thickness of the crust.

Water samples for salinity determinations

Were collected at different depths by lowering

the weighted end of 12.5 mm hose to the

required level anil then slowly pumping water

up through the hese to remove that taken in

during the lowering. Further water from the

required level was then pumped slowly into 4

sumiple bottle, Densities of the water samples

were measured very accurately, amd their

salinities, in terms ol grams of Lake Evre salts

per litte of solution, were determined from

density-salinity relations previously established

by solution of known weights of Lake Eyre

salts in water, and by gravimetric measurement

of total dissolved solids per lilre of Lake Eyre

hrines,

Results

Mujor Filling, [974

The dry lake bed survey of sale erust thick-

fess and distribution (Dulhunty 1974) Was

comied out about 18 months before the major

filling commenced in February 1974, This pro-

Vided an excellent basis of pre-filling control

data for invesligation of the progressive solu-

tion ef sall crusts Which occupied almost two

years

‘The principal investigations were carried out

in Madigan Gulf where the largest and thickest

sul ceust occurred betere the fillmeg, Dulhunty

(1974) termed the place of thickest salt and

lowest beight value ‘Centre Point’ and noted it

to be about 1 km southerly from the point of

thickest salt and lowest height recorded by

Bonython (19561,

At various stages throuphour the investiza-

tion, salinities were determined for water fram

the surface and at different depths, as shown

in Pigs (Band iC. The 12 values shown along

the salinity curve in Fig. 1B ure measured

values. Five were obtained from water sumples

collected in Level Post Bay (2 in March 1974,

| in December 1974 and 2 in Mareh and Apri

1975), and the other 7 were fram sanyples col-

lected at Centre Point. The steep salinity

gradient in bottom waters overlying residual

salt crust, Wustraled in Fig, 1C, was obtained

by measurement of salinities; in samples col-

lected From botlom waters at Centre Point anid

along ling A-B in Fig, 1A, during July 1974.

Residual salt crust thicknesses shown along the

salt crust thickness curve in Fig, 1H, were

measured by borings at Centre Point. at stages

during progressive solution of the crust,

The curve for water depth nm Fig, |B repre-

sents the mean of a large sumber of gauge

board readings in Level Post Bay and sound-

ings at Centre Point, The depth values shown

along the curve are mean values at different

places to facilitate reading of the diagrum. The

fall of about 04 m between November 1975

and March 1976, is based on interpolation

between readings in October 1975 and April

1976, Unfortunately no readings were made

between these dates, but the level must have

fallen, as a result of evaporation, by ut least ax

much as it did dunng the same period in 1974—

75 (see Fig. LB). No water entered the lake

during cither of these summer periods, Also

observations of shoreline wave cut features,

beneath water level later in 1976, indicated that

the luke bad fallen to 4 level equivalent to a

depth of about 3,2 m at Centre Point, There-

forte a fall to this depth is believed to have

oceurred erly in February 1976. An

appreciable volume of water entered the luke

from the Macumba and Neales Rivers and

Frome Creck late in February 1976, raising

jake water towards the levels measured in

April, May and June of that year,

The margin of the original sall crust hefore

the 1974 filling, is shown in Pig, LA extending

almost to the shorelines of Madigan Gulf. To

the northwest the salt crust thinned out and

disappeared where the Gulf opened on to the

Slush Zone of very thin discontinuous salt over-

lying soft mud nd slush extending geruss the

full width of the lake (Dulhunty 1974),

It had been generally assured that the sali

crusts of Lake Byre rapidly passed into solu-

Uoo ws soon as 4 Volume of waler in excess of

that Which would have been sufficient ta dis-

solve all the salt, entered the luke, The |974

filling of Lake Eyre North commenced in Feb-

ruary, and after six months filling with a

volume of water (ir in excess of that neces-

SALT CRUST SOLUTION, LAKE EYRE 149

HAMBIDGE BROOKS\

POINT 1 7ZON E

sg LUSH

ARTEMIA

POINT

MADIGAN

CAMPBELL POINT

FIG. 1A

SALT CRUST IN MADIGAN GULF

BEFORE FEB.1974, AND AFTER

SOLUTION AT STAGES TO MAR, {976

BLUEBIRD LY SHELLY I.

—

KILOMETRES

FIG, 1B satt crust SOLUTION, WATER DEPTH & SALINITY AT CENTRE POINT, FEB. 1974 to DEC. 1976

¥O so

WATER DEPTH ar CENTRE POINT

metres

SALINITY —gramsAitre

ALT CRUST THICKNESS,

Ss.

NG GRADIENT TO SURFACE

FIG. 1C

| RESIDUAL SALT CRUST AND

SALINITY LAYERING in BOTTOM

WATERS of MADIGAN GULF, —

JULY, 1974

{GRAMS per LITRE

MARGIN of SALT CRUST + ei

bLt_ji jf

RESIDUAL SALT CRUST HORIZONTAL

CENTRE POINT. SECTION ALONG A~B, FIG 1A bie

Fig. |. Relations of salt solutions to water depth and salinity in Madigan Gulf.

1st)

sary to dissolve ull the sall, a residial area of

crust remained undissolved im the central ares

of Madigan Golf Its margin, determined by

pole test Survey in July 197d, was as shown

in Fie. 1A. Further surveys in October 7974

und August [975 showed progressive, but slow-

ing solution of the residual crust to the posi-

tiony shown if Pig. | A. Pole tests by R. Clarke

and M. Atkinson in August 1976 and by the

author in Novernber 1976, established that the

whole of the residual crust had gone into solu-

tiou.

Measurements of salinity at different depths,

in Fuly 1974, established a saturated brine layer

from 10 to 20 cm deep on the solution surface

of the salt crust, Above the brine layer, there

eaisted felatively high salinity water up to 5O

em decp with a high salinity gradient, The

satiated brine layer did not extend more than

T km, afd the overlying high salinity water

more than 5 km beyond the edge of the sult

(Fiv. 1C), Away from the area of residual

sult crust, where salt had been completely dis-

solved, the lake Waters exhibited very little. if

any, salinity gradient, and no saturated brine

or high salinity water existed on the bottom.

The bottom saturated brine layers were

always almost opaque black in colour, rich in

dissolved hydrogen sulphide, and carried finely

divided organic debris amd clay which

appeared lo he almost colloidal, but could be

sepnrared hy filtration

Minor Filling, 1973

The minor filling of Lake Eyre during May,

Jone and July 1973 covered Belt and Jackboot

Bays and an atea west of Hunt Peninsula to a

level of -13.6 m A.H.D. Tt gave maximum

Water depths of 1.58 m to the base of salt

crusts in Belt Bay and 1.43 m in Jackboot Bay.

Some water, when aided by westerly winds,

flowed cust round Hambidge Point into Madi-

gan Gulf covering parts of its bed to depths of

less than O.5 om.

During July 1973, pole test surveys for salt

in Jackboot and Belt Bays indicated the

presence of relatively large areas of residual

salt crusts beneath the water cover, where the

thickest crusts were found in 1972 (Dulhunty

1974) and the deepest wiiter occurred in 1973,

Thicknesses of residual crusts were measured

hy boring and water samples were collected for

salinity determinations. Approximately one

third of the original thickness of crusts Jed

been dissolved by late July 1973, under the

J. A. DULHUNTY

conditions which existed during the filling, pro-

ducing salinities of 186 and 235 g// in Bell

and Jackboot Bays, respectively, The small

amount of water which reached Madigan Gulf

vovered part of the salt crust in central areas

of the Gulf and extended almost to Willow

Head. Measurements 3 km northeast of Willow

Head showed that about 12 mm of crust had

been dissolved by the end of July when water

salinity reyched saturation at about 325 //,

Sault solution in Jackboor and Beli Bays,

1974-75

Only limited opportunity wag available tor

investivation ef salt erust solution in Jackboot

and Belt Bays during the 1974 filling. Surveys

ol these bays were made early in September

1974, to ascertain the extent to which sall

solution had occurred. Pole test traverses in

Belt Rav indicated a roughly circular residual

urea of salt crust, about 2.5 km in diameter,

where the thickest salt occurred in 1972,

immediately 10 the northwest of a silercte

island situated 1.5 km west of Bonython

Head. Ata point 0.4 km west of the southern

tip of the island, water depth was 5.8 m, salt

29 cm, upper water salinity was 9.4 g// and a

gradient from 11 to 325 g// occurred in 1 m of

hottom water lying on the salt crust. This md}.

eated a layer of saturated brine on (he solution

surtuce of the salt crust asin Madigan Gulf, and

the 23 em of salt dissolved was slightly less, bur

comparable with the 29 cm clissulved In Madi-

gan Gulf by early September 1974. A pole test

for salt immediately west of the siferete island

in Belt Bay, by R. Clark, and A, & M. Atkite

son in August 1975, indicated that the Crust

had completely dissolved, and a surface water

sample collected at the same live gave a

salinity of 29.2 w/J,

In Juckboot Bay it was found that the mara

salt crust had dissolved by early September

1974, with only oceasional small patehes of

salt remaining on the bottom. Maximum water

depth was $.75 m and salinity of upper water

was 11.8 g// with a gradient from (3.4 to 15.0

g/l in 1.0 m of bottom water, where the salt

crust had originally been 23 em thick. This

means that solutian of 23 em of sult had

occurred in Jackboot Buy. as in Belt Bay, and

that it had just removed all the main crust. No

botton: brine layer was found in September

1974. due possibly ta the small area of thick

crust in Jackboot Biry. or dispersion afley solu.

lion of The salt

SALT CRUST SOLUTION, LAKE EYRE 1S}

Conclusions

Salt crast solution in Madigan Gulf, 1974-76

It was evident that the saturated brine layer

and overlying high salinity water regulated rate

of solution of the crust, Where water depth was

appreciably less than half the wavelength of

wind waves on the surface, disturbance from

wave action would have been sufficient to

prevent the formation of a brine layer, and salt

crust would have dissolved. At depths con-

siderably in excess of half the wavelength, dis-

turbance of bottom water would have been

negligible and a brine layer could have formed

over the salt. During the strongest winds

experienced on the lake, wavelengths of 8 m

were measured suggesting that salt crusts at

depths of less than 4 m would dissolve rela-

tively quickly. but at greater depths they could

be protected by overlying brine layers, There-

fore the history of salt crust solution in Madi-

gan Gulf during the 1974 filling, appears

evident [rom relations between salinity, water

depth and rate of solution of salt illustrated in

Fig, 1B.

With early vigorous inflow of relatively

shallow water across the lake bed during Feb-

Tuary and March 1974, solution of salt was

very active. Crust thickness was quickly

reduced from its original thickness of 46 cm at

Centre Point, to about 28 cm, producing water

salinities in excess of 30 g/l. Water depth

increased rapidly until the end of May, reach-

ing its maximum of 6 m in early June 1974,

Depths in excess of half the maximum wave-

length of wind waves were soon established.

With lack of wave disturbance in bottom

waters, bring layers Conimenced to form on the

residual salt crust, slowing down rate of solu-

tion. This, and dilution of upper waters by

cnormous volumes of fresh water, reduced

salinity of the upper waters to as low as 10

g// in August, From September 1974 to Octo-

ber 1975, water depth and salinity increased,

and the rate of salt solution slowed down and

almost ceased by September 1975, Water level

continued to fall reaching a depth of less than

half the maximum wavelength during October

where it remained until April 1976, In August

1976 no salt erust remained, and it is believed

that it finally dissolved during the period from

October 1975 to March 1976, as illustrated in

Fig, 1¢.

Acknowledgments

Tt is Wished to acknowledge (i) valuable

agsistance of Muloorina Station in field investi-

gations including that of Mr M. O. Hughes in

gauge board readings and collection of water

samples, (ii) the assistance of the Engineering

and Water Supply Dept, 8. Australia, in field

activities and discussion, (iii) the co-operation

of R. Clark and A, & M. Atkinson with their

boat Ibis, and (iv) research facilities provided

by the Department of Geology and Geo-

physics, University of Sydney.

References

—Its occurrence in Madigan Gulf and its pos-

sible origin. Trans. R. Soc. S. Aust., 79, 66-

92,

BuonyTHon, C. W. & Mason, B. (1953) The filling

and drying of Lake Eyre, Geogr, J, 11903),

371.330

Dutuunty, J, A. (1974) Salt crust thickness and

lake bed conditions in southern areas of Lake

Eyre North. Trans. R. Soc. §. Aust., 98(3),

125-133.

DuLwnunty, J. A. (1975) Shoreline shingle terraces

and prehistoric fillings of Lake Eyre, Jbid,

99(4) 183-188.

COORONGITE, BALKASHITE AND RELATED SUBSTANCES —- AN

ANNOTATED BIBLIOGRAPHY

BYR. F. CANE

Summary

A resurgence of interest in coorongite has been occasioned by the search for renewable hydrocarbon

energy sources. Coorongite is a biogenic polymer originating in profuse blooms of an aberrant alga,

Botryococcus braunii (Kutzing). B. braunii can exist in three physiological states, two of which

produce large quantities of polyene hydrocarbons. On the death of the colony, the hydrocarbon

metabolites oxidize and polymerize into a dark coloured rubbery mass, called coorongite, from

which a hydrocarbon oil can be obtained by pyrolysis.

COORONGITE, BALKASHITE AND RELATED SUBSTANCES—AN ANNOTATED

BIBLIOGRAPHY

by R. F. CaNng*

Summary

Cant, R, F. (1977) Coorongite, balkashite and related substances—an annotated bibliography.

Trans. R. Soc. §. Aust. W016), 153-164, 31 August, 1977.

A resurgence of interest in coorongite has been occasioned by the search for renewuble

hydrocarbon energy sources. Coorongile is a biogenic polymer originating in profuse blooms

of an aberrant alga, Bolryoceceus braunti (Kutzing), B. braunii ean exist in three physiological

states, (wo of which produce large quantities of polyene bydrocarbons. On the death of the

colony, the hydrocarbon metabolites oxidize and polymerize into a dark coloured rubbery mass,

called coorongite, from which @ hydrocarbon oil can be obtained by pyrolysis.

The lack of 4 comprehensive bibliography on coorongite and related bio-elaterites hus

been felt by workers in this field, The present survey, which covers the period 1866-1976,

makes good this deficiency,

Introduction

For more than a century it has been recog.

nised that algae provide useful information on

the genesis of petroleum, Increasing upprecia-

tion Of this fact, together with advances in the

lechniques of analytical chemistry, has resulted

in the chemistry of algae becoming an impor-

tant study for organic geochemists. One out-

come of these researches has been a much

better knowledge of the geochemical processes

which occur during the diagenesis of earth

hydrocarbons,

In addition to biologically derived liquid pet-

roleum, there are other types of hydrocarbon

deposit which have received recent attention,

These may be arranged into three groups,

namely:

(i) material resulting from the alteration of

petroleum to yield brittle, largely insoluble

solids such as wuttzilite, grahamite and

gilsonite:

(ii) abiotic hydrocarbons:

(ii) polymeric hydrocarbons

extant algae,

The purpose of this paper is to review the

literature of the polymeric group and to present

a bibliography of an important South Austra-

lian representative called coorongite.

Coorongite is a naturally occurring rubbery

polymer arising from a specific alga and, like

ansing fram

balkashite from Kazakh (Siberia), has its

genesis in prodigious blooms of an atypical

colonial alga, Botryococeus braunii, There are

only three known representatives of this type

of substance, all of which are of the same algal

origin and, presumably, have the same chemi-

cal composition, but only coorongite has been

fully examined. These representatives and their

main habitats are:

(i) Coorengite—The Coorong (South Aus-

tralia) +

(ii) Balkashite—Lake Balkash (Siberia) |

(iii) N'Hangellite—Lake N’Hangella

Africa),

There are dubious reports of other occurrences

but these have not been authenticated.

Until a decade ago, the problem of the

chemical composition of coorongite had

received only scant attention because it had

been tacitly assumed that the work of Stad-

nikoy (1929 eft seq) and others had established

the nature of the “algal oil”, The availability

of gas chromatography and computerized mass

spectrometry has now shown that the previous

assumptions were wrong and, over the last few

years, there have been important publications

on this topic. As recently as 1976, it has been

suggested (Hillen 1976) that the alga should

be fully investigated as a possible future source

of hydrocarbon energy.

(East

* QOueenshind Institute of Technology. Brisbane, Old 4000,

Lad RF, CANE

Because of Lhe present interest wm) hydro-

curhon-producing algae, it was decided to col-

lect together all reliuble information on the

nature of coorongite and related niwterlals,

with particular refenee to the role of Bo hrannit

in the formation of such bio-claterites, Tt

inevitable thal some citations will provide only

passing reference but, for completeness, such

material has been included, References dealing

with Boirvecocens purely from the viewpoint

of taxonomy or algology have been excluded

Unless the text has some direct bearing on

coorongite and its allies.

History of Coorongite

Although coorongilte was feporfed in 1852

(see Scrutton 1874—Suppl.), the first deserip-

tin was that given by Francis (1866—Suppl.),

followed by ag initial scientific report by Dyer

(1872). Prior jo these dates, the only micrest

in Coorongite (balkashite) had been us a squrce

of fuel by the Khugese tribe of Turkeston

(Morgan 1921). Some aboriginal tribes of

South Australia had aceasionally burned

coorengite to provide light.

Because of the appearance of coorongile and

its burning churaeteristics, Dyer (1872)

believed that it might be of mineral origin and

hence indicale the presence of underlying

mineral off. Arguments over the oryin of

coorongite Were to extend over the next half

century (Colyer 1974), If of mimeral origin.

it was argued, it was likely that “inillions of

tons” of petroletim was \nderlying the

Coorong. Hf it Were a Vegetal growth, it could

be grown to produce much oil, so it was said.

Early chemical analyses showed that the pyro-

lysate of coorongite Was largely Unsapormfiable,

This information was used by the "mineral oil”

protagonists as evidence of the relation of

cooronpite to petroleum. More reliable work

hy Roodle (1907) and Jater by Cuming (1902)

showed that samples of coorongite did contain

appreciable saponifiable matter gnd the field

observations of Broughton (1920) left no

doubt of the vegetal origin of cooronyite and

of its mode of formation,

After the establishment of the origin of

coorongite, the assumption that the “aleal” oil

was fatly ester temained the consensus for

nearly half a century, Scientific opinion was

strengthened by the comprehensive botanical

papers of Blackburn (1936) and Temperley

(1936), and the extensive organic 2eochemical

studies of Stadnikov & Weilzmarin (1924) anc

Stadnikey (1930). fn order to provide a suil-

able model for |is studies of the ongin of

cooronegite and torbanite, Cane (1967) used

elieostearic yeid for the synthesis of “syn-

thetic” coorongite, basing his premise on the

assuroption that the decarboxylated dimer

(Co Hs.) was the main “building block", His

hypothesis Was feasonably satisfactory but

there were still wnexplained features. Later,

Muxwell et af (1968) using conputerizect

mass Spectrometry, showed that the algal lipid

matter (ol the orange form of the alga) wias

uot fatty ester, as previously supposed, but two

isomeric §=polyene hydrocarbons (Co,H-2)

which they eulled botryaceecene and ie

hotryococeenc. At that stage it seemed whyious

thay botryocaccene polymer was the matrix of

coorongite. Further work by Cane & Albion

(1973) led te the conclusion that coorongite

Originated, not from the botryococcenes, but

from alkudiene hydrocarbons produced from

the green form of the alya allhough botryqeoc-

cene has a small but still important role. Later

work has shown that the nature of coorongite

varies more widely than previously supposed

and that the compositiun can be dependent on

the ecological conilitions during formation,

Post-depositional changes are also important,

Nevertheless, it is now clearly established that

coorongite is essentially a hydrogarbon poly-

mer arising directly by algal imetabolisin,

The nature of Botryococens braunii

Baryocercus diraveii is an ubiquitous abyn

Wistributed from the tropical to temperate

climates |i) nrany parts of the world, As well us

having a wide geographical occurrence, Af.

braunii ocevrs, ay fossil forms, in many geo-

logical eras as far back as the Ordoviciin,

mieed, as Dulhonty (1944) writes “that

organic evelution should have allowed the

Organism fo remain unchanged over the

immense lenath of time since the Permian coal

measures were laid down, is extraordinary”,

The alaa also occurs in w fossilised condition

in the Kukkersite of) shale of Estonia, and as

the main cantributur to the torbanites of New

South Wales, Scotlind and South Africa, Fossil

B hrawnlh has been found ih English and

Americal) peats. in the mud of elacial lakes,

in carbonaceous clays, and in some recent eur

bonate rocks: In extant form it is usually found

in fresh-water areas ranging in size from small

ponds to Iwkes alihough it can also be found

in brackish sWartps, hows and even In salt

Water. Under, as yet, undefined conditions, 2,

heave “blooms” to yield large areas of a foat-

CUUKONOGILE, BALKASHITE AND RELATED SUBSTANCES 155

ing muss of colénics which, on drying at shore

jines, have been reported as being of hundred

of square metres io extent and up to several

centimetres thick. The dried material after

some oxidgtion afd ageing is called coorongite,

The ubiquity of B. braunit has given rise to

some contusion because algolovists at various

places have nol recogtised the genus with

which they were working. Thus, the species has

been “rediscovered” under the generic names of

Pila, Reinvehia ant Elavephyton, The family

affiliation of the genus bas also been coantro-

versial, however, the work of Belcher & Fogg

(1955) using phytochemical criteria, has

placed the genus unequivocally in the Chlero-

phyceae..

Borryacoceus brawii exists in three distinet

growth states {Belcher (1968). Brown,

Kuights & Conway (1969)), each with quite

different lipid coniposition. The ¢hree states

wre;

(W) a green coloured, thin cell-walled rapid

growth stage, The hydrocarton metabolite

consists of ahour 30%) of straight chain

Cy7-Cy4 dienes:

{b) un orange coloured thick walled resting

stage. The hydrocarbons gre largely

branched chain C,, polyenes.

Both forms are colonial, and

contuining little hydrocarbons,

The morpholozy of B, brawl! has been well

documented, the classical contribution being

the collaborative papers by Blackburn (1936)

and Temperly (1936), These authors showed,

beVond doubt, that tarbanite and bowhead coals

bad a common origin in vast growths of this

alga und that the lipid mutter of the algal cups

provided the mionemer for the organic matter

of the minerals. The first instance of the recog-

wition of A drauagi as the oil forming alga of

these rocks was hy Zalessky (1926), although.

eatlier, he had believed the slza belonged to

the genus Pilla (Zalessky 1914), Zalessky's

research was mainly concerned with the identi-

fication of the organic matter of Esteanian on

shale and like others, kiter, be recognised that

bulkashite (coorongite) was the ‘peat staye™ in

the dlagenesis of algal organic rocks, Bath

falessky (1926) and Stadnikev (1930)

believed that the "oil” of this ulga was fally

matter and this opinion remained unchallenged

until Maxwell's (1868) work in the fast

decide,

Utilization of Coorongite

It was mentioned in the Introduction that

the burning of coorongite has been used by

primitive people us a source of heat and light,

The substance burhs very readily and it has

been stated that the infrequent and sporadic

finding of coorongite ts caused by the destruc-

tion of previous deposits by bushfires. In early

newspapers, one reads of layers of coorongite

ash many inches thick (there may be an

element of imagination if this).

After the establishment of the origin of

coorongite und the death of the “mineral oil”

theory, from time to time it has been suggested

that this material might provide a renewable

source of energy in the form of liquid or solid

fuel. The cultivation of the alga was suggested.

by Basedow (1925—Suppl.) and Sir Douglas

Mawson is believed to have put forward a

scheme at the 17th ANZAAS Congress in

Adelaide. At the 47th ANZAAS meeting in

Hobart, Hillen (1976) presented a case for the

further study of £, Aratnii as a renewable

hydrocarbon source and a_ potential fuel

supply.

Bibliography

The bibliography has been compiled from

first-hand study of all references except one of

Zalessky’s papers, In general, the verbal tense

ol the abstract is the present, except where the

past tense more suits the original publication.

A BIBLIOGRAPHY OF COORONGITE

AND COGNATE SUBSTANCES

containing abstracts of papers and occasional

nates

BReicHea, 1, H, (1968) Notes on the Physiology

of Botryoeoceuy braunii (Kuteing). Arch.

Mikrohlol, 61, 335-346,

A sludy on the growth of Boiryecocens under

yueying conditions. The rate of early growth

is exponential and the green colonies are

heuvier than water. After a few weeks a

stalionary phase is reached at which the

colonies ure red and floating, when subcul-

tured, they revert to the green phase. Under

nonmbloom conditions growth is slow and

large irreguku’ colonies are formed The paper

Uiseusses the effect of the environment on the

chemical composition of the wslga, The

rubbery substance found in lakes in Australia,

Africa and Siberia is w by-product of this alga,

These deposits fémain on the edge of lakes

and are the result of hydrocarbon residues

left behind en the death of the organism,

136 R, F, CANE

Bevcuer, }, H, & Poos, G, B, (1955) Biochemical

Evidence of the Affinities of RBarryececcus,

New Phytol. $4(1), 81-83.

The paper points out the Uncertainiles in

assigning the taxonomic position of

Botryocoecus, Extracts of B, hraunit were

chromatographed and the eluents examined

spectrographically, The absorption spectra and

other plytochemical criteria show “clearly

that the genus showed he placed in the

Chloraphyceae".

BLAckBURN, K. B. (1936) Botryococcus and the

Algal Cols. Part L—A reinvestigation of the

alza Berryvococens braunié (Kutzing), Trans.

R. Sac. Edin, 58(3), 841-854,

This is Part I of the classical paper dealing

with the role of Botryococcus in the formation

of balkashile, coorongite and boghead coals,

This paper traces the confusion of early

workers regarding the taxonomy of the genus

and notes how these misconceptions lead to

misnomers, The algal morphology is treated

in some detail including a discussion on the

cell anatomy. Comment is made that the

lipids of coorongite are very unstable and that

widely different analytical results may be

obtained depending on the age of the sample.

The material of the cell wall is chemically

highly unsaturated but, of exposure lo air,

the lipid matter becomes more inert and in-

soluble, This very thorough paper Jaid the

groundwork in this field for the next three

decades.

Boopte, LL, A. (1907) N'Hangellite and

Coorongite. Bull, Mise, Inf. R, bat Gdas

Kew 5, 146-151,

A description of the occurrence and nuture of

& gelatinous deposit found near Lake

N'hangella in Bast Africa, Examination af

specimens showed conclusively thal the

rubbery material originated in algnl growth Tt

Was suggested that the alga belonged “almost

certainly to the blue green algae”, Although

chemical data are net given, a separate des-

cription and examination of coarongite

(based partially on Dyer's observations) left

no doubt that both coorongite and n'bangel-

lite had a common Origin and diagenesis,

Boodle expresses indecision as to how the

“mucilage” of the alga became transformed

into “material showing the characters of bitu-

men” but assumes extensive chemical change

with loss of oxyden, He notes that n'hangel-

lite is divectly comparable to certain organic

minerals and mentions a likely connection

with the kerosene shales (of N,S,W,).

Brouctvon, A, C. (1920) Coorongite. Trums, R.

Soo, 8, Austr. 44, 386,

This short note occurs in “Miscellania" and

consists of only two paragraphs. The note

provides a first-hand description, one of the

few, of the actual formation of courongite

from the green alga in The Coorong, Tn view

of the uniqueness of this information, por-

tions of the submission are quoted verbatim:

“a thick scum, like green paint, is forminu.

This scum is drying on the water in places to

a semi-clastic substance, forming around reeds

.. « Like green paint, a quarter of an inch

thick, it covers hundreds of square Yards of

water, and as it drys it forms a skit like lin

seed oil drying on an overturned mass of

paint. This skin in places is yards in arca.''

“Today it is there in thousands of gallons, Tr

ds eoorongite in process of formation .

Every stage from the green, liquid, paint-tike

substance to the tough, elastic, sund-contain

ig conrongite muy be Observed, Scooped

with the hand from the surface of the lake

this substance, within a few minutes, chunges

before the eyes from a green liquid, which

Urops from the fingers, to » brown, plastic

solid, Large areas ore now drying in sheets of

coorangite.”

Brown, A. C., Kyuicuts, B, A. & Conway, E-

(1969) Hydrocarbon Content und its Rela

tionship to Physiological State in the Green

Alga, Botryococeus braun. Phrytochem, §,

543-347.

Botryocneeus is a peculiar alga characterised

by the production of hydrocarbons which

vary in Composition With its three pbhysioloxi-

cal states. ‘Vhe greet active-growth foin con

tains three fonalotous series of ulkadienes

in the Co7-Ca) range, substantially of straight

chain comfiguration with the vetreral formula

CHoy.2 where n = 27, 29, and 31, There és

a second setles, C,Ho,4, where, largely,

3%.

The brown resting stage contains large

amounts of hivhly unsaturated botryocec-

cenes, which may be up to &6% of the dry

weight of the colony. The dark-green urge

cell final stage has little hydrocarbon content

The chemical jjterrelationship of the stages

uncertain as the botryacoccene producing

stage reverts ta diene productiom when sub-

cultured, The paper establishes the main

chemistry of cach stage and shows that the

hydrocaybon content varies with growing con-

ditions.

Brown, H. ¥. L. (1908) Rec, Mines S. Aust. 4.

350-35),

This annual Seporr Contains a short review on

the occurrence of coorongite ond the presence

of mineral ofl, Wt records that coorongite “is

COORUNGITE, BALKASHITE AND RELATED SUBSTANCES 157

wn oindiarubber material” found near Salt

Creek which empties into The Coorong. After

Qn cxaminution of two bores in the area, the

Government Ceolugist dismisses any likeli-

hood of petroleum being found in the

Coorone ares,

Burotss, J, D. (1975) Batryececens Deeurrence

asain Ant in interpreting Paleoenvironments.

Abstract only ta Geasei, Man th. 154,

The full paper is a Useful stidy of the mor-

pHuloyy of Borryseocens vs oun indicator i

delermining the water environment during

sedimentation processes, The occurrence of

coorongite is discussed in the dight of likely

emvironmental conditions during. the bloom

stages. Tl appears that the main criterion in

determining whether the thin wall or the thick

wall stage ts predominant is whether the ulea

is inhwbiting fresh or brackish walters.

Cane, Ro F. (1967) The Constitution and Syn-

thesis Of Oil Shale. Proc, 7th World Petrol,

Congr (Mexico) 3, 681-489 (Elsevier: Bark-

ing) -

Although this paper is concerned with the

drigin of oil shale kerogen, ibe seetions

enciled “The Nature of Coorongite™ und

“Polymerisalion of Patty Oils’ deseribe the

likely reactions leading to the formation of

evorongite. Coorovgite analyses an its infra-

red spectrum are published. The important

coyttibution made by this pauper was the

hypothesis thal the covrongiic (and kerogen)

unit was a Cyyhsy polyene hydroacarbon. This

uniL was believed to be a dimer of a decarb-

oxylited trienoic Cy, acid. Later work showed

lat voorungiic urises directly from algal

hydrocarbuns and not tram an acid alkyl

chai, The CayH>, molecule corresponds

exuetly to the same elemental composition as

ihe Uimer postalated in this paper,

Cane, KR. PF. (1969) Coorongite and the Genesis

al OF} Shale Geeehim. Cosmochim. Aete 33,

257-245

the fistory and accuirrence Of Cobrongite are

reviewed ok theory is pur forward that

Courongile Gonsisis Jargely of a polymer of

“hydmcurbon chains containing some tutisal-

iwatioan” Carboxylic groups oceur wh chitin

Lerminauiens and whe non-polur end of the

livdrecarbon chain appears lu contain a dbene

structure. Ether cross-linkages are an impor-

tint feature of the mucromolecule. The

molecular weight of the monomer (or the

slimmer) iw about 490 Coorongite mass speotru

shuwed aw range of aliphutics, Alkaline — per-

mingunite oxidalion produced a waxy solid

with properties corresponding ta fatty acids of

hioh motucukw weight,

Cant, R. BF. & Arnnion, PR, (1971) The Phyto-

chentical History of Torbanites. Proe. R. Sec.

NSW. 104, 31-37.

The paper shows that the phytochemisiry of

Boryococcus leads to ah explanation of the

formation of coorongite. The alga is ¢excep-

fional in that it produces large quantities of

insaturated hydtocarbons which cun be either

branched or straight chain, depending on the

physiological state. ‘This state appears to

depend on food reserves und environmental

conditions, Coorongite undoubtedly arises

from the polymerisation and oxidation of

wyal hydracacbons. Spectral studies leave con«

siderable doubt as to the number of methyl

vroups wllached to the carbon chain “back

bone” of coorengite. Some alkyl oromatic

structures seem to occur in coordngite in uddi-

tion to hydroxyl and carbonyl yroups.

Can. R. F. & Aton, PLR. (1973) The Organic

Geochemistry of Torbanite Precursors, Geo-

chinr. casmochim, Acta 37, 1543-1549.

This paper extends the work of Cane who had

Suggested that the building block of eooron-

Hite wus ua decurboxylated polyene acid cor

responding to CyyH5x. Recent research had

shown that the alga produces Cy,Ho» directly

und not fram carboxylic acid. Further work,

ising mathematically derived data from pro-

ton yesonunce spectra indicates that long

chain dienes from the green form of

Bolryococcus are the precursors to coorongite

rather than the highly branched botryococ-

cenes, The orange form of Botrvececeus pro-

duces botryococcenes and these, in turn, form

“botryococcus rubber" but not coodrongite.

The opinion is put forward that, in the field,

dhere may be some contribution trom

Hraneched chain hydrocarbons as well as car-

boxyli¢ acids. Post-depositional suierobial

transformations also occur,

Carne, J. FE. (1903) ‘Phe Kerosene Shale Deposits

OF N.S.W. Mem. geol. Surv. NSW, 333.

‘this Moneeraph is devoted ta an exhaustive

study of the occurrence and natwre of the

NSW, oil sbales, The author quotes earlier

opinions that there is no real evidence af flow

gil in The Coorong, Reference 1s made to

coorongite (p. 109 and p. 302) ih connection

with other possible oil sources in Australia.

There is little additional pertinent informa-

on.

Corver, F. (1974) Fools Gold, Petrol. Gaz.

Mell, T8, 58-63,

The text provides no scientific mformution

but the generul historical survey is of much

interest, ‘The article gives the history of

138

Kk, F, CANK

eoormiupgite trom its discovery in 1852. ‘The

paper traces the vicissitudes in belief of the

interconnection between coorongite and flow

oil und the endeavours of carly entrepeneurs

to raise share capital to drill for oil, Mention

is made of previous mleas regarding the cul-

tivation of algae as oil producers,

Conacher, H, R. J. (1958) Coorongife and iis

Occurrence, Jn A, EB, Dunstan (Bd) ‘Oil

Shale and Cannel Coal", 42-49 (Institute of

Petroleum; Loodon)

The writer reviews the literature, pointing out

that, because of the sporadiv occurrence of

coorongite, definitive information on the

occurrence of deposits is lacking. Conacher

Visited The Coorong in 1935 and, althongh

he Was unsuccessful in observing coorongite

in the natural state, he provides » very good

description of {ts habitat. The article points

out that coorongite is also found in Western

Australia and discusses the environmental

conditions which appear to favour the growth

of the alga,

Cox, R. By BuriIncAME, A. L.. Witson, DL M.,

Ectiinton, G., & Maxwert, TI. R, (1973)

Botryococcene—a_ Tetramethylated Acyclic

Triterpenoid of Algal Origin, J, Chem, Sav,

D, 284-285.

As a result of the use of 18C muclear mass

fesonance spectroscopy, with pulsed Fourier

transform operation, a structural formula of

hotryococcene is suggested, It appears that

botryococcene contains eight methyl, eight

saturated und five unsaturated methylene, five

suturated and three utmsaturated methine car-

bon atoms. One saturated and four un-

anipatee quaternary carbons were also iden-

uhed.

Comina, «A. C. (1902) Cooronglte—A South Aus

tralian Elaterite, Prac. R. Soc, Vicd, 15, {n.s.)

(2), 134-140.

The paper gives a general description of

coorongite and provides references to its early

history and discovery. Cuming’s investigations

showed fhat this substance could be separated

into two portions depetiding on solubility In

carbor bisulphide. The soluble portion was a

Wwax-like solid, from the general properties

and chemical analysis of which the formula

(CyoHi20)., with x about 8, was assigned,

The insoluble portion, amounting to about

three-quarters of the sample, was given the

elemental formola Cy yHoyOs. As the solubles

were readily oxidised, Cuming suggested

{quite correcily) that the soluble portion may

become insoluble by the combined effects of

ageing. and oxidation. Ash analysis showed

that coorongile was not of animal origin,

Cumine, A.C. (1903) Courongite, A South Aus-

tralian Elaterite. Chen, News (London) 87,

306-308.

A recast of the previous reference in an

abbreviated form,

Daviv. T, W. K, (1890) Note on the Origin of

“Keruscne” Shale. Proc, Linn. Sac. N.S.

4/2), 483-500,

This paper, devoted to lhe origin of the tor

banites Of W.S.W, mentions the coorongit

theory of biogenesis of oil shale. This is a

Most important paper As in it David initiates

his algal theary of origin of oil shales. Micro

acopieal evidence on the origin of coorongite

strongly points t0 a Vegetative source bit

David leaves the matter undecided, He points

our thit, if allowance is made for the elimina-

tlon of oxygen, there are some striking poiuls

of resemblance between the chemistry of

coorongite and the “kerosene” oil shales of

Australia, The coorongite studies represent

only a amall portion of the work.

Doucias, A. G., Dowracm-ZapeR, K_, Ecuinten,

G. (1969) The Fatty Acids of the ales

Sberpococedy braunfi, Phvtochem. % 285-

The orange resting stage was purified, the

lipid extract frydrolysed and the methyl esters

examined by gas/liquid chromatography. The

extract was shown {0 contain a variety of

monocarboxylic acids ranging from Cy, 0

Cao With appreciable amounts of palmitic,

oleic and octacosenic acids, The total fatty

acids of the alga are in relatively small

amounts. A “synthetic coorongite’ derived

from botryococcene “rubber” was shown to

contain traces of various carboxylic acids.

Doucias, A. G,, Rotinron, G,, Maxweie, J, R

(1969) The Hydrocarbons of Coorongit

Geochim. cosmochim. Acta 33, 569-ST7,

A brief survey of the literature is given

followed by experimental data on the com-

position of coorongite extracts. The hydro-

carbon distribution ranged from Cy, to Cor

und consisted of alkanes, terminal alkenes and

some aromatic constituents. The fatry acids

ranged from n-Cy, to n-Cog with marked

even/odd preference and major components

at mCyq, Chg, The Cys isoprenoids con.

tained phytane and = pristane, but no

bolryococeenes. No n-alkanes were detected

im the orange bloom state of Botryoceceus.

The paper shows that hydrocarbons amounted

to 0.53% of the sample. As the extract

amounted to S0% of the sample it is unfor-

tunate that no attempt was made to charac

terise the whole extruct.

COORONGITE, BALKASHTITE AND RELATED SUBSTANCES 159

Dusncnry, 1. A, (1944) Origin of the N.S.W.

‘Torbanites, Prac, Linn, Son NSW. 69, 26-

ak,

The section (p. 31) entitled “Coorangite and

is Yelation to ‘Torbanite™ gives a short

necount of previous work and deseribes

samples collected by the writer, Dulhunty

defines coorongite as the “peatstuge” in the

formation of torbanile and gives convincing

feasons [Oo suppart his argument. The proper-

ties OF torhanile and coorongile are compared

and discussed,

Dyer, W. T. ‘Trstreron (1872) On a Substance

Known as Australian Caoutchoue. J. Bor.,

Lend, 10, 103-106,

This paper provides the first reliable descrip-

tion of coorongite. The pupet states “Lt con-

sists of sheet-like masses-—more than one inch

thick and is confined to a depressed partion

of the district, the bottum of which is sandy

and grass covered . , , or on the sides of

island-like elevations”. Dyer quotes ¢xaminu-

tions hy various workers which revealed “a

granular and cellular structure”. The sugges

lion is pul forward that it might be a crypto-

gamic plant but this is then discounted

because uf the inexplicably small amount af

oxygen, Dyer states, prophetically, that

coorongite “is practically o hydrocathow™ and

that the origin of the substance is likely to

Cause A great controversy.

Germ, E., Ord, I Scusemer, H. |}. & Bennett.

BR. a (1968) Olefins of High Molecular

Weielt in Two Microscopic Algae. Serence,

N.Y, b63, 700-702,

Laboratory cultures of B. braunii have been

shawn to contain alkenes with carbon num-

hers ranging from Ciz to Cys with one, two

and three double bonds, The Caz, Con and

Cy diolefins were predominant and hydro-

carbon distribution was similar to that of the

kerogen of certain ol! shales,

De Haurtrics, FE. (1923) Coorongite, A Petroleum

Product. Min, U., Loved, 142, 375.

A short contribution discussing the Origin,

oecurrence and properties of coorongite. The

writec errs in dismissing its vegetal origin on

the hasis of growth, This puper illustrates the

conceptual difficulties of early investigators

who could not reconcile a “pure” hydro-

curbon being produced by algae. Unsaponi-

fable oils were considered part af the mineral

kingdom and therefore must indicate le pos

sible presence of petroleum. His report to The

Coorong Oi] Company (14 pp., issued June

(9, 1925 Adelakle) farther elaborates of the

alleged interconnection between coorongite

and the occurrence of petroleum. De Haut-

pick urges further boring in the Coorong area.

Haurtpick, BE. (1926) Mote sur Je minéral

bitumeneux dit “Coorongite’ ¢t sur son

témaignage de la formation du pétrole. Bull.

Soc, eéol Fr. 26(4), 61-66,

The writer reviews previous work and men-

tians there are many previous papers includ-

ing a bibhography by L. Wattall (the present

reviewer can find no record—R,F.C.). Men-

tlon is made {hat coorongite appears to have

heen transported from Where it was formed.

\t 19 stated that the forniation of this “migra-

tory” mimeral has been recorded only in 1865

and 1920, after heavy rain, The paper gives

the physical properties of coorongite and

hotes that. on destructive distillation, the

material yields i whole series of “petroleum”

products, pone of which are saponifiable to

any extent, Although coorongite is of vegeta-

tive origin, de Hautpick reaches the conclu-

sion that “here is the true source rocks of

petroleum”, i.e. oi! glolmles inside vegetation,

This paper again Wlustrates the enigma facing.

early workers Whe couldn't reconcile the non-

saponifiable “fats” of coorongite with its plant

origin,

This is an important paper bat it is particu-

Jarly unfortunate thar no literature references

are provided, It might be mentioned, /nfer

alia, that Captain Hautpick was associated

with early share raising efforts in connection

with coorongite and petroleum in South Aus-

tralid.

Hinten, L. W. (1976) Prospects for Liquid

Hydrocarbon Fuels from Solar Energy via

the Alea Borryacocens braunii, — 47th

ANZAAS Conference (Hobart) May 1976.

This paper reviews the occurrence of

coorongite and states that there are four well

documented areas on the coastal sandy low-

lands of Australia where this deposit is found,

A study of the growth of coorongile suggests

thal this alga bas potentialities as an energy

source, however, the large Water areus

required for commercial production would

present difficulty in any large scale wnder-

taking,

Jackson, I. R- (1872) Coorongite or Mineral

Caoutchouc of South Australia. Pharm. J.

31, 763-4 & 785,

Portion of the paper is based of the eariler

observations of Francis {l86h—Suppl.)

regarding the occurrence and probable nature

of coorangite, Jackson found coorongite to be

‘resolvable into two educts: (1) saft semi

160

R. F. CANE

uid hike balsam , , , resembling vegetable

wax, and (2) @ tough pulverulent substance

... like) ~~. a medified form of cellulose”.

Kxtcnrs, BAL Beown, A. C, Conway, BE &

Livinsky, L, L. (I¥21)

Miboreprrern, B.S. (1970) Hydrocarbons

from the Green Porm of the Freshwater Alga

Bairyorsecus branati. Phytochem. 9. 1317-

13294.

Botevococeus Braanii occurs in two distinct

forms, the brewn resting stage containing up

to 70% of its dry weight of two isomeric

hydrocarbons. The green exponential growth

form is shown lo contain about 20% «iene

hvdtocarbons i the Cuz-Cs) range,

Ozonolysis, gas chromatography and mass

spectral data showed that the diene com-

pounds tad the general formula

CHeCH, (CH) ,CHLCH. (CHsly.CHs

where nm == 17, 19, and (5, in order of whin-

dunce. The disubstituted double bend is civ

form. in the same position as in oleic acid,

Balkash “Saprepelite”,

Petraleum (Berlin) 17, 437-440,

Supropelite from Lake Balkash is a dark

colonred substance alleged tu be produced by

the alga B. brannii, It burns with a sooty

yellow flame with a peculiar odour, The

newly tormed supropelite is green bul quickly

changes lo a yellow brown viscous muss

which can be cut by a knife. Organic solvents

may be used lo separate the material into a

hurd paraffinic wax-like substatice Which may

amount to 42% of the raw material. Destruc-

tive distillution gives a series of hydrocarbon

fractions, Even when kept for five years there

is no change in its physical properties,

Maxwrar, J. R.. Dousias, A, G, EGLinton, G.,

é& McCormick, A. (1968) The Botryococ-

cenes—Hydrocarhons of Novel structure

from the Alga Sotryocceeus brannii Kutaing.

Pitches, 7, 2U87-2171.

The nature, occurrence, and previous work on

B beuenil are discussed, Barly anulyses

showed a high lipid content containing a laree

ainount Of unsaponifiable matter, Hy the

application ef column chromatography, muss

wad infpa-real spectroscopy it has been shown

that the oily matter is not fatty ester as

previously believed. The lipids consist langely

of jwo polyene hydrocarbons of novel struc-

lure, Which have been called botryococcene

znd pobotryoceccene, The elemental campasi-

tion carresponds (o CeyHyss. Infra-red spectra

show the presence of exomethylene and viny!

groups tovether with much unsaturation. High

resolution spectraseopy indicate lwo terminal

vinyl! groups, six methyl growps, and perhaps

Mawson, D.

Morcan, Ro J,

Repwoop, B.

four exomethylene groups. A suggested struc-

ture is put forward, The behaviour of the

dead algal colonies and the formation of

coorongite can well be explaied in terms of

these hydrocarbons.

The paper presents entirely new evidence on

the composition of coorongite and marked

the end of the Stadnikov/Cane fatty acid

theory.

(1938) Further Discoveries of

Sapropelic Deposits in The Coorong Region

of South Australia. Of! Shale and Cannel

Coal, 50-52 (Institute of Petroleum: London),

The geological and topological features of

The Coorong arew ure given and observations

made on the flora of the saline lagoons, Tt is

stated that the locality is famous for the

occurrence of coorongite. Apart fram a use-

ful discussion on the area's geology, littl por-

linent information is presented of coorongile,

. J. (1921) The Occurence al

Coorongite in Central Asia, Chem. Engng. &

Min, Rev, 348, July 5, 1921.

This paper gives wo interesting description of

the bulkashite area in Siberia, Balkashitc is

found in Lake Ala-Kool which is a saline

extension of the fresh-water Lake Balkush,

Coorongite (balkashite) occurs above and at

the shore margin. Tt is a yellowish spongy

material which burns with 4 smoky flame

with a disagreeable odour, The deposits vary

from 2 feet to 10 feet in width and from thin

sheets up to 2” thick. Balkushite is always

mixed with wlgal remains and other general

sapropel, Morgan states that the alga grows

prolifically at the margin of the lagoon but

only in shallow water and preferably where

there is some surface disturbance, such vs

thut caused by breezes. He stutes that balka-

shite is never found in the main lake and it is

interesting to note (R.F\C) that cooronyile

fas never been recorded as growin in The

Coorong itself, Similar deposils have been

observed in Turkestan and elsewhere in

Siberia,

(1907) Report on a Sample of

Nhangellite from Inhambane, Portuguese

East Africa. Ball. Mise. Inf. R, hot. Gdas

Kew 5, 151-153.

A short description of N'hangellite together

with proximate analyses. Redwood showed

that destructive distilition produced, i addi-

tion to an aqueous phase and voke. an ally

product resembling mineral oil, The elemen-

tal analyses showed figures comparaliie to

those of coorongite avd that the Wwo depots

ure similar products of the same origin,

COORONGITE, BALKASHLITE AND RELATED SUBSTANCES 161

Stmrson, E. S. (1926) Coorongite, Rep, Dep.

Mines, Wes), Als, 234-235,

Coorengie has been found in many localities

near the South Coast of Western Australia.

Sheets of cuorongiie, up to one inch thick,

have been gathered from a swamp al Marta

callup, Dr Simpson, the government analyst,

dismisses any connection between the occur-

Tenves Of coorangifc and petroleum, He points

out that oily fiquids can he obtained by the

distillation of “almost any organic substance

fram coal to cucumbers”.

Stannimov, CG. 1, & Weizmann, A. O. (1929)

Transformation of Fatty Ackls During Geo-

logien! Periods TI. Brennsr-Chem, 10, 401-

403,

Patmets of hoghead coal were shown to con-

alst maloly of polymers of unsaturated fatty

acids. Such fatry ucids are shown to be the

parent substances of both coorongite and

balkashite, Armlysis has shown that coorongile

has a high content of saponifiable and un-

suponifiible organic acids. Polymerisation in

cooronpite is not extensive enough to make

it botally insoluble,

Srannikoyv. G. L. (1930) Die Entstehung von

Koble und Erdél—Die Umwandlung

der ovgaiischen Substanz im Laufe der

geologischen Zeitperioden. Sclir, Geb. Brenn-

stoffovel. 254 pp. (Rnke: Stuttgart),

Seetions of the book discuss the oxidation and

polymerisation of fatty acids into rubher-like

materials. The author Outlines the role of

csorongite ind balkashite in the formation of

hoghead coal. Mnvestigations show that balka-

shite is not a wax or of mineral origin bul ao

algal byproduct (wrongly termed Elavophyton

eaoradgiana) derived from fatty matter.

Courongite is similar in constitution and both

can be separated into soluble and insoluble

portions. The selubles are a thick yellow oil

‘Whereas the residuc is a rubbery soli. Stad-

nikey did a greal deal of fundamental wark

shawing that Oxidation and polymerisation of

rinsatirated fatty acids could give rise ly

either rubbers of brittle solids. His hypothests

of the decarboxylation of un acid to yield

hydrearbons was well supported by existing

evidence, Recent results, showing that hydro-

carbons Were the direct algal metabolites.

were possible onty after the perfection of

Vapour chronratopraphy. Stadnikov aserlibes

somewhat different diagnetic routes ta

eooroncite and Palkashite depending on con-

ditions of sedimentation and on the ratio of

uerohic to unaeroble environments,

STapNikov, G. L,, & Wozzeinska, 7. TI. (1930)

Transformation of Fatty Acids during Geo-

logical Periods IV. Brentist- Chen UW. 414-

416

Balkashite, like coorongite is denved Irom

Botryococcus brawnii and the remnants of

algae can be easily distinguished in samples of

both deposits. Fresh balkashite oxidises In the

aumosphere and undergoes a slow hardening

process. Some fally acids also show a great

tendency to polymerise to insoluble rubber-

like materints. Microscopic cxamination of

bopheuds and Moscow cannel coal also shows

evidence of jlgal origin,

Tempercey, B. N. (1936) Botryovoceus and the

Algal Coals. Part I], The Boghead Contro-

versy and jhe Morphology of the Boghead

Algae. Trans, R. Soe, Edin, 5803), 855-868.

The second part of this paper (see Blackburn

1936) deals with the interrelationships

between Bofryococens braunii, coorongite and

torbanite. The morphology and mode of

reproduction ot the alga are discussed and

illustrated with diagrams und photomicra-

gruphs, Temperley states that, in coorongite,

the characteristic cup-in-cup structure has

coalesced into a structureless rubbery mass.

The important conclasions of this paper laid

the groundwork for most later research on

the phytochemistry of algal oil shules and

showed that the “yellaw bodies" of Scottish

boghead and the torbanite of N.S.W. were,

in fact, remains of Borryococeus, The paper

also discusses the vegetal origin of coorangite

and its close relationship with balkashite.

Tryesses, R- (1925) Origin of Boghead Coals,

Pref. Pap, US. peol, Surv, 132, 121-135.

This publication deals largely with the origin

of the kerogen of bogheads and torbanites.

A section (pp. 127-130) is specifically

devoted to coorongite, ils hislory, occurrence

and composition, Thiessen failed to recognise

the genus but applied a new name of Elaco-

phyton caorengiana, Samples fram The

Coorong were eximined and described, Data

on the chemical properties of coorongite are

given including clemental and proximate

analyses. Thiessen believed thal it Was the oil

in the cell wall of the living plant which pro-

vides the “yellow hadies” of boghead coal and

the matrix af coorangite,

Traverse. A, (1955) Occurrence of the Oil Form-

ing Alva Borryococcus in Lignites and other

Tertiary Sediments. Micrepal. 1, 343-350,

Althotiah balkashite and coorongite receive

only uw mention, the paper presents a goad

review of (the ubiquity) of the occurrence of

R. F. CANE

Betryococcus and its role in organic rich

deposits. Traverse pre-empls the Later dis

covery that the alga itself may give rise to

hydrocarbons. The author also states that

other fossil genera described in the literature

are often Varieties of Botryococcus and that

it ocurs in a wide variety of geological ages

und geographical situations.

Wako, L. K. (1913) The possibilities of the Dis

covery of Petroleum on Kangaroo Island and

the Western Coast of Eyre’s Peninsula, Jul.

geol. Surv. 8. Aust, 2, 15-20.

The history of coorongite is given and pre-

vious investigations ure discussed. The opinion

is put forward that coorongite is Hot a petro-

leum product and onalyses would indicate a

hydrocurbon-like matenal of unsaturated

nature. Coorongite occurs on the north-

western shore of Murrays Lagoon on Kan-

garoo: Island where a scum is to be found

on banks below Hood level. The deposits are

associated with much vegetal detritus, Tt is

believed that coorongite might be an oxidized

product of “some pre-existing hydrocarbon”,

At the time of publication, this paper was the

best general survey and the references are

nearly complete. A bibliography is given.

Warp, L, K, (1915) The Supposed Oil Bearing

Areas of South Australia. Bull. geol. Surv. S.

Aust 4, 36-37,

Section 3 of this publication discusses the

alleged connection between cooronpite and

petroleum, Coorongite is found on the shores

ot Murrays Lagoon (Kangaroo Island) and

close to the Coorong area, It is found a “few

feet down on old shores on Which sund and

debris has subsequently collected” or at the

surface on banks associated with ponds after

wet seasons, The paper gives a good descrip-

tion of the occurrence of cooronzite and

asserts Lhat all facts predicate against its being

associated with petroleum seepages.

The opinion is put forward that coorengiic

Oonpinates from lowly vegetable organisms

which grow on the lagoons and that it bas a

acnesis similar to WN'hangellite (see ref.

Boodle, 1907),

Warb, L. K. (1916) A Review of Mining Opera-

tions in the State of South Australia, Issue

No. 24. p, 43, Department of Mines (Govt

Printer: Adelaide).

The report emphasises the falsehood of

associating coorongile with the occurrence

of petroleum. The report goes on to state that

pieces of coorongite which were placed “on

the crests of calcareous sand dunes... (was)

regarded as evidence that the material was

deliberately placed there with fraudulent

iitent™,

Warp, L. K. (1944) Search fer Oi! in South Aus-

tralia. Ball, peol. Surv. S Aust. 22,

There is a short reference to coorongite (p,

12) in the discussion of the occurrence of

petroleum in South Australia, A review is

given (p. 17) of the prospecting boreholes

which were put down in The Coorong in the

belict of the ussociation between mineral oil

and coorongite,

Witson, R. C. (1926) Reported oil at Kenderup.

Rep. Dep. Mines West, Aust., p. 78,

Mr. Wilson visited Luke Martagallup at the

request of local residents and collected pieces

of coorongite “about the size of dinner plates”

at the edges of the lake from shorelines which

have now dried up,

ZaLessky, M. D, (1914) On the Nature of Pila,

the Yellow Bodics of Boghead afd on Sapro-

pel of the Ala-Kool of Lake Balkash. Bull,

garte Géol, St Petershourg 33(248), 495-

507,

The paper supports the opinion that the

“yellow bodies” of bogheads were not algal

in origin but highly sculptured walls of the

spores of cryptogams, At that lime there was

confusion between the genus Pila and the

alga of balkashite. Zalessky records that,

along the Ala-Kool, this alg comes to the

surface of the water and it contains a con-

siderable amount of oil. Decomposition of

the alga on the shores of the luke generates

much hydrogen sulphide while the green plant

residue changes to a brownish rubber-like

mass,

ZaLessky, M. D. (1917) On Some Sapropelic Fos-

sils. C. r. & Bull, Sec, géol, Fr. 4th Series,

17, 373-379,

Because of further werk, Zalessky believed

that Estonian kukkersite is an oil shale

derived from Botryoceceus. This alga is simi-

lar to that found growing in Lakes Bieloe

and Kolomenskoe in the Tyer district of

Siberia. The sapropelite is also found in the

Ala-Kool gulf of Lake Balkash. In Lake

Ricloe, areas up to nine metres square ure

covered with a type of rubber humic jelly.

This sapropel has been used as a source of

ummonia for agricultural applications. Other

more Mature torms of the sapropelite ore

found in the Kamenkarita valleys in Siberia

und known as Kouswvriaslo,

COOURONGITE, BALKASHITE AND RELATED SUBSTANCES 163

Zanessky, M, D. (1926) Sur les nouvelles Algues

découvertes dans le Sapropélogéne du Lac

Beloe et sur une Algue sapropélogéne. Reve

gén, Bot, 38, 31-42.

A deseription is given of the deposits at Lake

Balkash. The deposits are formed by the

coalescence of vast numbers of the colonial

olga B. hraunii, which later dry on the shores

of the lake. The deposits are very thick and

resist decay. Zalessky gives a description of

the variety of Borryecoecns inhabiting the

Siberian lakes. Zalessky shows that the alga

has various forms depending on the ecological

and environmental conditions,

Supplementary chronological bibliography

Although not always of sclentific merit. early

references to the heated controversy over the

origin of coorongite are included m this survey,

The violent arguments urose because of dif-

ference of opinion as to whether coorongite

was of mineral or of plant origin. If the

former, it was tilleged that the discovery of a

large petroleum deposit could not be dis-

counted. Tf, on the other hand, the substance

was a vegetative growth, then it was said it

should be possible to cultivate the plant and

harvest the “oil”. It is interesting to note that

this latter possibility has been recently put for-

ward (Hillen 1976).

As it proved difficult to ascertain the author

of some vewspaper articles, a selection of the

main contributions is set down in chronological

order,

Francis, G. (1866) The Substance found near

The Courong. The §. Aust. Register 8.5.66.

This. well presented letter (p. 3, col. 3) gives

a general description of coorongite including

a reliable examination of its physical proper-

tics antl chemical reactions, Francis states that

all evidence points to a vegetative origin as,

under the microscope, coorongite has a cellu.

Jar structure, Francis considers it “to be

neither caaiitchoic, clastic bitumen, asplate

or petroleum but a peculiar fungoid growtle

and that it has no connection with coal or any

other combustible mineral”. The information

in this carly article is remarkably troe and,

although no quantitative data are provided,

the qualitative observations ure lurgely still

valid. The writer suggests that the substance

probably has somie financial value \f sufficient

quantity were available,

Mupeckn (1869a) The ddelaide Observer 3.7.49.

A letter (p. f col. 7) decrying any possibility

of a connection between coorengile and petro-

leum. Dr Muecke stated that coorongite has

been found on the top of recent sands and

shelly limestones and that it never had any

connection Wilh the Underlying strata. He sug-

gested (hat cooronyite «rose from allies of the

the grass trees because of its resinous and in-

flammable nature. “The damp yellow juice

exudes from the knot and bottom stalks dur-

ing the summer heat and flows on ihe sand

where it becomes hard, as every caoutchouc

does,”

Mupcke (1869b) Caoutchouc, The 8. Aust, Regis-

ter 30.7.69.

A turther letler (p. 3, col, 8) In reply to

another letter reaflirming his opinion on the

vegetal origin of coorongite and stating that,

under no circumstances, can coorongile be

regirded as of mineral oright—see also The

Adelaide Observer 7.8.69 (p. 13, col. 5).

Anon (1871) The S&S. Auut. Register 29.8,71-

The article (p. 2, col. 4) contains reproduc

lions of letters from J, Hooker of Kew Gar-

dens and from M- J. Berkeley regarding a sub-

stance called “mineral gamiboge" which is

believed to he # “collemal” in an imperfect

slate. The general opinion was that the

material (coorongite) is of Vegetal origin but

no firm views are given,

Awon (1871) A Singular Vegetable Formation,

The Advertiser 29.8,71.

An article (p. 2, col. 5) concerning the

dichotomy of opifien on the origin of

coorongite, pointing out that a “good deal of

money" had been spent in the belief that

“mineral gamboge” was an indication of

petroleum. Samples had been sent to Kew

Gardens and examined by M. J. Berkeloy.

Berkeley's opinion was that the substance

(coorongite) was “a collemal in an imperfect

sinte and au thin slic¢ shows necklaces of

eonidia”, The famous Dr Hooker, who had

written) to Adelaide “sets the matter at rest”,

Lc. Qorongite belonged to the vegetable king-

dom

A similar article also appears Ir The 8, Aust.

(nitials only) (1871) Coorongite—Vepetable

or Not? §, Aust, Express & Telegraph 19-71.

FLY,

A letter {p. 3, col. |) in reply to the previous

abstract. F.Y. affirms thal the descriptions

gives can onty he applied to globules of

mineral oil which are dispersed in water. "The

evidence is conclusive that fixes! petroleum ail

floating on water... forms a cnat of varnish

164

R. PF. CANE

or gum more or less thick according to the

accidents of position’. F.V. appears as a

strong adherant of the “mineral oil" theory.

This letter also appears in The Adelaide

Observer of 1.9.1871 (p. 3).

Wuirteti, H, A, (1871) The Adeluide Observer

30.9.7 1,

A reply to the previous reference, which

agrees that coorongite has an organic

structure, but, stating that microscope thin

section views cannot be explained in terms of

the plant origin suggested by M. J. Berkeley.

Further, any suggestion of coorongite being a

lichen is dismissed on account of the occur-

rence of diatoms embedded in the matrix.

Scrutron, T. U. (1874) Petroleum or Coal in S.

Aust. The S. Aust. Chronicle & Weekly Mail

Suppl. to issue of 21,2,74.

This article reports an address by T. U. Scrut-

ton (p. 1, cols 1-4) to the S.A. Chamber of

Manufacturers extolling the many virtues of

petroleum whilst calling for further invest-

ments in gil drilling. Scrutton refers to the

value of coorongite (which he confuses with

elaterite) and completely dismisses any pos-

sibility of its plant origin. He states that,

because of the high oil yield from coorongite

on heating, it is likely that, in the past. “mil-

lions of tons of oil have been projected from

subterranean sources” and it only needs

money to find it, Many aspects of the report

had no substantial basis at the time, and sub-

sequent efforts have shown that they were

erroneous. Nevertheless, some _ interesting

information on the early discovery of

coorongite is given as well as descriptions of

the area. The address by Mr Scrutton was

also reported in The South Australian Regis-

ter of 16.2.74 (pp. 5 and 6). The report was

also issued as a separate pamhplet under the

same title.

Basepow, H. (1925) The Adelaide Observer

14.8.25.

A contribution stating that authorities in the

United States had confirmed that coorongite

“consists in part of vegetable organism which

is oil bearing”. Basedow explains that he had

grown the alga under laboratory conditions

and “the little plants developed so plentifully

that the material grew up the sides and neck

of the bottle . . . If this can be done on a

small scale, why not apply it to the large?”

He further states that the material “could be

as valuable to the State as a gusher of liquid

oil” but no one seems to have given credence

to his suggestion.

Acknowledgments

The writer acknowledges the assistance

afforded by Mr L. S. Marquis of the State

Library of South Australia in providing some

of the early newspaper references, The writer

would also like to express his appreciation to

the Queensland Institute of Technology for

granting financial support to assist the publica-

tion-of this bibliography.

VOL. 101, PARTS 7 & 8 30 NOVEMBER, 1977

TRANSACTIONS OF THB

ROYAL SOCIETY

OF SOUTH AUSTRALIA

INCORPORATED

CONTENTS

Ludbrock, N. H. Early Tertiary Cyclammina and Haplophragmoides (Foramini-

ferida: Lituolidae) in Southern Australia - - - - - 165

Houston, Terry F. A new species of Diporiphora from South Australia and geo-

graphic variation in D. winneckei Lucas & Frost (Lacertilia:

Agamidae): - - - - - - - - - - 199

Annual Report of Council - - - - - - - - - - 207

Award of the Sir Joseph Verco Medal_ - - - - - - - - 208

Balance Sheet - - - - - - - - - > - - 209

PUBLISHED AND SOLD AT THE SOCIETY’S ROOMS

STATE. LIBRARY BUILDING

NORTH TERRACE, ADELAIDE, S.A. 5000

EARLY TERTIARY CYCLAMMINA AND HAPLOPHRAGMOIDES

(FORAMINIFERIDA: LITUOLIDAE) IN SOUTHERN AUSTRALIA

BY N. H. LUDBROOK

Summary

The genus Cyclammina is represented in southern Australian Tertiary deposits by five species

whose internal and external morphologies are described: C. complanata Chapman, C. otwayensis

n.sp. and C. paupera Chapman, which are restricted to sediments of Palaeocene to Middle Eocene

age, and C. incisa (Stache) and C. rotundata Chapman & Crespin, which usually occur together over

a wide geographical range and have a long stratigraphic range from Palaeocene to Early Miocene.

Their palaeogeographical and palaeoecological significance and their stratigraphic utility are

discussed.

EARLY TERTIARY CYCLAMMINA AND HAPLOPHRAGMOIDES

(FORAMINIFERIDA:LITUOLIDAE) IN SOUTHERN AUSTRALIA

by N. H. LupBROOK

Summary

Lupsprook, N. H, (1977)—Early Tertiary Cyelammina and Maplophragmeides (Foramini-

ferida: Lityolidae) in southern Australia, Trans. R. Soc. S. Aust. 10107), 165-197, 30

Novetnber, 1977.

The genus Cyclammina is represented in southern Australian Tertiary deposits by five

species whose internal and external morphologies are described: C. complaneta Chapman, C.

otwayensis nap. and C. paupera Chapman, which are restricted to sediments of Palaco-

cene to Middle Eocene uge, and C. incisa (Stache) and C. retundata Chapman & Crespin,

which usually occur together over a wide geographical range and have a Jong stratigraphic

range trom Palaeocene to Early Miocene. Their palacogeographical and palaeoecological siz-

nificance and their stratigraphic utility are discussed.

A species of Haplophragmeides occurring in Late Cretaceous assemblages and with

Cyclammina in (he Palaeocene is described as Maplophragmoides taylori n.sp.

Introduction

Since it was first recognised by Chapman

(1904) m ochreous brown clay from Brown's

Creck in Victoria, the genus Cyc/ammina has

occupied a prominent place in the literature on

early Tertiary sediments of southern Australia.

Tt occurs abundantly and in some parts of the

sequence, particularly in the Gambier Embay-

ment of the Otway Basin and in the Torquay

Basin, is the dominant and, apart from marine

dinoflagellates, almost the only marine micro-

fossil occurring in Palaeocene and Eocene

paralic silts and sands. In the past, its strati-

graphic potential Was discounted and only

superficial attention was paid to its internal

suucture, Knowledge of the internal structure

of Cyclammina species has been greatly

udvanced by the work of Bronnimann (1951),

Voloshinoya & Budasheva (1961), Serova

(1964) and Banner (1966, 1970),

The present paper is designed to vindicate

the early work of Chapman in correctly recog-

nising the genus Cyclamntina, separable into

several species, and to support the conclusions

of Taylor (1965) that the species in the Otway

Basin have stratigraphic value, Glaessner’s view

(1951) that Cyclammina was not a reliable

index fossil appears to have prompted Baker

(1953) and Harris (1965) to discount its

stratigraphic potential. Taylor's contention that

arenaceous forms previously assigned to

Cyclammina are, in fact, Haplophragmoides, is

shown to be based on a misunderstanding ol

the internal morphology of the species and to

have been influenced by bathymetric and

ecological interpretations.

Five species of Cyeluntmina are recognised:

C. complanata Chapman, C. inciva (Stache),

C, otwayensis nsp,, C. paupera Chapman and

C. ratundata Chapman & Crespin, A species

described by Taylor as Haplophragmvicdey sp.

B was correctly placed in Haplophragmoides.

Tt occurs in the Late Cretaceous with a small

benthonic assemblage and in Palaeocene assem-

Dlages with Cyelammina, and is here named

and described as Haplophragmoides taylori

n.sp.

Abbreviations used are as follows:

§$.4,D.M. Department of Mines, South Aus-

tralia

E.&W.S. Engineering and Water Supply

Department, South Australia

V.M.D, Mines Department, Victoria

B.PLN.L. Beach Petroleum No Liability

O.D.N.L. O1l Development No Liability

PAC, Point Addis Company

S.E.0.8, South East Oil Syndicate

CPC Commonwealth — Palaeontological

Collection, Canberra

GSSA Geological Survey of South Aus-

tralia Collection

166 N. H. LUDBROOK

GSM Geological Survey of Victoria Col- Chapman’s figures. The specimen figured as

lection Haplophragmium canariense (d’Orbigny) (pl.

NMV National Museum of Victoria Col- 22, fig. 2) is however, not on the slide: the

lection specimen on square 2 is not “H. canariense”,

WAM Western Australian Museum Col- as indicated on the slide, but a distorted

lection juvenile of Cyclammina complanata.

NZGS Geological Survey of New Zealand Chapman & Crespin (1930) described

Collection Cyclammina rotundata and C. longicompressa

Historical records of Cyclammina in southern

Australia

Chapman (1904) recorded Haplophrag-

mium latidorsatum (Bornemann), H. glomera-

tum Brady and H. canariense (d’Orbigny)—

and described Cyclammina complanata and C,

paupera—trom Brown's Creek. His sample was

collected by Kitson from a locality between

Rotten Point and the mouth of the Johanna

River 13.6 km northwest of Cape Otway, Port

Campbell Embayment of the Otway Basin, in

the lower 1.3 m of the Johanna River Sands

(see map and section, Carter 1958, p. 8). In

the section exposed between Rotten Point and

Brown’s Creek described as Section 28 by Rag-

gatt and Crespin (1955, p. 134), the lowest

25.6 m (84 feet) comprise the Rotten Point

Sands, and the overlying 24.4 m (80 feet) the

Johanna River Sands (Carter 1958, p. 10)

from the 0.6 m (2 feet) bed of “grey to

purplish-brown shale with Cyclammina” of

which Chapman’s material is presumed to have

been collected (Carter 1958, p. 10; Taylor

1965, p. 151).

Taylor (1965, p. 157) considered that the

presence of C. complanata and C. paupera gave

evidence of a Palaeocene age for the lower part

of the Johanna River Sands, and, while this is

possible, there is no firm supporting evidence,

and the age could be somewhat younger. Some

support for an age younger than Palacocene is

given by Harris’s (1971, p. 83) recognition of

his Proteacidites pachypolus Zonule (Middle

Eocene, P10 to P13 of Blow, 1969) (McGow-

ran et al. 1971, Enclosure 14.1) im dark

purple to black carbonaceous silts, sands

and clays he referred to the Johanna River

Sands, without identifying the sediments with

those described by Carter (1958). Harris

reported that derived Palaeocene species were

also present. However, preservation of the

Cyclammina spp. described by Chapman

(1904) is such that it is unlikely that they were

derived from older sediments.

Chapman’s figured specimens (1904, pl. 22)

are, with one exception, mounted on NMV

Slide P26049; they are clearly identifiable from

from subsurface micaceous marls (Micaceous

Marl Member of Carter 1964, pp. 22, 58,

Table 1) now renamed the Metung Marl Mem-

ber (Hocking 1976, p. 259) of the Lakes

Entrance Formation. The association of

Cyclammina species with Victoriella conoidea

and Almaena gippslandica (Carter 1964, pp.

22, 56) establishes an Oligocene to Early

Miocene age (Janjukian Stage) for the unit,

Globigerina euapertura zone of Ludbrook &

Lindsay (1969) equivalent to P21 (—N2) to

N4 of Blow (1969).

Chapman & Crespin (1932) recorded

Cyclammina incisa (Stache) from the same

unit.

Parr (1938) briefly described Cyclammina

incisa (Stache) from sediments he believed to

be of Late Eocene age from deep borings in

King’s Park, Perth, but which are now defined

as the King’s Park Shale, of Palaeocene age

(McGowran 1964).

Singleton (1941) erected the Anglesean

Stage for “the interval of time represented by

the deposition of the dark-coloured sands with

Cyclammina of cliff sections between Anglesea

and Point Addis”, which he considered to be of

Oligocene age. However, Singleton (p. 13 and

correlation chart) correlated other not neces-

sarily contemporaneous Cyclammina-bearing

carbonaceous sands with those at Anglesea.

Crespin (1943) recorded in detail the distri-

bution and stratigraphic range of Cyclammina

incisa, C. rotundata and C, longicompressa (=

C. incisa) in subsurface sediments of the

Gippsland Basin, Cyclammina incisa being

selected as the zone fossil for the Janjukian

Lakes Entrance Formation in which it was said

to be persistent (Crespin 1943, pp. 8, 10, 13,

78, Table 1).

Crespin (1950, p. 72, pl. 10, figs 3, 4, Sa, b)

described species occurring in the stratotype

Anglesean at Demon’s Bluff. The specimen

figured as C. paupera (pl. 10, fig. 4) appears

to be an immature C. incisa, and not C. pau-

pera as interpreted by Taylor (1965, p. 151, fig.

4 (la, b)) and in the present paper.

Baker (1953) recorded Cyclammina from

the Princetown Member of the Dilwyn Forma-

167

EARLY TERTIARY CYCLAMMINA AND HAPLOPHRAGMOIDES

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EARLY TERTIARY CYCLAMMINA AND SA PLOPHRAGMOIDES

tou, According to Hurris (1965, p, 78) palyno-

logigul evidence indicated a Late Palacocene

age for this member, and the possibility of

the microfloral zonule present in the upper part

of the Dilwyi Formation being as young as

Early Bocene was not excluded (Taylor in

Singleton 1968, 1973, p, 116); Harris 1971,

pp. 70, 78; Taylor 1971, p. 226). The Prince-

town Member contains Planeroralites cf,

pyeudomendrdti (Taylor in Singleton 1968,

1973, p, 116) and a latest Palaeocene to Early

Eocene age (latest P6 to P7) is indicated

(MeGawran et al, 1971, Enclosure 14.1),

Rayggatl & Crespin (1955) defined the

Demon's Bluff Formation in which C'yelant-

ming was the dominant and abundant form in

exposures Of the formation between Torquay

and Easter: View tn the Torquay Basin,

From palynological data presented by Har-

ris (1971, pp. 72, 84), the microflora of the

Demon's Bluff Formation falls withes bis

Triovirey Midenifieus zonule, which in its full

range ts equivaleal to upper Glohigerapsis

Index index to "Verhorotalia’ aeuleate plank-

tonic foraminiferal zones of Ludbrook and

Lindsay (1969) corresponding fo P13 to PIG

of Blow (1969) and of lute Middle to Late

Eagene age (McGowran et al, 1971, Enclosure

14.1), Sinpleton’s (1968, 1973, p. 116) chart

would indicate s Late Eocene age, PLA io PLT,

and Abele et al. (1976, p, 232) a Late Eocene

to Karly Oligocene age for the Anglesea Met-

her

Ludbrook (1963) reported Cyelanunine us

hemny well-represented in the early Tertiary of

the Gambier Embuynient,

Taylor (1965) in studies of subsurface sedi-

ments of the Port Campbell Embaynrent,

claimed that the genus had been misinterpreted

and that apparent labyrinthic jnternal structures

Were fot primary morphological features but

the result of replacement of agglutinating

cement by pyrite, and that quartz plucking was

responsible for o cancellate appearance of the

wall surface (Taylor 1965, p, 9), Using

evidence that the livitg Cvelanumina caneelluia

Brady was restricled Lo depths greater than 200

metres und that the Cvelammine-beuring sedi-

ments of the Otway Basin were laid down

under fairly shallow shelf to estuarine condi-

Hons, so that the presence of Cyelanunina in

the Dilwyn Formation would be contrary to

cnviroumental interpretations (p. 198), Taylor

transferred the species previously recorded in

C\yclummina to Haplopliragmvides. Unfor-

tunately. Taylor's reclassification was aceeptud

by other workers such as McGowran (165. p,

18), Singleton (1468, 1973, p. 117). Banner

(1970. p. 277) and Harris (1971, pp. BOL 83,

84), and the stgnificance pf Cyclammina in the

Australian early Tertiary was temporarily

pluced in abeyanee, Taylor, however, demen-

strated mainly from subsurface sections that

the Cyclanmina (°Maplophragmeaides’) species

had characteristic stratigraphic ranges, In this

he is Supported in the present paper.

Ludbrook (1971) wsserted that the species

were correctly placed in’ Cyelwmmind by

prewious authors. MeGowsran (1973) recorded

a Cyclammina facies in the Lacepede Foenva-

tion. Lindsay & Bonnett (1973) recorded and

figured “Cyeclamnina®’ cf, taewa from subsur

face sediments of Gligocene age in the

Waikerie urea of the Murray Basin,

Cockbain (1974) deseribed and figured,

including a thin section, Cyelunmina ineive

from the Late Eocene Pallinup Siltstone,

Plantagenet Group, southwestern Australia.

PLALE |

Figs S-8. 13. Crclamminag inctva (Stache), Fig. 5

GSSA F597. O.D.NLL. Mt Salt No. |, 939-942

m, Dartmoor Formation, Palacocene to Early Eocene, X46, Fig, 6—GSSA FIS96, FE. & WS.

Kingston No,

3, 65.2--69.2 m, Lacepede Formation. Late Eocene, umbilical view, X30, Fig, 7

—GSSA_ PAS96. FE, & WS. Kingston Nov 3, 65.2-69.2 m. Luccpedy Formation. Late Eocene,

uperminul view, X30, Fig: &-GSSA FIS95, O.D.N.L. Mt Salt No. |, 777.780 m, Dartmoor

Formation. Palaeacene to Eurly Eocene, %350 Fig. (3—GSSA FfA03, O D.NL. Met Salt No,

|, 750-753 m, Dartmoor Pormation, Palueocene to Early Bovene, natural dissection, View

inte chamber lumina showing hypodermal alveoluc in walls of two chambers, X46.

Figs 4 12,

Salt No. |,

14-17, Cyvelumenina rodundam Chapman & Crespin. Fig, 9—GSSA FIAl3. O.D.NL. Mi

966-969 mm, Dartmoor Formution. Pulueocene to Furly Foceng, NSO. Fig. [O—

GSSA_Fl6l2. B.P.N.L. Geltwood Beach Wo. |, 454 m, Tartwaup bormation, Middle Eocene,

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ing war

Formation, bite

small areal mpertares on the left side. X50. Fig. (2—GSSA Ff616b, Demon's Bluff

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S.A LUM, Waikerie Bore 28W, 146,3-147.8 m. Ettrick Pormation, Oligacene. juvenile speci-

men showing sherl open aperiure, XS. Fig

15—GSSA F612, BPN LO Geltwood Beach

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sseuyory) uswin| Jaquibyo ssauyoryd yeniur Sioqureyo SSOUYSIYT, isyaweEld ise] ur Jaquinpyl “ONL “Sdy

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170

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EARLY TERTIARY CYCLAMMINA AND HAPLOPHRAGMOIDES

NORTHERN

TERRITORY

WESTERN

AUSTRALIA

171

QUEENSLAND

SOUTH

AUSTRALIA

BREMER

Sibi

Obs. Bore F,

Por! Gawler

Workerne Nos. 2,

Glenelg River

Browns Creek

Castle Cove Coanalpyn No!

Mogg: Creek Kingston No

Demons Blut

Fossil Blutt

Kings Pork Bores

Beachport! No.l

Gellwood Beach No}

Mr Soll No]

CGP, County Grey

Wongoom Noé

South Stirling lac, Sé46

Adelaide, Morphew Sr

North Adelaide

wewrworaaonF wns

—_

Caiclew 20 Loong Nol

0 500

TORQUAY i

BASIN gyMELBOURNE

Lo Trobe No!

Ph. Bumberrah Nol

1000,

KILOMETRES

NOH ludbtock 1976

76-734 SA Department ol Mines

Figure 1. Map showing Cyclammina localities.

and recognised the pertinence of Robinson's

(1970) observations to the apparent dis-

crepancy between records of Cyelammina in

early Tertiary shallow water deposits and the

predominantly deep water depth range of the

genus.

Quilty (1974) briefly described and figured,

including a thin section, Cyclammina cf. incisa

from Fossil Bluff, south of Table Cape. Tas-

mania.

Source ef material

Details of the localities shown in Figure 1

are as follows:

Outcraps

1. Glenelg River, 2.4 km downstream from Kil-

lara Bridge, 14 km SW of Casterton, HAMILTON

1:250000 geological map sheet, 37°39°56'S,

141°17'23"E, Gambier Embayment, Otway Basin,

base of Dartmoor Formation, Palatocene to Early

Eocene (see Casterton 1:63360 geological map

sheet).

2. Brown's Creek, between Rotten Point and

mouth of Johanna River, 13.6 km NW of Cape

Otway, COLAC 1;250 000 geological map sheet,

38°46'°22"S. 143°23'14"E, Port Campbell Em-

bayment, Otway Basin, base of Johanna River

Sands, ?Palaeocene to Early Eocene (section des-

cribed by Raggatt & Crespin 1955, p. 134).

EARLY TERTIARY CYChAMMINA AND HAPLOPHRAGMOIDES 173

3, Casile Cave, LO km NW of Cape Otway,

COLAC 1.250000 gevlogical map sheet,

BRA URS. [43°26495S°B, Pert Campbell m-

hayment, Olway Basin, Johanna River Sands,

UM\dule to Late Eovene (see Carter 1958. p, 135

Ahele et. ul. 1976, p. 224),

4. Mogo's Creek, 4 km E of Gastern View,

QUEENSCLIFF |°250 000 geological map sheet,

48° 28°15"S, i44°04'19°B, Torquay Basin,

Demons Blult Formation, Late Eocene (seeuca

deseribed hy Raggatt & Crespin 1955. p. 108),

5, Demon's Bhof, Atglesea, QUEENSCLIFF

1:250 000 geological mop sheet, 38°24'36°S,

144° 139°R, Torquay Basin, Anglesea Member,

Demon's Bluff burmation, Late Eocene (section

deseribed by Raggatt & Crespin 1955, pp. 113-

117)

6. Fossil Bluff, Table Cape, BURNIE 1:250 000

ueologieal map sheet, 40°S8'55'S, 145°44 545,

Bass Basin, Freestone Cave Sandstone, Table

Cape Group, Barly Miocene N4/5 (Quilty 1974.

p 33).

Borcholes and Wells

7. King's Park Bores T and 2, Perth. PERTH

(<250 N00 geological map sheet, 31°58'S,

115°50'E, Perth Basin, King’s Park Shale, Palaeo-

cene (Mctiowran 1964).

8. South Stirling. Water Bore, Plantagenet Toca-

wen 5666, acar South Stirling, 20 km S of Stir-

ling Range, MOUNT BARKER 1;250000 map

sheet, 34°36'S, 118°08'20°R, at 12-21 m depth.

Bremer Basin, Pallinup Siltstone, Plantagenet

Croup, Late Eovene (Cockbain 1974).

Y SADM. -ldelaide, New Morphett Street and

Vicloria Bridges, Bore 11, Adelaide Railway Sta-

Won 25-25.6 m; Bore 12, south bank Torrens

Lake 16.76-17 m; ADELAIDE 1:250000 map

sheet, J4°SS'41'S, 138°38'02"E. St Vincent

Basin, Adelaide Plains Sub-Basin, tndifferemtiated

Yortuchilla Limestone-Blanche Point Transitional

Marl, Late Eucene (Lindsay 1969, pp, 34, 54).

10. SA.D.M, “Carclew", Narth Adelaide, section

TA74¥, hundred of YWatalu, ADELAIDE 1:250 000

eeological map sheet, 34°59°24"'S, 138°35'02"E.

19.8-19.9 m, St Vineent Basin. Adelaide Plains

Sub-Basin, Blanche Point Banded Marl, Late

Eocene (lindsay 1969, pp. 53, 58),

11, S.A.D.M, Ghseryation Bore F, Port Gawler,

T/A, hundred of Port Gawler, ADELAIDE

1250000 geological map sheet, 34°45%9'S,

138°27'B, 253-2594.5 m, St Vincent Basin, Ade-

luide Plains Sub-Basin. Blanche Point Banded

Marl. Late Eocene (Lindsay 1969, pp. 52, 55)-

12, S.A.D.M, Waikerie Bore 2, section 692, hun-

dred of Waikeric, 3.6 km SSW of Waikerie, 149

m; Hore 28W, section $53, himdred of Waikerie,

2.5 km southwest of Wajkerie, 146-148 m. REN-

MARK 17:250000 geological map — sheel,

34° 1318'S, 139°5730' EK Murray Basin, “glau-

gonitic clay unit", Ellrick Formation, Oligocene

(Lindsay & Bonnett 1973),

13, E. & WS. Caoonalpyn No. |, section $6, him-

dred of Coneybeer, PINNAROO [7250 000 geo-

logical map sheet, 35°41°05”’S, 139°49'S3"E,

69-71 m, 105-107 m, Murray Basin, Buceletch

Beds, 4, B, Late Eocene (Ludhrook I9AI, pp. 16,

17).

14. F. & WS. Xineston Ne. 3, section 374, hun-

dred of Lacepede. NARACOORTE 1:250 000

geological map sheet, 36°S0°S. 137°51E, 652-

69.2 m, Gambier Embayment, Otway Basin,

Lncepede Formation. Late Eocene (Ludbrook

L971. pp. 36. SR).

PLATE 2

Figs 18-20. Haplophrazmvides taylori psp. Fig. 18—Holotype GSM 64829 (|), La Trobe No. 1 Well,

292.41 m,

Dilwyn Formation, Palaeocene to Early Focene, X48. Pig. 19-—Paratype GSM

f4829 (5) Wangoom No. 6, Core 12, 596-601

m, Dilwyn Formation, ?Early Bocene,

umbilical view, X48, Fig, 20—Paratype GSM 64829 (5), apertural view. X48_

Fips 21, 22, 26, 27. Cyclammina puupera Chapman Fig. 21—Topotype GSM #4828 (13), Brown's

Creek, base of Johanna River Sands, ?Palaeocene to Early Fovene, N50. Fig. 22—GSSA

Ffe07 V.M.D, La Trobe No. 1, Core at 298.7 am, Dilwyn Formation, Palaeocene to Early

Eocene. umbilical view, X50. Fig. 26—GSSA F616, V,M.D. La Trobe No. 1, Core 5?

298.7 m. Dilwyn Formation, Palaeocene to Early Eocene, apertural view, X90, Fig. 27—

GSSA. Ff6sq, O-D.N.L. Mt Salt No. 1, 954-957 m,

Early Gocene, X90,

Parimoor Formation, Palacocene (|o

Figs 23-25. Cyclammina oiweyensis nsp, Fig. 23—Holotype GSSA_ Ff6d8, Glenelg River, base ot

Dartmoor Formation, Palaeocene to Early Eocene, X50. Fig. 24—Paratype GSSA_ Ff609.

V.M.0 La Trobe No. |, Core at 298.7 m. Dilwyn Formation, Palavocene to Early Eocene,

X50. Fig. 25—Paratype GSSA Ff610, V.M.D, La Trobe No, 1, Core al 298.7 m, Dilwyn For-

mation, Palacocene to Barly Bocene, X50

Fics 28-30. Cyelammina complanata Chapman. Fig. 28—GSSA_ #f593, S.B.0.S. Beachport No, 1,

602-604 m, Jeavings from Dartmoor Formation, Palaeocene, shawing appearance of distal

ends of alveolae as seen through the epidermis and afier crosion of epidermis, X40. Fig,

29—GSSA F594. §.6,0.8. Beachport No. |, 661-663 m, ?cavings from Dartmoor Formu-

tion. Palaeocene, showing areal apertures and thin, fine-grained epidermis with distal ends

of alveolae exposed by erosion of epidermiy, X37. Fiy, 30-—GSSA PES¥4, X26.

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EARLY TERTIARY CYCLAMMINA AND HAPLOPTRAGMOIDES 177

15. SE.O.S. Beachport No. 1. secon 20, hundred

of Lake George, PENOLA 1°250 000 geological

map sheet, 37°26°55"S, (40°02'1S"E, Gambier

Embayment, Otway Basin: 238-240 mo Ganibier

Limestone, Late Eocene; 259-271 m Lacepede

Formution, Late Eocene: 311-417 m Tartwaup

Formation. Middle Eoeene; 343-546 m Durimoor

Formution, Palaeacene lo Early Eocene; 602-664

m %cavings from overlying Darmour Formation

(Ludbrook 1971, pp, 52, 5).

14, BPN. Gelimooad Bench No. |, section 157,

hundred of Mayurfa, PENOLA 1:250000 geo-

logical mapyheet, 37°39'44"'S, 140°14'35°E, Gam-

bier Enmhoyotent, Otway Basin; 238-277 mo Gam-

tier Limesione, lite Eocene; 454 m Tartwaup

Formation, Middle Eocene (Ludbrook 1971, pp,

52, S@)-

17. O.D.N.L. Mr Salt No. J, section 783, hundred

of MacDonnell, PENOLA 12250000 geological

map sheet, 37°97'25"S, 140°37'43"E, Gambier

Einbaymentt, Qlway Basin. 585-954 m Dartmoor

Formation, Palaeocene to Early Bocene; 954-972

in Bahgalluh Formation, Pulaencene (Ludbraok

1971. pp 52, 56).

18, SA.MD. CGY, County Grey lignite inves-

ligutions, section 819, hundred of Young

PENOLA 11250000 geological map sheet,

A7 4449S, 140°37'54"E, Gambier Embayment,

Otway Basin. 28.96-29.26 m, Burrungule Merher.

Turlwanp Formation, Middle Kocene (Horns

1966, p. 2, 1971, p. BT).

19. VM.D, Hangeom Ne, 6, Warrnambool, water

exploration bore, PORTLAND 1:250 000 geo-

logical map sheet, 38°23'S, |d2°29' 1B’ E, Tyren-

Jarre Fimbayment, Otway Basin, Core 12. 596-

601 ny Dilwyn Formation, ’Rarly Eocene (Glenie

1971, Enclosure 13.6),

20 VIM.D. Lewng New 1, 28 km east of Warr-

nambool, water exploralion bore, COLAC

1250000 peologicul map sheet, 98°23°S,

I42°48°33°E, Tyrendarra Embaymeni, Olway

Basin. Core 12, 654-656 m, Nirranda Sub-Group,

Naruwaturk Marl Late Eovene (Taylar 1985,

fig, 5, p. 155; Glenie $971, Enclosure 13.2).

21. V.M. £o Trobe No, T Princetown, COLAC

122510 geologiea! map sheel, 38"4)49°S,

147 1049" E. Part Campbell Embayment, Otway

Basin, 297.6-298,7 m Dilwyn Formation. Palaeo-

cene to Barly Eocene (Taylor 1965, Fig. 5, p.

155; Glenie |971, Enclostire: 13,2).

2 PAC. No. 1 Bore Parivh of Bumberrah,

Metung, BAIRNSDALE 12250000 geological

fap sheet, 37°53°34"S, 147°50'14°B, Gippsland

Basin, 394.7 17 Lakes Entrance Formation, Oliga-

ene,

Stratigraphic utility

Allowing for differences in nomenclature and

taxonomic interpretation, Table 3 gives support

to the data presented by Taylor (1965, p, L55,

fig, 5), Palacocene to Middle Eocene faunas

arc represented by three species: C. camn-

plunata, C. otwayensis and C. paupera, These

species are not found im Late Eocene to Mio-

cene sediments where only the long-ranging €,

incisu and €\, rerundata occur,

The stratigraphic relationships of — the

Cyclammina-bearing formations are shown in

Figure 2. For the Otway Basin, the chart has

been considerably simplified and for the Aire

District slighly modified from those presented

by Abele et al. (1976) which should be con-

sulted for greater detail and for illustration of

the diachronous relationships between most of

the formations. The position ef the Glen Aire

Clay approximates ta thal expressed by Lud-

brook & Lindsay (1969, p. 371). The name

“Knight” (Sprigg 1952; Sprigg & Boutwkoff

1953) has been retained for the Group of Early

Tertiary non-marine and paralic sediments of

the Guntbier Embayment, in conformity with

its continued use by the Souwlh Australiun

Department of Mises in hydroxecloyical

studies of the Embayment and its use by most

wwuthors (Kenley, Rochow, Ludbrook, Taylor)

in the Bulletin on the Otway Basin (Wopfner

& Douglas 1971) and on the geological maps

ccompanving the Bulletin, tt is beyond the

scope of the present paper to disentangle the

nomenclatural priorities of the units eamprising

the Koight and Wangerip (Baker 1950)

Groups which have already been discussed at

some length by Kenley (1971), Glenie (1971)

PLATE 3

Pies 71-35, Cyolumruna ineisa (Stache). Pig. 31— GSSA F621. Demon's Blull Formation, Deman's

Bluth, Late Eovene, microspheric specimen. equatorial section, X30. Fig: 32—GSSA F621,

enlargement of part of bist whorl showing hypodermal and septal alveolac, X75. Fig, 33—

GSSA PEG33. Demon's Bluff Formation, Mogs’s Creek, Late Eocene, microsplreric speci-

men. equatonal section, X13. Fig. 34--GSSA F644, Demons

Blull Formation, Demon's

Bluff, Late Eocene microspheric specimennm, equatorial section, X73. Fig 35—GSSA Fisiz.

B.P.NA,, Geltwood Beach No. |

274-277 in, base of Gambier Limestone, Late Fuvene.

part of 2 chambers of collapsed specimen, equatorial section, X75.

Vig, an

Cy lannind cotendae Chapman & Crespin C?C. ineisa (Stuche). GSSA PfA38, S.b.0.S,

Beaehport No. I. 262-265 m, Lacepede Formation, Late Hocene, rriccospheric specimen,

equutorl seetian, X75.

N. H. LUDBROOK

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EARLY TERTIARY

and Abele ef al. (1976). Until the lithological

and pulaconrologicul relationships of all the

units of the two groups are fully defined, at

seems desirable to retain the nomenclature cur-

rent for the Gumbier and Port Carnphell

Embuyments ts Shown in Figure 2,

Sectioning techniques

On the whole, specimens of Crelauntnine

fron) the southern Australia Terhary are

abundant and reasonably well-preserved. Dis-

torion ts relatively uncommoan, and, apurt trom

pyrite infilling, most tests have fot been sub-

jecled lo chemical wction such as the secondary

silicification described hy Serova (1944). They

ure, nonetheless, difficult to section since the

eryinal cementing material was very thin,

apparently organic (see pl.o4, fig, 44). and

usually not preserved at all, particularly in out

crop specimens, Cireal cure has to be exercise

to uvend quickly reducing the specimen to an

unrecvanisable mass of quurte grains,

Scrova (1964) distinguished between hall

sections or “grinds” (shlifavaniyal achieved

by grinding down to the equatornal plane so

that the interfal structures are viewed in dircet

fight (see Serova 1964. pls. 2 and 3), and thin

sections (shlify) completed in the normal wey

hy turning the specimen over and grinding the

other side, the result being vieweu! by trans

mitted tight (see Serova pls. 410 7),

Taylor's figures (1965, p. 146, fig. 2) were

driwn from dissected specimens {b) or

“erinds” (a, ¢, d, “thick sections” of Taylor),

No thin sections were cut.

The first prablem encountered in sectioning

js fo Keep the specimens intact and to prevent

the agglutinated quartz grains from dispersing

during the gtinding process. Sectioning was

done Under the microscope, using one ground

glass slide, with or without grinding powder, to

grind the speeimen mounted in Lakeside

Cement on the other, The specimen was. kept

under constant observation during grinding,

aml at the first sign of a brewk through the

cement to the lest wall, the slide was reheated

and the cement redistributed over the specimen

hy fine needle, filling any exposed cavities

This method of recementation was continued

throughout the sectioning process, and turning

the specimen over for erinding was done while

the specimen was completely immersed ijn the

heated mounting medium. At oo stage was

grinding done on Lincemented test wall or on

empty chamber lumina, Any surplus cement

was removed when sectioning was completed

CYCLAMMINA AND MAPLOPNRAGMOIDES

and a drop of Nam pluced on the specimen

hefore covering with the cover shy. The inst

successful sections were those of specimens in

which the organic lining of the chambers and

alveolue were preserved (pl. 4, fig. 43) or

Where the chamber lumina and alveolae had

heen filled with pyrite and organic material (pl.

4+. figs 37-42). No distorted spevimens were

used,

Morphology

Species show a wide range of development

from the quile simple to non-alveolyr structure

of Cyclammine poupera to the highly-

Both of these extremes are prevent in’ Palueo-

cene sediments. and the only evidence of

evaluionary developrient is that both the

simple and the highly-developed forms do not

persist beyand the Middle Eocene. The internal

struciures become clear only when thin sections

ure cut, though SEM photographs are valuuhle

aids, Externally, species can he guite dificult

to separate, as they almost all appear to iter

grade, Considered at the adull well-developed

end of the dimensional range, species are Curly

readily separated [rum ane ynother, hut some

populations contain a hith percentage ol

immature individuals which, without the aid of

thin sections, are specifically identifiable with

only a mild degree of confidence, The overlap

of species determined on their external features

and plotted according to their relative dimen-

sions (diameter thickness) iy clear from Figure

3, and the overlap of three of the species when

the average diameter of the measured speci-

Mens 1s plotted against the number of chambers

in the Jast whorl (Fig, 4).

Overlap is particularly the case with C. iecise

together, and with C. camplanata and C, inéivu,

Which frequently occur together. As specimens

stratigraphic range (Palacocene to Miocene) wf

fossil Cyelamming in southern Australia and it

is not practicuble to cut thin sections to can-

firm the identity of all the specimens, its range

may perhaps be open to question; the species

has a knoWn range of Encene and Oligocene in

New Zealand (Hornibrook 1971). C. inetva

has been described ay a “common Oligu-Mio-

cene species of the ¢ireum-Pacifie region”

(Chang 1953) and recorded by several authors

from the northern Pacific margin (Asano

19ST, p. 6, figs 18. 19: Voloshinova & Buda-

sheva 1961, p. 207. pl. 10. figs la, fb, 3u, 4a,

dh; Chang 1983, p. 34. pl. 3. figs 3-10; Chang

1056. pl b. figs G8). but it has not been pos-

sthle i dhe present study ta confirm these

peeords,

Diagnostic external characters

Measurable parameters are tabulated in

Table | and shown graphically in Figures. 3 and

4, The 172 specimens of C. olwayensiy

measured are representative of some 600 indi-

viduals all of approximately the same dimen-

sions.

CC, vamplanata has a large, Nattened,

discoidal test, partly evolute, with 15 to 16

chambers in the last whorl of the fully-grown

adult and slightly sinuate sutures. [t has o thin

epidermis through which the alveolar hypor

dermal structure appears a4 a fine punctate to

vermiform pattern (pl 2. figs 28, 29), The

apertural face is high, laterally flattened. with

large supplementory apertures (pl. 2, fis 29,

40h).

torted has a moderately lorge hiconves invalute

test compressed at the periphery with 11 to 12

chambers in the fast whorl. straight sutures. and

2 finesgeained epidermis with scattered large

quartz grains. The apertural face is moderately

high and covered with coarse quarte grains,

Supplementary areal apertures are sometimes

visible between the grains (pl. |. fiz. 7, pl. 6.

fies 47-49).

C. onvevensts is a small, bicenvex, rather

inflated species with usnally 8 to 9 chambers

in the Jast whorl, a thin epidermis through

which the distal ends of the alveolae are visible

m 2 or 3 radiating series in cach chamber. Phe

apertural face ts a fairly high rounded arch:

small supplementary areal apertures ure some-

times visible,

C. pauperd is conspicuous in a population as

a osmall, fattened, biconvex, commonly

collapsed pauperate test with & to 10 chambers

in the last whorl avd a very fine-grained cham-

ber wall. Alveolae are visthle through the epi

dermis in the holotype and topotypes, but more

commonly the alveolar hypodernis appears

not to have developed (pl. 4, fig, 37). Tt would

appear to be a primitive type of Cvelamnina

similar to the small species close to C. e/egenis

figured by Banner (1970, pl. 13. figs J. ta).

notable for the absence of g supra-apertural

zone.

C. rotundata is a large, inflated species, with

a coarse-grained chamber wall and 8 to 11

chambers in the Jast whorl of the lully-grown

adult. The apertural face is a low broad areh

at all stages of development (pl, 1, figs 10, 11,

12, 14, 16): areal apertures are frequently

visible between the the coarse quartz grains

which cover the face (pl. 1, fig. 11).

Diagnostic internal characters

Parameters measurable from thin sections

are shown in ‘Table 2.

Almost the complete range of morphological

varnation illustrated by Banner (1970, pl. 13)

is present in the five species. The structure of

the hypodermis of southern Australian C vel

mine is alveolar, as deserrhed for the type

species Cyelannine cancellata Brady by

authors such as Bronnimann (1951), Serova

(1964) and Banner (1970), and not lubyring

thic as the genus has been conventionally

deseribed (e.g. by Loeblich sud ‘Vappan 1964).

The septal walls are perforated by septal arcal

PLATE 4

Fiz, 37, Cvelammina paupera Chaproun, GSSA Fi649, V.M.D, La Trobe No.

1, 298.7 m Dilwyn

Formation, Palaeooene to Earty Kocene, microspheric specimen, the black areas are pyrite.

X75.

Bias 48-40. Cyclameminda efwayenxis osap. Fig. 38-—-GSSA Fiads8, V.M.0. La Trobe No. 1, 298.7 m,

Dilwyn Pormation, Palieocene to Early Eocene, meguspheric specimen: black urens pyrite,

XT5- Fig. 49—GSSA FPf636, V.M.D. La Trobe No, 1. 298.7 m. Dilwyn Formation, Palaco-

cene to Early Hocene, vertical section; black

wreas pyrite, X75, Fig, 40—GSSA_ Ff647,

V.M.D. La Trobe No, 1, 298.7 m, Dilwyn Formation, Palaeocene to Early Eocene, micta-

spheric specimen; black wreas

Figs 1-44. Cyclarmemina coniplanata

yrile, X75.

‘hapman, Fig. 41. GSSA Pf627. O.D.N.L. Mt Salt No. 1, 881-

R84 om, Dartmoor Farmation. Palacocene to Early Eocene, juyenile specimen; black areas

pyrite. X75, Fig. 42—GSSA FI631l, O.D.N.L. Mt Salt No. 1, 917-920 m, Dartmoor Forma:

lion, Paolaemcene to Early Bocent. juvenile, X75. Fig. 43—GSSA Ff623, S.E.0.8, Beachport

No. 1, 411-414 m, Tartwaup Formation, Middle Eocene, part of vertical section showing

organic Jining of alveolne, some pyre in himina and openings of alveolae into chamber

lumina, but alveolae mostly free of pyrite, X75. Fig. 44—GSSA F619, §.£.0.8. Beachport

No. |, 651-652 m, Ycavings from Dartmoor Formation, Palaeocene, microspheric specimen,

equatorint section, alveolae mostly

pruins and

chintbers of the last whorl, X30.

free of pyrite which is in the form of scattered small

ane small aggresute imdieated by the black patch between the tenth and eleventit

RARLY TERTIARY CYCLAMMINA AND HAPLOPHRAGMOIDES [83

apertures (pl, 3, fig. 32) similar to those illus-

trated for Ciyclamrmine cf. elegany Cushman &

Jarvis (Banner 1970, pl 13, fig. la)-

Each species has a chalinctive alveolar put-

tern when tully developed. although there is

the same range of intergradation as that

presented by the external features.

¢. petiperd is a primitive type with a thin

epidermis and a simple alveolar or non-alveolar

hypodermis: the chamber lumina are widely

open and in successive whorls are only slizhtly,

if at all, offset as shown in equatorial section

where the overall pattern is of unbroken radii

the type of Cyvelammina cf. elegans Cushman &

Jarvis ws illustrated by Banner (1970) pl, 13,

ts |, ba).

C. inciva is similar in structure to the type

species C€. cancellate Brady. The chamber

lumina ure widely open, separated by rather

Massive septal walls about equal in width to

the interseptal width of the chamber lurmina

(pl 3, figs 31-34 and Cockbain 1974, fig.

670). Che alveolar pattern of the hypodermis

cancellata, consistine of simple, more-or-less

parallel tubes opening into the chamber lumina

(pl. 4, fig, 33; pl. S. fig 46; pl. 7, fig SO) and

tending to bifurcate beneath the epidermis (pl.

3, fig. 32). The alveolae, chamber, lumina and

septal areal apertures are lined with organic

material (pl. 3, fig. 32), which, it is assumed,

acted also as cement in the chamber and septal

walls,

C. retundata appears to be a variant of C.

ineiva characterized by the development of very

thick walls and reduced chamber lumina. In

fully developed specimens the alycolae are fine,

This, more or Tess parallel tubes.

with alveolae radiating from the chamber

lurnina in series oF 2 or 3 per chamber and

bifurcating below the relatively thick, simple

epidermis (pl. 4. figs 38, 40).

C. complantte is a highly complex form,

with thin epidermis, thick alveolar hypodermis,

thick septal walls penetrated by areal apertures

and much reduced chamber lumina. The alveo-

fae are lined with pseudochitin (pl. 4, figs 43,

44).

The nature of the organic cement is not

known for any species. Staining did not reveal

any calcile, as also Found by Murray (19730),

und pyrite is present as an infilling of the cham-

ber lumina and alveolac, Hedley (1963)

described the cement of living agglutinated

foramiotters ax an aekl mutcopolysaccharide

with organically hound iran, the cement being

reinforced in Cyclammina by incorporating Cer~

ric iron. The presence of irun int any cement

surviving the processes of fossilization could

not be confirmed.

Palacogeographical and palaeoecological

interpretations

Sediments containing Cyclammina along the

margin of southern Australia Were deposited

mainly during the early stages of the final

separalion of Australia From Antarctica and the

development of the Southeast Indian

(Southern) Ocean, Rifling was preceded by the

extrusion during the Middle Jurassic of

tholeiitic dolerites and basalts in Tasmania,

Antarctica and Kangarao Island (McDougall

& Wellman 1976). The Otway Basin was

initiated in the Late Jurassic or Barly Creta-

ceous by the opening of a long, deep trough

(“Otway Rift Valley” of Griffiths 1971. p. 77)

into which « great thickness of non-marine

clastic sediments, mainly Feldspathic and lithic

greywickes, way deposited. This was followed

hy continued subsidence and sporadic marine

incursions in the Gambier and Port Campbell

Embayments during the Late Cretaceous anc

Palaeocene and into rhe Eocene.

According to Weisse| & Hayes (1972), the

Oldest lineation identified in the Southeast

Indian (Southern) Ocean is anomaly 2), the

age of which is 54 my. B.P, or gnomaly 27

(56,5 my. B.P.), so that the formation of the

normal oceanic crust which recorded the mag-

netic lineations began in the Late Palaeocene ta

Early Eocene about 35 million years age.

Rifting and later patterns of sedimentation

were diachronous events progressive from west

to east (Griffiths 1971, p. 77). Limited marine

PLATE 5

Fig. 45, Cyelummnina inciva (Stache), NZGS FLOLO7!, King’s Park Bore No. 2, King's Park Shale,

Pulaeocene, X65.

Fin. 46. Cyelammina incine (Stache), NZGS FI00894, Denon'’s Bhiff, Demon's Bluff Farnation, Late

Eocene, section through chamber wall and chamber lumen, X155,

184 N, H. LUDBROOK

”

iui

iad

NUMBER OF CHAMBERS IN LAST WHORL

Ludbraok 1974

76-734 SA. Deportment of Mines

Figure 4. Relationship of number of chambers in last whorl and average diameter.

influence from the west was éxperienced during

the Late Cretaceous, when the sea entered the

western part of the Eucla Basin (Ludbrook

1958), the Duntroon Basin (Boeul & Doust

1975) and the Otway Basin (Taylor 1964,

1971; Ludbrook 1971),

The Eucla Basin experienced an open sea

environment during the Middle Eocene when

carbonate sediments containing abundant

planktonic foraminifers were deposited (Lud-

brook 1963, 1969, McGowran & Lindsay

1969). Middle Eocene sediments in the Gam-

bier Embayment of the Otway Basin, as exem-

plified by the Burrungule Member of the Tart-

waup Formation, are paralic, highly car

bonaceous clays and silts with sporadic

planktonic foraminifers (Ludbrook 1971, p,

57) and also Cyeclammina otwayensis. Open

sea conditions reached the Port Campbell

Embayment in the Late Eocene and the Tor-

quay Basin shortly before the beginning of the

Oligocene (Abele et al. 1976). Carbonate

sedimentation began in the St Vincent Basin in

the Late Eocene, The full extent of the easterly

transgression did not affect most of the Murray

Basin and the Bass and Gippsland Basins until

the Oligocene to Early Miocene (see also

Deighton et al, 1976, figs. 9-14).

Cyclammina flourished in the period when

the Southeast Indian (Southern) Ocean was

passing through the immature ocean phase; that

is, in the period between the sporadic murine

ingressions of the Late Cretaceous and the

easterly spread of carbonate sedimentation

extending into the Early Miocene.

The occurrence of Cyclammina spp. in

southern Australia may be compared with that

of C. canecellata described by Akers (1954)

from the Louisiana Miocene where the species

occurs in certain zones with planktonic fora-

minifers and abundantly in other zones to the

exclusion of other foraminifers. Studying living

C. cancellidta from the Peru-Chile Trench,

Theyer (1971a) observed that small and com-

paratively wide forms occur between 500 and

1000 m, with temperatures above 3° to 4°C.

larger and proportionately narrow forms live

at between 1000 and about 2500 m, below the

permanent thermocline and in the oxygen mini-

mum zone with oxygen content below 3 ml/I:

PARLY TERTIARY CYCLAWMINA

specimens Tying in deeper waters of note than

YOO m with temperatures below 2°C and oxy.

ven values above 3 tnl/l decrease slightly yn

dimeter but widen considerably,

Gregarious in habit, ts is found mest abun

dainty ag a “onespenus” assemblage (Akers

1954), or with almost all other genera

edclusted, in Pilaeocene to Early Focene para-

lw sediments of the Dilwyn Formation of the

Fort Campbell Embayment, Knight Group of

the Gumbier Embiayment, and in the Late

Eocene to Oligocene. Demon’s Bluff Formution

of the Torquay Basin, In the St. Vincent, Mur-

ray, Buss and Ciippstind Basins it is associated

wilh the carly stages of the diachronous murine

Tratsgressinas ob the Late Bocene, Oligocene

aml Barly Miocene The sediments in which

ibis abumdant and almost exclusive are ustially

highly carbonaceous: those in which it &

asocited will other henthonie forarmpnifers

(ire frequently glaueonitic, They are all

issumieal deo obave heen deposited ino shallaw

water, though palavebathyeretric studies have

Hor been done in any detail,

It was {Mls apparently anomalous habitat fer

whal has been uccepted as a deep-water genus

(Brady 188d; Akers 1954; Theyer 197fa, by

Roltovskoy & Wright 1976) that Taylor

(1965) found siMentt to aceept. The living

Cyvelanunine eancellare is widely distributed: in

oceanic waters olf the continental shelves ut

Jepths between 114 and S800 m, with a tem-

pertlure range between L1.6" at depths 278

uid B82 mM aml 1.2° at depth S800 m in the

North Pacific Ocean (Akers 1954). It does not

aceur in Antarctic waters (Theyer, 1971a).

The only species living in Australian waters is

and 122 min bryozoal mud off Maria Island

and at 128 mm off northeastern ‘Tustunie on

the continental shelf (Parr 1950), These are

shallow occtirrences compared with depths of

from 393 to [718 m for the widely distributed

dredged by B.A.N.Z. Antaretic Expedition

(1929-1931) as well a8 by — eurher

“Challenger”, Deutsche Sudpolar and “Scotia”

Expeditions (Pare 1950, p, 274).

According to Theyer (1971b). C. arbicularis

is the most characteristic index of lower buthyal

to upper bathyal zones in the Pacitice-Antarectic

Basin. Abundant specimens begin to appear at

upproximately 1800 m and its lower depth timit

posite. whieh is an index of abyssal depths,

hucomes significant ac 3000-3500 m and dis-

AND HAPLOPHRAGMOIDES 1kS

uppears between 4500 and 5000 nm. Tt under-

pocs size changes which probably parallel those

ol €. cancellata in the Peru-Chile Trench area,

While the bathymetry of the Cyedlersne/ee

hearing sediments, particularly in the Otway

Basin, and the younger occurrences in the St

Vincent and Murray Basins remains to be

Studhed, some broad interpretatians uf a tinnited

area, based on microfaunas (Tavlur 197|, figs

10-5 to 10-14) and computerderived meon-

struchions of the continental margin based on

quantitative seafloor spreading data (Deighton

et al, 1976) have been made. ft is, however,

not possible to compare the patlerns of hathy-

metric distribution of southern Auseralian

Cyelwmming species with that «beseribed by

Rohinson (1970) for Late Miocene ty thela-

cene species in the Gull of Mexico. Robinson's

(Vig. 8) distribution patterns shiw thet the

bathymetric distribution of Cyrelamilie spp.

decreased From a dominant abunlanee in the

ouler nentic—upper bathyal zone, herween 147

and 384 m depth, during the Late Miocene

Euirly Pliocene toa normal disterbution of ran

bul persistent examples trom the lower bathyal

gone (500 to 2000 on) inthe Barty Pleistucere,

The relevance of Robinson’s studies is to ¢is-

pel the conviction that the presence of Cycle

ating is indicative of deepwater sedimenturian

and to emphasise that the use of generic «dis

Irnibutions in palacoecdlogical interpretations

must be made with caution, partionlurly iF tie

abundance of the generic proup differs

mutkedly through time (Robinson 1970),

Robinson also noted tht, al the generic level,

spectinens associated with neritic asseimblaves

ure usually smaller, lighter in colour, coatser-

eouned, and with fewer chambers Jian those

associated with bathyal assemblages. tn the

opinion of Mutray (1973b) deepwater forms

of Cyelkimmina are larger that the shallow

Water representatives, Boltovskoy & Wright

(1976) show the depth distribution of C'velam-

ining as from the outer shell to the abyssal

zone Quoting Sigal (1952), Pokey (7958)

and Betienstacdt (1962). they state that genera

such as Maplaphragmotdes, Troehamrmnime,

Cyclammina and Bathysiphon require little

oxygen to survive,

The distribution and faunal associations of

southern Australian species are shown in Fable

3. Neritic assemblages in which Cyelammnina is

associated with planktonic and benthonic, other

than agglutinated, forms oecur principally in

fale Eocene or younger sediments. Phe two

Species represented i these assemblages, CC.

EARLY “TERTIARY CYCLAMMINA AND HAPLOPHRAGMOIDES 187

ineixa and C. retundata, are in general

moderately large ta large coarse-grained

species, the greater proportion of which have

relatively few chambers (see Table 3 and Fig.

4). The small fine-grained species C. otweryert-

vis and C, paupere with fewer chambers and

planata with more numerous chambers are

restricted to Palueoeene to Middle Eocene

paralic silts and sandy clays which, in the

Wanegerrip Group at Jeast, are of shallow

Witer origin (Baker 1950), Compared with the

neriti¢ assemblages the associated microfaunas

are poor,

The Jargest examples of C. conplunata were

recovered fron) the Dartmoor Formation mter-

sected in 8.2.0.8. Reachport No, | Well and

from oulerops an Glenely River. Consistently

laree examples of C, ineiva were collected from

the Demon's Blufl Formation al Mogg’s Creek,

There are oo data to suggest that these are

deepwater representatives of those occurring at

other localities.

Systematic descriptions

Order FORAMINIFERIDA Eichwald, 1830

Suhorder TEXTULARIINA Delage &

Hérourard, 1896

Superfamily LITUOLACEA de Bainville,

| 825

Family LITUOLIDAB de Bluinville, 1825

Subfamily HAPLOPHRAGMOIDINAE

Mayne, 1952

Geous HAPLOPHRAGMOIDES Cushman,

L910

Haplophragmoiues taylori n.sp.

PL. 2, FIGS 18-20

Haplophraymvides sp. Bo ‘Taylor, 1964: pl 59,

fio. 4: Taylor, 19G5: 181. fig. 4 (Sa, bh),

flolotype: GSM 64824 (1); figured paratypes

GSM 64829 (5).

Type localitv: VM.D. Ta Trobe No. | Weill,

Poncetown, 38'4)'49"S, 143°10 24°F, CO-

LAC 1:250000 geological map sheet, Core

S53B at 292,61 m, Dilwyn Pormatien, Palaeo-

cene to Early Eocene.

Material; 244 specimens of which 3 were

measured as typical; the holotype and 6 pira-

types from V.M,D. Wangoom No, 6, Core 12,

596-601 om, Dilwyn Formation, ?Early

Eocene; 48 paratypes O.D.N.L, Mt Salt No, 1

509-590 m, Daritioor Formation, Palaeocene

to Furly Eocene; 186 specimens ODNL. Mt

Salt No. t, 1533-3061 m Sherbrook Group,

Late Cretaceous.

Description: Test small, inflated, umbilicate,

involute to slightly evolute, with 6 to 7 cham-

bers in the last whorl, sutures straight, deeply

incised, periphery lobulate, umbilicus deep aud

broad.

Wall finely agglutinated, Apertural face

broad, high, aperture an interiomarginal slit

without lip or with a slight lip.

Dimensions) Holotype diameter 0.35, thickness

0.20 mm; average of 13 uncollapsed specimens

diameter 0.45; thickness 0.23 mm.

Remarks: Taylor (1964, p, 564; 1965, p. 151)

described (his distinclive small species of Map-

lophragmotdes, which is common in Late Cre-

taccous sediments and occurs also in the

Palacocene of hath the Port Campbell and

Gambier Embaymeuts of the Otway Basin. It ts

named in his hanour

Divtribulion; Olway Basin, Port Campbell and

Gambier Embayments —Sherbrook Group,

Belfast Mudstone und Paaritte Formations and

their equivalents, Late Cretaceous (Turonian-

Santonian (Taylor 1964) ); Wangertip Group,

Dilwyn Formation, and Knight Group, Dart-

moor Formation (Late Palaeocene to Early

Eocene),

Subfamuly CYCLAMMININAEF Marie, 1941

Genus CYCLAMMINA Brady, [874

Cyclammina cémplanata Chapman

PL, 2, FIGS 28-30; PL. 4, FIGS 41-44

Cyelummina complarata Chapman, 904; 228, pl.

22. fig. 7.

floletvpe, NMV Slide P26049 No, 6,

Type locality: Brown's Creck, between Rotten

Point and mouth of Johanna River, 13.6 km

NW of Cape Otway, 38°46'22"S, 143°23'| 4B,

PLATE 6

Pies 17, A Csclammina inciva (Stache), N@GS F 100894, Demon's Bluff, Demon’s Bluff Formation,

Late Eocene, aperlural view showing areal apertures; 1, X65; 3, enlargement of centre of

apertural Tee, X240.

Fig, 48. Cychimming ineisa (Stache), NZGS F100937, Deman's Bluff, Demon's Bluff Formation, Late

Evcene, apertural view showing areal apertures and coarse grains of apertiral Face, X34,

188

base of Johanna River Sands, ?Palaeocene ta

Early Eocene,

Marerial; (OL Specimens of which 54 were

measured and & sectioned. From outcrop—

Glenelg River (7): from boreholes—O,D.N.L,,

Mt Salt No, 1, 585-954 m (84); S.B.O.S.

Beachport No. |, 408-603 m (9); B.P.NLL.

Geltwood Beach No. 1, 579 m (1),

Description: Test large, composed mainly of

quartz grains, planispiral, flattened, diseoidal,

partially evolute, with 2 whorls in the mega-

spheri¢ form dnd 3 to 4 in the microspheric

form: chambers 6 to 16, but 15 or 16 in the

fully-grown adult, periphery narrowly rounded,

very stuhuly lobulate, umbilicus well-defined,

hallow, sutures incised, sinuate.

Wall ayglutinated, thick, epidermis thin,

smoothly finished, imperforate; hypodermis

thick, alveolar, with a series of parallel alveo-

luc distal to the septal wall but branching from

the chamber lumen, all alveolae bifurcating

just below the epidermis which they da not

penetrate, the distal ends of the alveolae can

be seen through the translucent epidermis as a

fine punctate pattern or when they have been

expased by erosion of the epidermis.

Septal wall thick, arcuate, thickness. as much

as 6 Times the interseplal width of the chamber

Jumen at its maximuny width in equatorial sec-

tions im section cach septal wall showmp at

least 7 more or less. parallel alvealac extending

from the supplementary apertures.

Chamber lumen mueh reduced, curved, both

chamber luming and = alveolae lined with

pseudachitin ("teetin”, a combination of pro

tein and carbohydrate (Hyman 1940) ).

Apertural face high, more or less fattened

lnterally and rounded at the periphery, covered

with fine quartz grains and with conspicuous

supplementary areal apertures, cach sur.

rounded by a rim. Aperture an interiomarginal

slit at the base of the apertural face.

Dimensions, Holotype diameter 2.0 mm, OF

54 specimens measured, diameter 0.45 to 3.75.

average 1.08; thickness 0.18 to 1.12 mm, aver-

ae O39 mm: average ratio diameter > thick-

ness 2.87)

Remarks: This i8 4 rare species, occurring only

from the Palaeovene to ?Barly Bacene in the

Otway Basin, [It is a complex form of Cyelam-

mine with greatly reduced chamber lumina

and an extensive alveolar pattern similar to

that of the Cyclanunina pilveensis Yoloshmova

& Budasheva group (see Banner 1970, pl 4,

figs 11, 12: pl, 13, figs $-7) and Cyelomemina

N. H, LUDBROOK

aff. rani Ishizake (Banner 1970, pl. 13, figs 3.

4). From its external features it is obviously

very close to Cyclainming sp. of Chang from

the Shihliufeng sandy shale of the Liuchungchi

oilfield, Taiwan (Chang 1956, pl. 1, figs 1-3).

Distribution; Otway Basin—Dartmoor Forma:

tion (Palucocene to Early Eocene) and lower

part of Johanna River Sands (?Late Palaeocene

to Early Eocene).

Cyclammina incisa (Stache)

PL, 7, PIGS 5-8, 13: PL. 3, FIGS 31-35;

"FIG. 36: PL. 5, FIGS 45, 46; PL. 6, FIGS

47-49; PL 7, FIciS 50-51; PL. 8, FIGS

52-53.

Haplophragmium inetsim Stache, 1864: 164, pl,

2I. fig. 1.

Haplapragrium maericum Stache, 864) 166, pl.

Tl. fig, 2.

Cyclommind paupera Chapman, 1904: 229 (in

paupera Chapman, (904, yense stricto). Rag-

paupera Chapman, t904, sensu stricto).

Cyclarnming ineisa (Stache) Chapman, 1926: 29,

pl. 2, fig. |. Chapman & Crespin, 1932: 14, pl.

14. fig. &. Parr, 1938: 89, text fig. 1. Crespin,

1950; 72, pl 10. fig. 3. Raggatt & Crespin,

1958: pl. 7, fig. 3. Hornihrook. 19612 30;

Hornibrook, 1971: 34, text fig. 9, pl. 6, figs.

RAG). Cockbain, 1974; 107, figs. 67A,B,C.

Huplopheagmium canariense Chapman, 1926: 28,

pl 2, fig. 2 (not Nerionina — canarterisis

d'Orbigny, 1839).

Cyclummina lonelcompressa Chaproun & Crespin,

1930; 97, pl, 3, figs 3, 4.

Haplophragmoides of, incisa (Stache)

1965: 150, figs 2d, 3 (3u,b, fab).

Haplophragmeides cf paupera Taylor, 1965: 1ST,

fig, 4 (2a, Zh) (not Cyelammina patipera

Chapman, 1904).

“Cvelarmmina” cf. inviva (Stachbe) Lindsay & Bon-

nett, 1973; 33, pl. I, fig. 4,

Cyelummina ef. ineiva (Staebe) Quilty, 1974, p.

33, pl. L, figs 1-3 (in part),

Holotype: Slide 64, Naturhistorisches Museum,

Vienna (Hornibrook 1971, p_ 25).

Type Jocalitv: Grid reference N6da/483465

(1948 ed.) Department of Lands and Survey,

NZMS 1, Te Kopapa Point, Raglan Harbour

(Whaingaroa), North Island, New Zealand,

Whaingaroa Siltstone, Whaingaroan (Early

Oligocene) (Hornbrook 1971, test fig. 1, pp.

9-10).

Materials 702 specimens, of which 390 were

measured and LO sectioned, from the following

localities; Outcrops—topotypes, Raglan Har-

hour (18). Demon's Bluff (98)., Mogg’s Creek

Taylor.

EARLY TERTIARY CYCLAMMINA AND HAPLOPHRAGMOIDES 189

(6%). Custle Cove (76), Fossil Bluff! (1),

Abbotsford, New Zealand (2). Boreholes—

O.D.N.L. Mt Salt No. 1, 719-1115 m (295)3

O.D.N.L. Mt Salt Structure Hole No, 1, 226-

229 m (1): S.E/0.8. Beachport No. 1, 241-

271 m (16h: B.PNLL, Geltwood Beach Nw, 1,

238-277 m (4); E, & WS. Kingston No. 3,

65-69 m (2). BE. & WS. Coonalpyn No. |

69-70 m, 105-107 m (2); SAM D. Waikerie

No. 2, 149-152 ny (20); S.A.M,D. Waikerie

No. 28W. 146-148 nt (1): Water Bore Plan-

tazenet 5666. South Stirling 11-88-21.03 m

(92), King's Park Na. 2, 216-219 m (4),

Parish of Bumberrah (Melung}, 394.7 m (3)

Description: Adult test of moderute to hinge

sive, composed mainly of quarte grains, with

Usiially 4 whorls ia the nyierosphene form, 2 in

the inegaspheric form: 7 to 15 ehambers in

the last whorl, plunispiral, biconvex, invalute,

more Or less compressed at the periphery

which is very slightly lohulalte, depressed

around the rather shallow umbilicus, sutures

inetsed, straight to slightly sinuate,

Wall finely agglutinated, thick, epidermis

thin, fine-grained, smoothly finished in line

sediment but varying according to the cuarse-

ness of the rnatris, of uniform texture but for

seallered large quart? grains, imperforate:

hypodermis thick, coarse-grained, coarsely

alveolie, not labyrinthie, the alveolae consist-

mig Of relatively simple tubes with a single

conspicnous opening inte the chamber lumen

and bifureating just below the epicermis,

which they do not penetrate, The distal ends of

the alyeolae are frequently seen through the

translucent epidermis of well-preserved speci»

mens. Alvealae lined with organic material

ipseudochitin or “teetia’, a combination of

protein and carhehydrate (Hyman 1940)),

and the test is probably held together with

orfanic cement,

Aperiural face moderately rounded to

roundly ogival, covered with coarse quartz

grains between which fine supplementary arcal

apertures are sometimes visible. Aperture an

intdriomuarginal narrow slit at the base of the

apertural face.

Seplil wall thick, ahoul as wide as the inter

septul width of the chamber lumina, oacea-

sionally perlorated by the areal upertures.

Dimensions: Holotype, diameter 1.44 mm,

thickness (1.90 mm (Hornibrook 197)). Of

390 specimens measured, diameter U.28 to

30 mm, megasphenc 1,12 mm, ivertige 1,15

miro; thickness O15 to 1 75 mm. averave 0.54

mm: overage ratio diameter ~ thickness 2.13) 1.

Diameter of early chambers 1.0 to 1.5 mm

(microspticric), Number of whorls. in carly

chambers (microspheric form) 3.

Remarks: C. fneisa is a ubiquitous qnd long-

ranging species with considerable variation in

shape from fairly fal, inostly due fo compres-

sion, with a diameter 2 thickness ratio of 4.5>1,

to the robust form with a ratio of 1,6: 1. Most

of the topotypes from Raylun Harhour kindly

Tent hy the New Zealand Geological Survey

are rather fattened as compared with the holo-

type and matched topotype (Ilornibrook 1971,

p. 35, pl. 6, figs 88, 91). The specimen NZGS

Reg. No. FP2078 sectioned by Hornibrook

(1971, text fig, 9) Appears to be a megaspheric

farm in Which the chamber limina have col-

lapsed, us Hornibrook suggested. The same

type of ulveolir pattern is shown i) the section

of the relatively poorly preserved megaspheric

specimen GSSA Fred2 (pl 3, fix. 35) and tw

CC. precaueellar Voloshinova as fivured by

Volashinova & Budasheva 1961, pl. 15, figs 4,

Sand Muyluert 1966, pl 42. figs 1-6. Well-

preserved specimens tpl. 3, figs 31-33) clearly

show an alveolur pattern and relutive disposal

of chamber lumina and septal wall of the same

shinova of Muylaert (1966, pl. 41, figs, 1--75 pl

42, figs. 7-9; pl, 43, fiz. 1). The section of C.

praecancellale illustrated by Cicha & Zapleta-

lova (1966, pl. 38, fig, 3c) shows w vonsider-

ahle reduction of the chamber lumina ahd jiere

complex alVeolur structure which is not readily

comparable with that of C. inetsa

The reeard of the species in the Late Cre-

taccous Curdies Formation (Ludbrook 1971,

fig. 3.3) is based ion probable contamination of

cuttings in Mt Salt Nu. | Well from the over-

lying Dartriaor Portation,

Distribunon: Widespread in southern Australia

and New Zealand from Cate Palacocene to

Marly Miocene.

Cyclammmina otway ensis rsp.

Pi. 2, FIGS 23-25: PL. 4, FIGS 38-40

Maplophragmoides canaricise Chapman, (904:

22M, ple 22, fiw. 2 (nor Nowionine canariensiy

WVOrbigeny, 1839).

Haplophragmoides complanata Taylat, (965; (48,

fies 2 fac). MiNi. (2) not Cyehimmina come

pPlanata Chapmun, 1904).

Holotype: GSSA F608,

GSSA FIqo9, FIG10,

Type locality: Glenelg River, Victoria, 2.4 km

downstream from Killura Bridye. 14 km SW

figured paratypes

FARLY TERTIARY CYCLAMMINA AND HAMLOINRAGMOIDES 1

of Cystertan, MAMILTON 1:250000 geo-

logical map sheet, 37°39'S6"S, 141°1723"E,

Gambier Embayment, Gtway Basin, Dartmoor

Formation, Palavocene to Eurly Eocene,

Material: The holetype, figured paratypes and

approximately 650 paratypes, of which |72

were measured and 4 seciioned, from: Qut-

crops-—Brown's Creek (3), Glenelg River

(43); Boreholes—V,M.D. La Trobe No, | at

298.7 m (500); SAM.D, €.G.9, County

Grey, 28.96-29.26 m (63): O.D.N_L. Mt Salt

No. |, 385-960 (cavings 1039-058 m) (23);

S.E.O.8, Beachport No. 1, 311-546 m (25),

Description: Adult test small. moderately in-

flated, composed mainly of fine quarte grains

wilh scattered medium grains which have a

tendency to be arranged radiully, planispiral,

biconves. involute, with 3 whorls in the micro-

spheric form, 24 in the megaspheric form, 7

to [1 chambers in the last whorl, hul wsnally

8 to 9 in the adult form, sutures moderately

well-defined, not incised, straight; periphery

roinded, only slightly or not lobulate, umbi-

fous well-defined and relatively deep,

Wall rather Joosely agglutinated, thin, with

Very little cementifg material, epidermis rels-

tively thick, smpoth or roughly — finished

according tu the coarseness of the grains of

the cnelosing sediment, imperforate; hypo-

dermis thin, simply perforated hy conspicuous

alveolae having a large opening into the

chamber lumen and branching distally from

the chamber lumen to the epidermis which

they do not penctrate but through which the

distal ends can be seen in 2 or 3 radiating

series per chamber, Septal wall moderately

thin and Occasionally punctured hy the sup-

plementary apertures. Apertural face a

moderately high rounded arch covered with

medium quartz grains between which very

smal] suipplementiry arcal apertures are occa-

sianally visible.

Aperture a well-detined interiomarginal slit

at the base of the apertural face, sometimes

with a slight lip on the absutural side,

Chamber lumen widely open.

Dimensiony, Holotype Ff608, diameter 0.75,

thickness 0.41 mm; paratype PIG09, diameter

0,85, thickness 0.42 mm; paratype Pf610, dia

meter 0.80, thickness 0.40 mm, OF |72 speci

mens measured, diameter 0,35 to 0.90) Min,

average 0.64 mm, thickness 0.12 to 0.42, ayver-

jive O30 mm; jverave ratio diameter + thick-

ness 201,

Remarks: Chapman (1904) identified this

species as Haplophragmium — eanuriense

(@Orbieny) (= Nonioninad — eanarlensis

WOrbigny, the type species of Haplophraz-

moles baie 1910) and figured a speci-

men (pl, 22, fig. 2) which suggests the pres-

ence of ulveolue below the epidermis. The

figured specimen is, unfortunately, not the one

now on Chapman's slide NMV P26049, but on

square 33 of a slide GSM 64828 with mounted

speeumens from Chapman's material very

kindly made available by D, J. Taylor, there

are four specimens almost identical with the

one figured by Chapman, These belong to

Cyclanmina as the open ends of alyeolac can

be seen on the inside of the chamber wall of

one partially dissected specimen, Taylor

(1965, p. 157) appears to have overlooked

Chapman's separation of the species from CL

complanata and interpreted the specimens on

square 33 (GSM 64828) a8 Haplophragiioides

complanwa He figured as “preservation

stages” (p. 146, lig, 2a, h, c) three specimens

from La Trobe No. I Bore at 298,7 m, which

are here reinierpreted aller examination of

the specimens as at) untogenetic series of C.

otwavensis, GSM 60464 (Pig, 2a) is a half

seclion or “grind” of a pyrite-filled immature

specimen, GSM 60465 (Fig. 2b) is a partly

dissected specimen showing the open chamber

lumina and relatively simple alveolar hypo-

dermis; and GSM 60466 (Fig. 2c) is a half

section or “grind” of a pyrite-filled specimen

on the reverse side of which the alveolar open-

igs beneath the epidermis ate visible. GSM

60466 is similar to the specimen GSSA

Ff609 figured on plate 2, figure 24,

The holotype is a rather cog rse-grained

specimen sclected from a sample taken from

PLATE 7

Fig. SO. Cyclurnenine incisa (Stache), NZGS F100894. Demon's Bluff, Demon's Bluff Formation,

Late Eocene, longitudinal view of alveolae from chamber lumen through hypodermis to epi-

dermis, X550,

Fig. SI,

Cyclarmina incisa (Stache), NZGS F100937, Demon's Bluff, Demon's Bluff Formation,

Late Eocene, distul ends of alveolae Viewed from eAdlerior Uirouph ruptured epidermis,

Most.

EARLY TERTIARY CYCLAMMINA AND HAPLOPHRAGMOIDES 193

the Darunoor Formation on Glenely River

because of the accessibility of the outcropping

material. The species is very abundant in the

Palaeouene to Early Eocene of the Dilwyn

Formition in La ‘Trobe No. 1 Bore, and com-

mon in the type section (Bore CG9) of the

Kurrungule Member of the Tartwaup Forma-

oon,

The specific name is taken from the Otway

Basm to which if ts restricted an present know-

ledge.

Distribution: Otway Basin-Dilwyn and Dart-

moor Formations (Palaeocene to Early

Eocene) and lower pan of Johanna River

Sands (?Palacocene to Early Eocene); Bur-

rongule Member of Tartwiup Formation

(Middle Eocene).

Cyclammina paupera Chapman

PL, 2, FIGS 21, 22, 26, 27; PL, 4, FUG, 37

Cyclamming paupera Chapman, 1904: 229, pl. 22,

fig. 6,

fHluplophragmoides panpera (Chapman) Taylor,

1965: 151, fie 4 (hab) (not Cyelanimina pau-

pera Crespin, 1950:72, pl 10, fiz, 4).

Holotype; SMV Slide P26049 No. 5,

Type locality: Brown's Creek, between Rot-

ten Point and mouth of Johanna River,

13,6 km NW of Cape Olway, 38°46'22"S,

143°23°14°R, base of Johanna River Sunds,

*Palatocene to Early Eocene.

Material: 86 specimens of which 52 were

measured, From outerops— Brown's Creek

(3), Glenelg River (7); from boreholes—

V.M.D, La Trobe No, 1 293-299 m (63),

V.M.D Wangoom No. &. 596-4600 m (49),

0.D.N.L. Mt Salt No. 1, 902-957 m (63),

Description; Adult test small, planispiral, flatly

biconvex, imvolute or slightly evolute, com-

posed of fine quarly grains, compressed ta-

wards the periphery which is ucule, rounded

or slightly lobulate; umbilicus well-defined,

deep. Six fo twelve chambers in the last whorl,

but usually 6 to 10 in the adult form: sutures

meised, straight or slightly arcuate.

Wall finely agglutinated, thin, epidermis very

thin, with fine ulveolae visible through the

translucent epidermis when the specimen is

wel, otherwise appurenily absent or ponrly

developed. Septal wall thin. Chamber lumen

widely open, subtrapezoidal in section, each

chamber tends to be set almost above the cor-

responding chamber of the previous whorl, so

that both chambers aud septa appear in equa-

torial section as unbroken radii (pl. 4, fig, 37).

Aperlural face high, narrow, subtriangular,

covered with fine quartz grains; uperture a

very narrow jnteriomarginal slit at the base of

the apertural face, without lip,

Dimensions: Holotype diameter 1,08 mm, Of

52 specimens measured diameter 0.37 to 1.08,

average 0.52 mm); thickness 0,12 to 0.25, aver-

age 0,20; average ratio diameter > thickness

2.6: 1.

Remarks: In texture of the epidermis, in size

and general shape, C, paupera may be com-

pared with C. yranget Finlay, which has a

similar stratigraphical range of Late Palacocene

ta Middle Bocene (Hornihrook 1968, p, 48).

€_ paupera is flatter and less clevated around

the umbilicus, and in C. prangei the alveolar

pattern as indicated by the distal ends of the

alveolae visible through the siliceous wall is

much coarser and mare strongly developed,

The specimen (CPC 645) figured by Cres-

pin (1950, pl 10, fig, 4) is # rather deflated

example of C. incisa, not unlike topotypes

(rom Raglan Harbour. The specimen figured as

Haplophragmoides sp. A (Taylor, 1964, p.

563, pl 9, fig. 3) and placed in synonymy

with Cyelammina paupera (Taylor, 1965, p,

15]) appears tu be a Haplophragmo@ides and

Late Cretaceous has not been confirmed,

Distribution, Otway Basin—-Dilwyn and Dart-

moor Formations (Palaeocene to Early

Bocenc) and lower part of Johanna River

Sands (?Palacocene to Early Eocene),

Cyclammina rotundata Chapman & Crespin

PL. 1, FIGS 9-12, 14-17, IPL. 3, FIG. 36

Chapman, 1904:

(not Neniforine latidersata

latntorsatim

fiz. |

Naplophragmium

227, pl, 22,

Bornemann),

Cyclammina retundaia Chapman & Crespin, 1930:

96, pl. 5, figs 1, 2- Crespin, 1950: 72, pl. 10,

ligs Sa, Sb. Raggatt & Crespin, 1955: pl 7, figs

Sa, 5b.

PLATE 8

Fig. 52. Cyclammina ineisa (Stache), NZGS F(00937_ Demon's Blulfl Demon's Bluff Formation. Late

Bogene, ulveolue viewed Cram chamber lumen, X50. ;

hig, $4, Cyclammina incisa (Stache), NZGS F100937, Demons Blatt, Dernons Bhifl Formation, Late

Focene, epidermis. 2500,

194 N. H. LUDBROOK

Maplophragenaides retoadain (Chapman & Cres-

pin) Taylor, 1965; 153, Nie 4 (4a. bh,

Cyelaninoia of. fiewds (Stiche) Quilty, 1974) 33,

pl. ty figs 1-3 Cin part ut least).

Nalolype; CPC 14.

Type localityy No, | Bore Parish of Bum

berrah, Metung., Victoria, BAIRNSDALE

1:250 000 geological map sheet, 37°53°34"S,

147°50°14°E, at 394.7 m depth, Gippsland

Basin, Lakes Entrance Formation, Oligocene.

Material’ The holotype and 107 specimens of

Which 93 were measured and 5. sectioned,

From outcropsDemon’s Blull (4), Castle

Cove (5). Fran: boreholes—O.D.N.L. Mt Salt

No. 1 908-1015 m (22), S.E.0.S. Beachport

No. [ 238-271 m (41); New Morphett Street

and Victoria Bridges, Adelaide, Bore 11 25—

25.6 m (2), Bore 12 20.1-20.4 m (1), Bore

14 16.7 -17,0 m (2); “Carelew” Bore 2 19.8-

19.9 m (1), Observation Bore F, Port Gawler,

253-254.5 m (2), Waikerie Bore 28W, 146m

(15), Plantagenet Location 5666, South Stir-

ling, LE.88-21.03 m (33).

Deseviprion; Adult test of moderate to large

size, inflated, planispiral, biconvex, involute,

composed mainly of coarse quartz grains, with

4 whorls in the microspheric form. 6 to 12,

rarely (3, chambers in the last whorl, periphery

rounded, umbilicus searcely or not depressed,

sutures. poorly defined, straight,

Wall agglutinated, thick, epidermis thin and

usually coarsely finished with numerous agelu-

tinated course grains, imperforate; hypodermis

thick, coarse-grained, alveolar, with a pattern

of fine roughly parallel! alveolac with only a

slight tendency fo bifurcate distally beneath

the epidermis. The distal ends of the alveolae

are not usually visible through the epidermis,

Septal wall thick, chamber lumen much

reduced,

Apertura] face a low arch covered with

course quartz grains hetween which small sup-

plementary areal apertures are frequently

visible. Aperture a slit at the base of the aper-

tural face, short and well open in inmmature

specimens,

Dimensions. Holotype diameter 1.4, thickness

0.82 mm, OF 93 specimens measured, diameter

05 to 2.12 mm, average 1.13 mm, thickness

0.30 to 1.50 mm, average 0.73 mm, Average

ratio diameter ; thickness 1,55;1.0,

Remarks; The specinven figures as MMaplo-

79, fig. 5) and placed m synonymy with

Cyelammina rotundata (Taylor 1965, p. 153)

uppears to be a Maplophragmoides and net C,

ratundata, he occurrence of which in the Late

Cretaceous has not been confirmed. The secord

of the species in the Late Cretaceous Curdies

Formation (Ludbrook 1971, fig. 3.3) is based

on probable contamination of cuttings in Mt

Salt No, | Well from the overlying Datimoor

Formation,

Distribution, Widespread in Austilia and New

Zeahind, associated with CL imeiva, Fram bade

Palaueoeene to Oligocene and possibly Purly

Miocene.

Acknowledgments

For the Joan of specimens and fur permis-

sion to examine material in thelr custody I

am most grateful to Mr D, J, Taylor whe

generously made available to me duplicate

material from the Port Campbell Embayment;

to Dr N, de B. Hornbrook of the New Zea-

land Geological Survey, the Assistant Direetar

(Geology) and Dr D. J. Belford of the Bureau

of Mineral Resources, Canberra, the Director

and De C. Abele of the Geological Survey of

Victoriv, the Assistant Director of the National

Museum of Victona, the Director and Mr

G. W. Kendrick of the Western Australian

Museum, the Direetor and Dr C. G, Adams of

the Britsh Museum (Natural History), Dr

P. G. Quilty of Macquarie University. For the

gift of material Tam indebted to Mr und Mrs

W. T. Grocock of Mount Barker, Western

greatly profited from discussions with Dr VT

Banner at the University College ef Swansea,

United Kingdom, and Dr M. Hamaoui of

CLN.P.A., Pau, France. Dr PLN. Webb, tor-

merly @f the New Zealand Geological Survey

and now of Northern Ulinois University.

U.S.A. generously provided the SEM photo-

graphs of Plates 5 to 8 Dr Abele critically

reud the manuscript and provided essential

data,

The study was done at the Department of

Mines, South Australia, and 1 wish to thank

the Director of Mines and the Division of

Biostratigraphy of the Geological Survey ot

South Australia for making facilities available,

lor photographic assistanee, and for permis-

ainn to publish the paper. The figures were

drown in the Drafting Branch of the Depart.

ment.

EARLY

TERTIARY CYCLAMMINA AND HAPLOPHRAGMOIDES 195

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A NEW SPECIES OF DIPORIPHORA FROM SOUTH AUSTRALIA AND

GEOGRAPHIC VARIATION IN D. WINNECKEI LUCAS & FROST

(LACERTILIA: AGAMIDAE)

BY TERRY F. HOUSTON

Summary

A new species of dragon lizard, Diporiphora linga, is described from western South Australia. It is

closely related to D. winnecki and, in order to facilitate comparison, the geographic variation of

winnecki is briefly reviewed with the recognition of two distinct races. Notes on the habitats of the

two species are included.

A NEW SPECIES OF DIPORIPHORA FROM SOUTH AUSTRALIA AND

GEOGRAPHIC VARIATION IN D. WINNECKEI LUCAS & FROST

(LACERTILIA: AGAMIDAE)

by Terry F. Houston*

Summary

Houston, ‘T. F, (1977) A new species of Diperiphera from South Australia and geographic

variation in 2, winneekei Lucas & Frost (Lacertilia: Agamidae). Trans, R. Soe. 8. Aust.

101(8), L99-205, 30 November, 1977.

A new species of dragon lizard, Diperiphera linga, is described from Western South Aus-

tralia. It as closely related to D, winneckei and, in order to facilitate comparison, the geographic

variation of wivnecked is briefly reviewed with the recognition of two distinct races. Notes on

the habitats of the two species are included.

Introduction

Specimens of an undescribed species of

Diporiphora Gray were first collected in 1921

at Immarna, $.A., on the Transcontinental Rail

Line and were identified us D. australis (Stein-

dachner) by Kinghorn (1924), These speci-

mens were apparently the basis of Cogger's

(1975) record of D. reginae Glauert from

western $8.A, Storr (1974) referred other speci-

mens of the species to D, winneckei.

My recognition of the new species followed

extensive ficld studies when many live speci-

mens were examined, from a study of speci-

menos of D. reginee, most of the ‘winneckel’

listed by Storr (1974) and additional material

in the Australian and South Australian

Museums. These studies also revealed that win-

neckei comprises at least two geographic races

and these are briefly defined in order to allow

comparison with the new species.

The following abbreviations of the names of

jhsututions or collections are used:

ABG Allen EB. Greer collection (presently

in AM)

AM Australian Museum, Sydney

SAM — South Australian Museum, Acelaide

WAM Western Australian Museum, Perth

Diporiphora winneckei Lucas & Frost

FIGS 1-5

TYPICAL EASTERN RACE

Diporiphora winneckei was described from

Charlotte Waters, Northern Territory, a locality

near the S.A. barder on the western fringe of

the Simpson Desert. All specimens examined by

me from the Lake Eyre and Lake Torrens

Basins (i.e, east of broken line A, Fig, 1) agree

with the descriptions and figure of Lueas &

Frost (1895, 1896) and represent a discrete

race,

Features distinguishing this race from

western populations are: no pre-atal pores in

either sex; gular area with three bold longi-

tudinal stripes, the median stripe continuous

with a pair of stripes extending down the chest

and belly (Fig, 2), the stripes grey in males and

either yellow or grey and yellow in females;

head with dark dorsal markings (Fig. 3).

Variation is slight, The lizards all appear

extremely slender, almost emaciated, The gular

fold is weakly developed and sometimes absent

medially,

At each of several localities where | collected

specimens the lizards were on sand ridges in

or onear bushes of Sandhill Canegrass

(Zygochlea paradoxa). This leafless plant has

wiry tangled stems and grows in hummocks

almost exclusively on sand ridges. Tts range

includes the Simpson Desert and bordering

areas, and extends south to Port Augusta and

east to the Darling River, New South Wales.

Thus the known distribution of typical win-

neckei corresponds approximately to the range

of the canegrass.

WESTERN POPULATIONS

Specimens of wirtnecket from western N.T.

and the Northwest and Eastern Divisions of

* South Australian Museum, North Tce, Adelaide, 8. Aust. 5000.

200

ee a ar eee

a 2

TERRY F. HOUSTON

@o D. winneckei

* D. ?bilineata

Fig. 1, Known distribution of Diporiphora linga, D. winneckei and some specimens of doubtful identity.

Solid symbols = specimens examined, open circles — literature records. See text for further

explanation.

Western Australia (see Fig. 1) differ from

those of the typical race as follows: 1-2 pairs

of pre-anal pores in both sexes (though weak—

rarely absent—in females); gular area with 4—5

narrow longitudinal grey stripes (Figs 4, 5).

These western winneckei are far more

variable than the eastern race and, as I have

only examined preserved material, I have pre-

ferred not to formally establish a new race at

this time.

While the majority of specimens have the

same extremely attenuated form of the typical

race, some are outstanding in being more robust

with thicker necks and larger heads. To

quantify robustness in preserved, often dis-

torted specimens is very difficult, but the ratio

of head width to head length provides an

approximation, It was found that the frequency

distribution of this ratio was normal, so that

the robust and very slender specimens connect

A. NEW SPECIES OF DIPORIPHORA FROM &.A, 201

through an unbroken series of intermediates,

the latter forming the bulk of the population.

Additionally, the unusually robust specimens

come from widely scattered localities on the

Pilbara coastal plain, the Hamersley Plateau

and the eastern desert of W.A.

The gular fold is also subject to variation,

being strongly developed in the majority of

specimens but occasionally feeble medially or

absent. Absence of the gular fold occurred in

specimens from across the range of the race

(except on the Hamersley Plateau) but

occurred more frequenily in specimens from

the Pilbara coastal plain. Development of the

fold also appears to be independent of robust-

ness,

The vertebral stripe (usually distinct and

grey) is occasionally bull brown, faint or

obscure, or divided medially, The ventral

stripes had faded in many specimens but their

number (4 or 5) on the gular area appeared

to be independent of locality.

Some variation muy have resulted from

preservation, For example the gular area, or the

whole ventral surface of some specimens, had

a. satin-like sheen and a few had dark dorso-

lateral stripes instead of the usual pale stripes.

One specimen (WAM R30433) from 16 km

S of Port Hedland combined characteristics. of

the two races; the gular area had three bold

stripes as in the eastern race but two pre-anal

pores were present.

Although I haye no personal observations of

the habitat of this race, notes accompanying

many specimens reveal that they were caught

amongst porcupine grass (Triedia spp.) on

sand dunes and sandy flats.

Specimens examined

The mitevial listed by Storr (1974) from the

collections of SAM and WAM plus the addi-

tional specimens listed below.

Typical eastern race; Northern ‘Territory:

Charlotte Waters, AM 2143-5. South Australia:

44 km WSW of Anna Creek HS, SAM R14522A—

EF; 7.5 km ENE of Bopeechee Rail Siding, SAM

R13947A-C: Hunt Peninsula, Lake Eyre North,

SAM R14643; shore of Lake Eyre, Muloorina Stn,

SAM R14796; 22 km WNW of Moralana HS,

SAM R14529; 5.5 km WNW of Myrtle Springs

Figs 2-5. Patterning in Diperiphora wihneckei

(in life, the areas shown jn solid black

are either grey or yellow). 2, ventral

pattern of typical eastern race; 3, dorsal

head pattern of same; 4, 5, two varia-

tions of ventral pattern in western race,

202 TERRY F. HOUSTON

— eee

bai pea

Figs 6-7. Diporiphora linga: 6, adult male (black speckling on body is an artefact); 7, adult female in

life.

“\ NEW SPECIES OF DIPORIPHORA FROM S.A.

HS, SAM R{4532; 10 km ENE of Stuart Creek

HS.SAM R14530 A-E.

Western race. Northern Territory: 62 km W of

Ayers Rock, AEG 311, 342; 62 und 82 km NW

of Chika Well, ABG 203, 235, 13 km W of ME

(les, AM 49711) Palm Valley, SAM R5047; 1.6

km Woof Refridverutur Bore, SAM RLII68..

Western Australias Marandoo, Mt Bruce, WAM

RS27U3, RS2757-8; 59 Km N of Neale Junction,

AFG 454, Nita Downs Sin, WAM RS51996,

R52009, RSIOLL; Well 38, Canning Stock Route,

WAM R44196.

Specimens excluded frum OB. winnecket

Amongst the specimens which Store (1974)

listed under winneckei were several from the

Kimberley Division of W.A. und from Joanna

Spring. The localities of these specimens. lie

ubove broken line B in Big, 1, Having

examined these and additional specimens from

wearby localities 1 feel that they are more

likely to represent an isolate of D, bilineata

Crray than of winnecker They differ from win-

weckei as Collows: gular fold) consistently

absent, gular seules mucronate; gular area with

Sar 6 dark longitudinal stripes ane chest and

helly with 4 dark Jongitudinal stripes (faint or

absent in many specimens); a dusky grey or

black area above insertian of fare limb; dorsal

pattern, especially dark cross bars, strongly

developed in some individuals, virtually absent

in others.

Typical and western winneeke? occassionally

lack the gular fold ouUL never exhibit blackish

putches above the fore limbs. However, such

patches ure characteristic of other species (ee

lalliae and hennerti),

‘The specimens of this unplaced form which

f have examined are the following (all in

WAM lnless indicated otherwise): Beagle Bay

Mission. R4G463: S81 miles (130 km) E of

Kroame, R36336: Derby (and ‘presumably

Derby’), R20262-4, R20317-29. R26834,

R4666). locujan Creek, La Grange. R27638;

Jounau Spring, SAM R1I4320 A-B, R146217 La

Grange, R46216; Point Coulomb, R40266.

Diporiphora tinga n, sp.

FIGS 6, 7

Holornpe: 3, SAM R1ISG20F, 23 km N of

Koonibha Mission, SS, Aust, 3427S,

133° 26°F, (I-VRALIY7S, collected amongst

fussucks of Triodia by C. & T. Houston, A-

Edwards and J. Herridee.

Diagnosiy: Morphalovien|ly and chramiticully

much like OD, winneckes differing as follows,

Mates always wud lemiades uswally with pre-anal

203

pores (absent in typical w/nneckel) ; back with-

out a preyish vertebral stripe and its dark cross

bands reduced to irregular spots alang dorso-

lateral stripes or virtually absent: ventral sur-

faces never longitudinally striped, white to

pale grey offen with weak ocelli on belly and

base of tail; males with bright pink flanks and

rump during spring (no similar coloration

recorded in winnecke?); form more robust than

typical winneeke? with thicker neck and fatter

bouy; tail, on average, relatively slightly shorter

(cf 227% ta 253% of SVL). Also very like

OD, reginae but of smaller size and not quite as

robust, lacking femoral pores, dorsal scales

more weakly keeled ynd not mucronate, belly

usually acellute and pink wash of spring mules

extending along full length of flanks, not just

on sides of tai) base,

Description

Small dragon lizards reaching a muximum

snoui-vent length (SYL) of SI mm in males

and 61 mm in females; maximum total length

195 mm} form slender (Figs 6,7).

Relative dimensions (expressed as %), Head

length: SVL, 25-37 (mean 30. n = 49); hind

limb Jength; SVL, 58-86 (mean 73, 9 = 47);

tail length: SVL. L6S-271 (mean 227, n =

47); head width: head length (measured from

Up of snout to angle of jaw). 58-89 (mean 68.

m= 48),

Dimensions of holotype (in mm): SVL, 48;

head length, 16: bead width, Uh: iad length,

106; hind limb length. 36.

No nuchal or dorsal crests, nor post-

auricular folds ov spines bul gular ond scapolar

folds well-leveloped; dorsal scales longi-

tudinally keeled but oot muvronate; scales of

buck homagen¢ous, the keel lines more or less

parallel or Weakly converging posteriorly:

scales of funks weakly keeled to almost

sniouth, the keel lines negrer forelinibs. con-

verging with dorsalateral stipes but pyuraltlel lo

them further back; chip and gular scales vir-

tually smooth; chest and belly scales weakly

longitudinally keeled and apically mucronate;

23-30 (mean 26, u ~ 48) subdipital lamellae

on 4th foe; usually 2 pairs of pre-anal pores,

wecisionally 1 or 3 pairs, rarely absent; no

femoral pores.

Celoralion ef Jemale; Dorsal ground colour

pale buf, greyish Brown or olive brown; head

patternless dorsally; a whilish stripe from eye

to top of ear margined above and below with

dark brown) @ qarrow dark brown line trom

eye extending forwards to snout through

204

nostrils uppee and lower hps whitish, no verte-

bral stripe; a pair of narrow white or yellow

dorsolateral stripes extending along back from

nape, converging an base of fail and continuing

for {/3 to 4 its length before merging inty

ground coloration; uswally 6-9 irregular dark

brown spots along inner edge of each dorso-

jateral stripe on back, merging on base of tail

to form a dark line separating the pale dorso-

lateral stripes; usually corresponding — but

smaller spots on flanks helow dorsolateral

stnpes! an irregular pale midlasteral stripe often

present: flanks speckled with dark brown and

dappled with whitish spots; limbs uniform

above or brownespeckled: gular area Usually

white: belly anc lower half of tail base white to

pale grey, commonly with dark-cdged white

flecks or distinct ocelli.

Coloration of male: Similar to that of female

except that dark spots along dorsolateral

stripes are reduced or obscure; in spring a dif-

fuse bright piuk wash extends from shoulders

along flanks to sides of tail and onto region of

rump and thighs. and dorsolateral stripes

become rich yellow.

Habitats The lizards inhabit Trivdia hom-

mocks in mallee Eucalyptus covered sand.

dunes. They regularly bask in the crowns of

the bushes and retreat into the dense growth if

disturhect. Most specimens have been encoun:

tered on the slopes of dunes bul a few were

observed on bard sand flats between dunes,

Distibution: Throughout a belt of sinairicge

country extending from near Maralingy,

through Ooldea, south-castwards almost as far

as Wirrulla and the Gawler Ranges on northern

Eyre Peninsula, S.A, (Fig. 1),

Eryntwloey: The specific epithet is a Pitjantjat-

jira aborivital wore meaning “litle lizard” and

45 used us. NOUN IN apposition,

Specimens examined {all localities in SAL:

Paratypes: 22 km E of Barton Rail Siding

e3y°3a'S, 132°42 2), SAM RIL4976 A-) Bates

119 &m W of Barton), SAM RI4855; [21 km N

of Ceduna, SAM R14459 A—B; Immarna (or 407

miles, F-W Jine}, AM S4624-7: 7 km W of

Immarna Rail Siding (30°29°S, 132°05°E), SAM

Ri4v98 AL; sanie data as holotype. SAM

RPSU2) A-BG, 1: 6.7 kim SW of turn-off to Lake

Everard LIS ov Wirrolla-Kingoonya road, AEG

431; Maralinga bomb site, SAM R14447: 8 of ML

Finke (39°4U°S, 193°S8'E und 31°I5'S, 134"00'R),

SAM RIS6OD-T: Watson (obviously im error),

SAM R10822-6; 28-29 km NE of Wirrulla, SAM

RISi74, Riss AG, RIS220: 23 miles (47 kn)

ENE of Wirrulla, WAM R24529- 90),

TRRRY PL HOUSTON

Discussion

The relative structural homogeneity of

Dipariphora species requires that maximun

usc be made of colour patterning (including

Ventral patterning and transient sexual colours)

for the purposes of their separation, However,

because of the tendency of parts of the pattern-

ing to fade after preservation mt is highly

desirable that any further taxonomic studics of

the genus be based on field studies of live speci-

mens.

While my studies ot preserved material could

find no basis for separation of western win-

necke? into Wo or More races, more specimens

are required, especially from the Hamersley

Plateau, to provide a clearer picture of varta-

tion. More material is alsa required from the

central southern region of N.T. to show how

closely the ranges of the eastern anc western

races approach each other.

Several species of Diporiphora are now

known to be closely associated with hummack

grasses! reginae, lalliae, linea and western wirn-

neckei with Triodia and typical winnecked with

Zyvochloe, Vhis pronounced habitat specifleity

has doubtlessly played a major role in cvolu-

tion of the species. Populations. of Jinga are

isolated from those of other hummock-dwelling

species by extensive tracts of unsuitable habitat,

In the west is the Nullarbor Plain and across

the north are vast arcas of mulga/tussock grass

ynd chenopod shrub habitats. The ventral pat

terning differences between the two races of

winnecke? are probably related to differences in

the plants inhabited. The ventral striping

probubly serves to camoullage the lizards while

they are perched amongst the straight leaves

or stems of the grasses and fewer bolder stripes

in the gastern race would suite the courser more

open growth of Zygechtoa whereas more

numerous narrower stripes in the western race

would suit the finer denser growth of T'rivdia.

Acknowledgments

For the loan of specimens in their care and

for information provaded 1 arm indebted to Drs

H. G_.Cogger and A. E Greer (A,M.), ond

G. M. Stort (WAM). LT also wish to thank the

following people for their energetic assistance

in collecting specimens in the field: My A. M.

Edwards, Mrs J, A. Forrest (nee Herridge),

SAM), Dr J. P. Jessop, Chief Botanist, South

Australian Merbarium, kindly provided useful

references conceming the distribution of Sand-

hill Canegrass,