VOL. 120, PARTS 1 & 2
31 MAY, 1996

Transactions of the

Royal Society of South
Australia

Incorporated

Contents

Bayly, I. A. E. _Inland-water calanoid copepods of Kangaroo, King and Eee Islands:
Biogeography and Ecology - - - - -

Robertson, G. B., Prescott, J. R. & Hutton, J. T. Thermoluminescence dating of volcanic
activity at Mount Gambier, South Australia - -

Davies, K. A. & Lloyd, J. Nematodes associated with Diptera in South Australia: a new
species of Fergusobia Currie from a fergusoninid and a new record
of Syrphonema Laumond & Lyon from a syrphid - - -

Campbell, E. M., Twidale, C. R., Hutton, J. T. & Prescott, J. R. Preliminary
investigations of dunes of the Gawler Ranges province, South Australia

Harvey, M. S. A revised systematic placement for sucerORT Rae Southcott (Acarina:

Hydryphantidae) - - - - - - - -
Barker, S. Seventeen new species of Castiarina (Coleoptera: Buprestidae) —-
Kolesik, P. A new genus and three new species of Srrmongies (Diptera) from

Olearia spp. (Asteraceae) in Australia - - - - -

Brief Communications:
Tyler, M. J., Barrie, D. J. & Walkley, R. W. First fossil record of the hylid pS: Litoria
raniformis (Keferstein) - - - - -
Hero, J.-M., Fickling, S. & Retallick, R. The tadpole of Litoria revelata Ingram, Corben
& Hosmer, 1982 (Anura: Hylidae) = - - - - - -

Obituaries:
Ladbrook, Nelly Hooper MBE, MA, PhD, DIC, FGS. - - - - - -
Edmonds, Stanley Joe BA, BSc, MSc, PhD, Dip Ed. - - - - - -

PUBLISHED AND SOLD AT THE SOCIETY'S ROOMS
SOUTH AUSTRALIAN MUSEUM, NORTH TERRACE, ADELAIDE, S.A. 5000

74
78

TRANSACTIONS OF THE

ROYAL SOCIETY
OF SOUTH AUSTRALIA

INCORPORATED

VOL. 120, PART 1

TRANSACTIONS OF THE
ROYAL SOCIETY OF SOUTH AUSTRALIA INC.

CONTENTS, VOL. 120, 1996

PARTS 1 & 2, 31 MAY, 1996

Bayley, I. A. E. Inland-water calanoid copepods of Kangaroo, King and Flinders Islands:
Biogeography and Ecology —- - - - : - - 4
Robertson, G. B., Prescott, J. R. & Hutton, J. T. Thermoluminescence aating of volcanic
activity at Mount Gambier, South Australia = - - © =
Davies, K. A. & Lloyd, J. Nematodes associated with Diptera in South Australia: a new
species of Fergusobia Currie from a fergusoninid and a new record of
Syrphonema Laumond & Lyon from a syrphid - - - -
Campbell, E. M., Twidale, C. R., Hutton, J. T. & Prescott, J. R. Preliminary fnivestieations
of dunes of the Gawler Ranges province, South Australia - “

Harvey, M. S, A revised systematic placement for Austrotrombella Southcott (Acarina:

Hydryphantidae) = - - = - ~ - - - - -
Barker, S. Seventeen new species of Castiarina (Coleoptera: Buprestidae) - -
Kolesik, P. A new genus and three new species of Cecidomyiidae (Diptera) from

Olearia spp. (Asteraceae) in Australia = - - - - ~ -

Brief Communications:

Tyler, M. J., Barrie, D. J. & Walkley, R. W. First fossil record of the tybe frog L Litoria
raniformis (Keferstein) - - -

Hero, J.-M., Fickling, S. & Retallick, R. The tadpole of Litoria revelata Ingram, Corben
& Hosmer, 1982 (Anura: Hylidae) - - - - - - -

Obituaries:
Ludbrook, Nelly Hooper MBE, MA, PhD, DIC, FGS, - - - - - -

Edmonds, Stanley Joe BA, BSc, MSc, PhD, Dip Ed. - - - - - -

74
78

PARTS 3 & 4, 29 NOVEMBER, 1996

Anstis, M. & Littlejohn, M. J. The breeding biology of Litoria subglanduiosa and L. citropa
(Anura: Hylidae), and a re-evaluation of their geographic distribution -
Dyson, I, A. Stratigraphy of the jeans ad Aruhna and Repet wing subgeups,
Adelaide geosyncline = -
Dyson, I. A. Stratigraphy of the Rocgarsacnede 4 Tent Hill Formation and Simmens
quartzite at South Tent Hill on the Stuart Shelf, South Australia-
Taylor, G. S., Austin, A. D, & Davies, K. A. Biology of the eucalypt ante aee fly,
Fergusonina flavicornis Malloch (Diptera: Fergusoninidae) and its
associated hymenopterans in South Australia, with a description of a new
species of Bracon (Hymenoptera; Braconidae) - : , ,
Bird, A. F Studies on the soil-inhabiting macigrete, secrpiirms ef, pseudohufelad,
from South Australia = - - - =
Kolesik, P. Rhopalomyia goodeniae, a new species of Cecidomyiidae (Diptera)
damaging Goodenia lunata (Goodeniaceae) in inland Australia - -
Nicholas, W.L. RKobustnema fosteri sp. nov., gen. nav. (Xyalidae, Monhysterida, hoe,
a common nematode of mangrove mudflats in Australia = - -
Smales, L. R, A redescription of Aspersentis zanchlarkynchi (Johnston & Best, 837
comb. nov. (Heteracanthocephalidae: Acanthocephala) - -

Brief Communications:

Baker, G. H. Seasonal activity of the earthworm, Gemascolex lateralis CengRerSe raat
in a Eucalyptus woodland in South Australia - -

O'Callaghan, M. G. & Beveridge, 1. Gastro-intestinal penaints of fel cats in i Norther

Territory - - - -s

Terrace, T. E. & Baker, G. H. Predation of earthworms by the land lal pune
sanguinea (Moseley) var. alba haem sensu at. 1981 (Tricladida:
Geoplanidac) - > - - - - -

Tyler, M, J. & Williams, C. R. Mass frog mortality at two localities in South Australia

trsent 15 Thansactions of rie Royal Sociery af South Austrafia, it 12!, pans | A 2, 30 May, 197

83

101

uy

155

INLAND-WATER CALANOID COPEPODS
OF KANGAROO, KING AND FLINDERS ISLANDS:
BIOGEOGRAPHY AND ECOLOGY

By I. A. E. BAYLY*

Summary

Bayly, I. A. E. (1996) Inland-water calanoid copepods of Kangaroo, King and
Flinders Islands: Biogeography and Ecology. Trans. R. Soc. S. Aust. 120(1), 1-6, 31
May, 1996.

Calanoid copepod identifications are provided for samples from 16 localities on
Kangaroo Island, 18 on King Island and 11 on Flinders Island. The number of species
found was five, three and seven, respectively. Conductivity data are given for most
localities. Species richness in relation to land area is tabulated and discussed.
Boeckella major is recorded from South Australian territory for the first time. The
occurrence of Hemiboeckella searli in temporary pools and amongst dense
macrophytes in lakes may be due to the absence of predators in young pools and the
difficulty encountered by predators in searching dense weed-beds in lakes. The
disjunct distribution of Calamoecia gibbosa is explicable on the basis of east to west
dispersal along a lowland plain during the Pleistocene when sea levels were low,
followed by marine inundation.

Key Words: Copepoda, Calanoida, biogeography, ecology.

Transactions of the Royal Society of 8, Aust. (1996), 120(1), 16,

INLAND-WATER CALANOID COPEPODS OF
KANGAROO, KING AND FLINDERS ISLANDS: BIOGEOGRAPHY AND ECOLOGY

by I. A. E. BAYLY*

Summary

BayLy, I. A, E. (1996) Inland-water calanoid copepods of Kangaroo, King and Flinders Islands: Biogeography
and Ecology. Trans, R. Sne. 8S. Aust. 12001), 1-6, 31 May, 1996.

Calanvid copepod identifications are provided for samples trom 16 localities on Kangaroo Ishind, 18 on King
Island and 1 on Flinders Island. The numbet.of species found was five, three and seven, respectively. Conductivity
data are given for most localilics, Species richness in relation to land area is tabulated and discussed, Beeckella
Major is recorded from South Australian territory for the first time. The oecurrence of Hemiboeckella searli
in temporary pools and amongst dense macrophytes in lakes may be due to the absence of predators in young
pools and the difficulty encountered by predators in searching dense weed-beds in lakes, The disjunct distribution
Of Calamoecia gibhosa ts explicable on the basis of east to west dispersal along a lowland plain during the Pleisingene
When sea levels were low, followed by marine ‘nundation,

Key Worps: Copepoda, Calanoida, biogeography, ecology:

Introduction

Following the glacial (and aridity) maximuny that
occurred in the Late Pleistocene at about 18 ka B.P.,
deglaciation, and the consequent rise in sea fevel
(Chappell 1978: Galloway & Kemp 1981), converted
several areas of land along the southern margin Of the
Australian continent into islands. The nature and fate
of the samples of the fauna of yreater Australia provided
by these islands 1s a matter of considerable interest.
Rawlinson (1974) studied this issue with reference to
the reptiles of Bass Strair islands and Tasmania and
showed that a whole series of islands in Bass Strait
and off South Australia became isolated during the
period 16.5 - 4.3 ka BP

Despite their small size and the ability of many of
therm to produce desiccation-resistani eggs, freshwater
calanoid copepods are widely acknowledged to exhibit
poor dispersal ability (Bayly & Maly 199]; Bandrescu
1990, Bayly 1995), As a consequence. the
biogeography of calaroid copepods is of considerable
interest, with dispersal playing a smaller role than has
hee suppased by (he mummerous workers who
simiplistically equate the possession of resting egas with
good pawers of dispersal (Bayly & Morton (978),

This paper aims to exanyine the relationship of the
calanoid fauna of three offshore islands (Fig. 1) with
that of mainland Australia and Tasmania and to
consider the role of historical and ecological factors
ih Observed differences and similarities.

+ Dupartinent ol Beolopy and Evolulioniry Biology, Monash
University Clayton Vie~ 3168,
Current address; Hd Belyraye: Hallam Road Belpre South
View Si)

Methods

Rach body of water was thoroughly sampled with
a zooplankton net oF mesh size 150 jan. Collections
were preserved in 10% formalin. Conductivity of a
water sample taken from the field in a polyethylene
bottle was determined in the laboratory with a
Radiometer CDM2e conductivity meter, Where the
K,, exceeded 5.0 mS em!, the conductivity value was
converted to a salinity value using the method of
Williams (1986). For the Ring Island localities, pH was
measured with a Metrohm E599 portable pA-meter.

With two exceptions, the Flinders Iskind tnealities
were sampled by the author alone at various Limes
between 1985 — 198, and by the author working with
a limnological team from 10-12 February 1993, The
Ring Island samples. with dne exception, were taken
by a team of workers including the author during the
period 2-5 December 1991, Wilh two exceptions, the
Kangaroo (simd samples were collected by the author
alone during the period 24 June - 3 July 1994,

Results

Physico-chemical and biological resulty are sumitiar-
ised in Tables 1 - 3. Five calanoid species were record-
ed from Kangaroo [sland, with only two species,
Boeckella triarticulaia and Calameecia elitellara,
occurring at those lovalities with a sulinity of 3.4 g 1!
or more. At the less saline localities, B major was
resuicted to Temporary Waters. Only three species were
fuund on King Island, and one of these, B pseudo-
chelae, occurred in the sole temporary water body that
was sampled, C° fesranica Was the only. species found
in Waters with a recorded pH ol less than 6.0. None
of the King Island waters included in the survey was
sdline. Seven species were recorded from Flinders
Island with C asmaniod most common, B sywmetrica
was the unly species common to all three islands.

2 I. A. E. BAYLY

TaBLe 1. Occurrence of calanoid copepods on Kangaroo Island.

Locality Sampling Permanency* Conductivity Salinity Species?
date [K,.] (gl) Bm Bs Bt Ca Cec
(mS cm?)
Dam 1 near Penneshaw" 20.viii.1991 P x x
Dam 2 near Penneshaw* 20viii.1991 P x
Waterhole edge Edwards Lagoon 30.vi.1994 T 0.21 Xx
Pond Roper Road 1.vii.1994 T 0,22 xX x
Lake at Karatta 28.yi.1994 SP 0.31 x x
Pond nr rush Lagoon 27.vi.1994 T 0.44 x
Pond south end Roper Road Lvii,1994 SP 1.23 x
Ditch east Kingscote Airport 27.vi.1994 T L71 x x
Small Grassdale Lagoon 28.vi.1994 SP 3.00 x
Big Grassdale Lagoon 28.vi.1994 P 3.65 Xx
Kaiwarra Cottage Pond 2.vii.1994 T 58 3.2 x Xx
Duck Lagoon Lvii.1994 P 6.0 3.3 x
Discovery Lagoon 27.vi.1994 T 9.0 5.1 x
Lake Ada 3.yii.1994 P 12.8 75 x
Murray Lagoon 3vii.1994 P 15.3 9.1 x
White Lagoon 27.vi 1994 SP 75.3 513 x

a. P = permanent; SP = semi-permanent; T = temporary

b. Bm - Boeckella major Searle; Bs = B. symmetrica Sars; Bt = B. triarticulata (Thomson),
Ca = Calamoecia ampulla (Searle); Ce = C. clitellata Bayly.

c. collevied by N. Frick

Kangaroo Is.

King Is. (}

\ Flinders Is.

Fig. 1. Map showing location of Kangaroo, King and Flinders Islands.

CALANOIDS OF KANGAROO, KING AND FLINDERS ISLANDS 3
TABLE 2. Occurrence of calanoid copepods on King Island.
Locality Sampling Permanency" Conductivity pH Species”
date 5 Bp Bs Ct
(mS crv!)
Poo! or Currie“ 8.xi.1963 T - - x
Meatsafe Lagoon 5,xi7.1991 P 0.56 3.8 x
Dead Sea 2. xi) 199] P 0.56 47 x
Lake Martha Lavinia 3.401.199] P 0.75 §2 x
Seal Rock Lagoon (North) 4,xi\ 199] P 0,96 7.9 x
Pearshape. Lagoon 2.xii 199] P 1.04 78 %
Lake nr Surprise Bay Homestead 4. xii 1991 SP 1.10 14 x
Lake opp. Pearshape Lagoon 2.xi1 1991 P 113 8.2 x
Pennys Lagoon 3. xii 1991 P 1.15 77 x
Granite Lagoon 5.xii 199] P 1,26 64 x
Pioneer Lagoon 4. xii 1991 P 1.46 78 x
Lake btn Denbys & Pioneer Lagoon 2.xii 1991 P 1.47 72 x
Denbys Lagoon 2.xi1.199L P 1.57 6.6 x x
Lake east end Pioneer Lagoon 4, xi).199] SP L80 8.5 x
Seal Rock Lagoon (Middle) 4.xi1.1991 SP 1.90 45 x Xx
Cask Lake 3. 11.199] P 1.96 82 x
Lake Wickham 3.xii 1991 SP 2.15 8.9 x
Lake Flannigan 3. xii 1991 SP 2.28 99 x

a. P = permanent; SP = semi-permanent, T = temporary

b. Bp - Boeckella pseudochelae Searle; Bs = B, symmetrica Sars; Ct

c. collected by M. J. Littlejohn.

|

Calamoecia tasmanica (Smith);

Locality Sampling Permanency® Conductivity Salinity Species”
date [K,«] (gl!) BmBp Bs Bt Hs Cg Ct
(mS cm")

Brodies Lagoon* May 1962 T x
Scotts Lagoon” May 1962 T x
Pond nr Sticks Lagoon 15,1985 T x
Pond (1) ar Killiecrankie Syi l988 T x
Pond (2) nr Killiecrankie 5,vi, 1988 T x
Reedy Lagoon 9x) 1988 P x x
Shag Lagoon 12.11.1993 T 1.7 x
Lake btn N & S Patriarchs 19.-v,1985 SP 2.2 x x
Small lake (1)

nr Singleton’s Lagoon 10.11.1993 T 24 Xx
Small lake (2)

nr Singleton’s Lagoon 10.11,1993 T 5.1 27 x
Sticks Lagoon 15.1985 T 123 71 x

a. P = permanent; SP = semi-permanent; T = temporary

b. Bm - Boeckella major Searle; Bp = B. propinqua Sars, Bs = B, symmetrica Sars; Bt = B. triarticulata (Thomson),
Hs = Hemiboeckella searli Sars; Cz = Calamoesia gibbosa (Brehm); Ct = C fasmanica (Smith).

c. calleeted hy W. D. Williams

Discussion

It is probably not valid on the basis of Tables 1-3
to attempt straightforward and unqualified comparisons
between the faunas of any two of the three islands;
complications could conceivably arise from differences
in season of sampling, year of sampling, ratio of
permanent and semi-permanent to temporary waters
and number of localities sampled. However, it is
important to note that in Australasia, calanoid copepods
are nearly always present perennially in permanent
standing waters (a few glacial or high altitude lakes

are the only exceptions) despite wide fluctuations in
population density (Bayly & Williams 1973). In any
large sample at least some males and egg - or sperma-
tophore-bearing females are present and the species
determinable. This means that yearly or seasonal
differences in sampling dates should not unduly
influence the assessment of the fauna of the permanent
waters, This leaves a residuum of problems. for
comparisons which, hawever, are not so great as to
preclude the examination of a number of general
features and trends.

The much cited island biogeography theory of

4 I. AE. BAYLY

MacArthur & Wilson (1967) would predict that a
positive correlation should exis| between the number
Ol species found within a discrete area and the size of
that area. With respect to non-marine calanoids in the
Australasian region, Table 4 indicates that across the
whole spectrum of six land masses there is only a very
(ough correlation of the sort predicted. Several
unamalies call for consideration and explanation
King Island and Flinders Island ditfer only slightly
in area bur che former apparently has less than half
the number of species found of; the latter, This
Jifference, 1Fa1 1s nof an artefact. is diffigult to explain
bul it may be significunt that native habitat destrucnon,
including the removal of vegetation, has preceded to
a gredier extent an King (sland than Blindery stand
Kangaroo [sland is about three and a half limes larger
than Flinders island but has fewer calanoids (if the
halobiont species C) clitellanz is omitted tt has unly
fuur species). [tas difficult nowailays to find a large
natural body of fresh water on Kangaroo Island,
Extensive Vegetation remoyal in the period 1945 - [955
and the consequent rise in water tables and mabilisation
of salt has resulted in the salinisation af several lowland
jakes thal were formerly fresh. Murray Lagoon
originally contained fresh water but today it is saline
(salinity 9.1 g 1! on 3 July 1994; Table 1). Several of
the likes on Kangaroo Island visited during the winter
of (994 were found ta be highly saline and were not
sumpled for that reason. Half of che natural fresh waters
that were located were very small and temporary in
character It is concejyable that species like Calamoecia
gibbosa and ©, lasmanica, which oceur in sett
eastern and soulh-wester Australia and typically
inhabit permanent fresh waters. no longer oeeur on
Kangaroo Island as a result of recent salinisatiin
New Zealand is about foor times large’ than
Tasmania but has fewer calanoids, However, during
the Oligocene, some two-thirds af the area of modern
New Zealand was covered by sea (Stevetis 1YX0)
feeckella symenetrica, which becurred on all thee
islands. und A trianiealara, which was found on
Kangaroo and Flinders Islands. are both common and

TAwhn 4, daar area und species nehness

Naone ot land mass Ata Number of
(hie) calanoid species”
King Csland 1200 3
Flinders Island 1,330 7
Kangaroer taland 4,400 4
Tasinania 67.300 8s
New Zealand 269.000 10
ANustralia T6RLOOD cs)

a, Belonging to the family Centropagidae and ecatrieted i
the penera Boevkella, Hemiboeckella and Culamvecia.

widely distributed species (Buyly 19924). The
occurrence of BL propingua on Flinders Island only
(Table 3) is consistent with the existing evidence (hal.
within Australia, this species is restricted to the fir
castern [ringe of the continent, previous Australian
records are from the east coast of ‘Tasmania and coastal
New South Wales. 8. major (Kangaroo and Flinders
Islands; Tables. | and 3) is characteristic of temporary
waters and has heen recorded previously trom New
South Wales, Victoria and Tasmania (Bayly !992a).
The present record from Kangaroo Island ts the first
from South Australian territory but it is likely that tlic
merely retiects.a lack of ifvestiyation in (his Stale of
the copepods of temporary ponds and pools. A
psendochelae (King Vsland; Table 2) is another
lemporary Water speciilisr previously noted trom
southern New South Wales, Victuria and Tasmania,
Calumorvia taxmanica (King and Flinders Islands) and
C aripulld (Kangarou [sland) are widely distributed
species known [rom south-easteth and suuth-westert
Australia (Bayly 1992a)-

Remiboeckella searli typically occurs in temporary
ponds and pools, but, as with the present record fren
Flinders {sland (Table 3), it also occurs ip littoral weed
beds in permanent ar semi-permanent waters. This
cammonality of occurrence is nel as incongruous as
it Fifst appears. Water permeating dense vegetation in
the littoral regaon of a permanent lake his an ecological
similarity io that in a shallow, temparary pool
(including those entirely devoid of vegetation) that is
not commoanly appreciated pamely the exclusion of
predator: It is well appreciated thal, ina newly formed
pool, flying insect predators such as notonectids may
tke Same time to arrive and, until this occurs, the
habitat may be largely predator-free. The fact that
Hemiboeckella characteristically occurs very carly in
pool successions (Bayly 19926) suggests a high degree
of predator susceptibility. However. as painted put by
Connell (1975) some prey species have evalved the
ability tr live in refuges. that the predator cannit invade
because the Aabitat structure is toe difficult tO search,
It is presumably for this reason that dense littoral
vegetation offers 4. sear/ia refuge from limnctic fish
and insect predators in lakes, H searli is widely
distributed, occurring in south-eastern and sourh-
western Ausiralia.

Calamoevia gibbosa has the most intriguing
distribution of all the Australian freshwater calanoids,
it o¢eUrS in Tasmania, on Flinders Tsland. along the
coastal fringe of south-eastern South Australia betwee
Mt Gambier and Salt Creek and on two granne
outcrops near Balludonia in Western Australia (Bayly
1984. 1992a)_ A previnusly unpublished record js froin
near Mt Rough to the south-east of Salt Creek in South
Australia. The two Western Australian populations
were (reated (Bayly 1979, 1992a) as belonging to a
scparate subspecies from the easiern form.

CALANOIDS OF KANGAROO, KING AND FLINDERS ISLANDS ‘

The most rcasonable explanation for current
disjunctions im the distribution of C gibbosa is that,
at the time of one of the three glacial minima during
the Late Pleistocene (Chappell 1978), probably the last
one at LS ka BP, it was continuously distributed atong
a coastal lowland plain to the south of the present
southern coastline of Australia. It may be supposed
to have extended from easter Bass Strait to the western
limit of the Great Australian Bight (cf. Nelson 1981,
fig, 2). A subsequent-rise in sea level of more than
IO) TChappell 1978) would then have been
responsible tor the present day disjunetions. The
morphological evidence suggests lhal the Westem
Australian form is a derived rather than ancestral form,
Thus it js proposed that, some time within the
Wisconsin glagiation when sea tevels had been lowered
by about 10m, gradital east (o West dispersal of C.
elbbaxa occurred along a broad coastal plain that fs
nvw submerged. This proposal of east to wesi
dispersal, followed by subspeciation in the west as a
result of vicariaat isolation, parallels the pattern of
specialion in Western Australian frogs first proposed
by Main er o/, (1958) and subsequently adopted by
Main (1968) and Littlejohn (1981), ft should be noted,
however, that nore recent molecular data an frogs is
said fot (a support multiple east to west invasions
dunn the Pleistocene as being the explanation lor
speciation in Western Australia (Roberts & Maxson
U5)

If we accept submergence of the southern plain as
the explanation for the disjunctions in the distribution
of © gibbosa, then three explanations may be offered
for the apparent absence of this spevies from Kangaroo
Island and King Island. First. the species does occur
ou these islands bul the present samplings were not
miensive enough to reveal it. Second, C. grhhose was
originally present on these islands but subsequen|
ecological changes (@.g, sdlinisation oa Kangaroo
Island) have brought about its local extinction. Third,
although the original distribution of C gibbasa alony
the now submerged plain was continuous in 4 browd
sense, i) was never(heless somewhat patchy. and the
persistent land samiples provided by these two islands
were nol sufficiently large to include his cutanoidl.

Ackiowledgments

During the fieldwork on King island in December
1991. | was accompanied by Russell Shiel, Grace and
Peter Tyler and Robert Walsh. On Flinders Island in
February 1993 | had the company of Peter Kew, Colin
Magilton, Russell Shiel, Peter Tyler and Robert Walsh
] thank al! of these people for their companionsinp and
assistance, The field work carried out on King Island
and Flinders [sland was supported by a Natonal Estat
Grant to P_ A. Tyler. Additionally, Monash University
contributed funds towards the King Island expedition

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Bary, AB. 19795 Further conmiburons ty a krowledge
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__ {19%4) A new species of Calamoedia (Copepoita:
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1992) “The Non-marine Centtopagidae (Copepoda:
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____ (1995) Distinctive aspects of the zooplankton of lange
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Structure in natural communtiies; a mode} and evidence
from field experiments pp, 460-490 Jn Cody. M, Lb, &
Diamond, Jo M. (Eds) “Eeology and Evolinon ot
Communities” (Harvard Univ. Press, Cambridge).

Gatioway, Ro W. & Kemp, G, M. (198)) Liste Cairo
environments in Australia pp. 52-0 Jn Keast, A, (Ed)
“Ecologial Btugengraphy af Austral” Vel | Gunk, The
Hague).

LivtLeioHn, M. J, (1981) The Amphibia of mente southern
Austen; 4 zoogeographic perspective pp 1305-1340, (nd,
Vel ou

Macarriiok, RoW. & Wirsen, BE. ©. (1967) “The Theory
ut Island Biugoogtaphy” |Menographs in population halo
No, |) (Princeton Univ: Press, Princeton).

Main, A, R. (968) Beology, systematics and evolution of
Australian frogs, ddewnc. bool Rey. 5. 37-Bb.

, Les, AK. & Lrrreaoun. M. J. 11958) Evolution
in three genera a! Australian frogs. Evolution LZ, 224-233,

Nrisow, £. C. (1983) Phytogeography of southern Australia
pp. 735-759 fn Keast. A, (Ed.) “Keological Biogeography
of Australia” Vol. I (unk, The Hague).

Raw inson, PA. (1974) Binzeopraphy and ecology of ihe
repliles of Tasmania and the Bass Strait area pp. 291-334
Jn Williams, W. 0, (Ed.) “Biogeography and Ecolnpy of
‘fasmunin” (Junk, The Hague},

6 I. A. E. BAYLY

Roserts, J. D. & Maxson, L. R. (1985) The biogeography STEVENS, G. R. (1980) “New Zealand Adrift” (A. H. & A.
of southern Australian frogs: molecular data reject multiple W. Reed, Wellington).
invasion and Pleistocene divergence models pp. 83-89 In
Grigg, G., Shine, R. & Ehmann, H. (Eds) “Biology of | WILLIAMS, W. D. (1986) Conductivity and salinity of
Australian Frogs and Reptiles” (Surrey Beatty, Sydney). Australian salt lakes. Aust. J. Mar. Freshw. Res. 37, 177-182.

THERMOLUMINESCENCE DATING OF VOLCANIC ACTIVITY
AT MOUNT GAMBIER, SOUTH AUSTRALIA

By G. B. ROBERTSON*, J. R. PRESCOTT* & J. T. HUTTONT

Summary

Robertson, G. B., Prescott, J. R. & Hutton, J. T. (1996) Thermoluminescence dating
of volcanic activity at Mount Gambier, South Australia. Trans. R. Soc. S. Aust.
120(1), 7-12, 31 May, 1996.

There are several products of volcanic activity which have the potential to be dated by
thermoluminescence (TL) such as lava, volcanic ash and glass, and layers of tuff and
sand lying beneath a lava flow. One of the most important factors in obtaining a
reliable date is the search for materials which have been sufficiently heated or
bleached by sunlight to reset the TL clock at the time of the eruption. We report the
investigation of a number of such materials from the Mount Gambier volcanic
complex. A date of 4.2+0.5 ka was obtained for the baked tuff underlying the lava
flow at Valley Lake. Other results suggest that an additional event may have occurred.
about 7 ka ago (1.e. during the Holocene). Lava and glass samples from nearby sites
had insufficient amounts of datable quartz and other samples had not been sufficiently
heated but the investigation has been valuable in providing evidence of the extent to
which TL dating can be applied to a context like the one at Mount Gambier.

Key Words: Thermoluminescence dating, volcanism, Holocene, Mount Gambier.

Transactions wf the Reval Society of 8. Aust, (1996), 12001), 7-12.

THERMOLUMINESCENCE DATING OF VOLCANIC ACTIVITY
AT MOUNT GAMBIER, SOUTH AUSTRALIA

by G, B. ROBERTSON’, J. R, PRESCOTT* & J. 'T, HUTTONT

Summary

Rosertson, G. B,, Prescorr, J. R. & Hutton. J. T. (1996) Thermoluminescence dating of volcanic activity
a Mount Gambier, South Australia. Trans. R. Soe. S. Aust. 120(1), 7-12. 3) May, 1996.

There are several products of volcanic activity which have the potential to be dated by thermoluminescence
(TL) such as lava. volcanic ash and glass, and layers of tuff and sand lying beneath a lava flaw. One of the most
important factors m obtaining a reliable date is ihe search for materials which have been sufficiently heated ur
bleached by sunlight to reset the TL clock at the time of the eruption, We report the investigation of u number
of such materials (rom the Mount Gambier volcanic complex. A date of 4.2 40.5 ka was obtained lor the baked
tuff onderlying the Java flow at Valley Lake, Other results suggest that an additional cyent may hive occurred
about 7 ka ago (i.e, during the Holocene), Lava and glass samples from nearby sites had insufficient amounts
of datable quartz and other samples had not been sufficiently heated bul the investigation has been valuable in
providing evidence of the extent to which TL dating ean be applied 10-2 context like the one at Mount Gambier.

Key Worbs: Thermoluminescence dating, yalcamsm., Holocenc, Mount Gambier.

Introduction

There has been a number of attempts at determining
the sequence of events and the time scale involved in
the formation of the voleanic complex al Mount
Gambier in South Australia's South-east. The findings
indicate that the volcano as we seo it today is the resull
of a very complicated series of events, possibly spread
over several thousand years. The earliest radiocarbon
dates obtained by Fergusson and Rafter (1957) indicated
two tain periods of eruption, one at 4700+ 70 years
B.P.. the other at 1410+ 90 years B.P. These dates were
incorporated into.a geological history of the eruptions
by Sheard (1978). Subsequently Blackbum er al, (1982)
concluded that the most likely C-14 age was about 4
thousand years (ka) although charcoal samples were
found covering the range 3.5 10 & ka. Barton and
McElhinney (1980) suggested that Mount Gambier must
pre-date 5-6 ka B.P. Recently published work by Leaney:
et al. (1995) on C-14 in the inorganic and organic
carbon fractions of Blue Lake sediment cores point ta
volcanic activiry at about 7 ka. On the other band
Sheurd (1995) now interprets the various C-14 ages as
indicating a penod of activity commencing 5-4.3 ka
B.P. and extending over perhaps 300 years. Nearby
Mount Schank has been dated by thermoluminescence
at 4.9 40.5 ka (Smith & Prescott 1987) in agreement
with @ palaconiapnetic measurement of Barbetti and
Shear (981) who placed the eruptions of Mount Schank
and Mount Gambier either between 1 and 5 ka or older
than 7 ka.

dn 1987, in collaboration with CSIRO, we enrbarked

* Department of Physics and Mathematical
University of Adetaide 5, Aust, 5005,
t Deceased

Physics,

On a programme to date the cruptions at Mount
Gambier using thermoluminescence. This technique
depends on the measurement of the energy imparted
toa mineral crystal over time by the ionizing tadiation
generated by radioactive elements in the environment
and by cosmic rays; this energy is released as light
when the mineral is heated in the laboratory. Its success
as a dating method depends on the fact that the TL
was set to zero by the event being dated, in this case
either by the heat generated by the volcanic eruption
or by the bleaching by sunlight of ejected material.

In the first tnstance samples were obtained from sites
where it was considered likely that sufficient heating
had taken place to reset the TL clock, and Where there
were likely to be sufficient quantities of quartz for an
analysis. The quartz is derived from country rock
sediments through which the volcanic conduit passed-
The sites chosen were at Valley Lake, at Brownes Lake
and at the Blue Lake crater, where there were layers
of heated material underlying the lava flaw. Later the
scope was extended to include maierial which may have
been blown into the air during the event, heated and/or
bleached during transport and deposited in positions
more or less remote from the central crater. Tulf
samples were collected from several sites in the Mount
Gambier and Mount Schank areas where it was thought
that they might have been associated with the lormulion
of ether of the volcanoes.

Details of Samples

Details of the samples collected from the Mount
Gambier complex are given in Table 1. The mineral
quoted refers to the material that was extracted for the
TL. measurements, The quinz simples. were collected
and prepared according to standard practice (Huntley

8 G. B. ROBERTSON, J. R. PRESCOTT & J. T. HUTTON

Taare |, Details of the samples collected in the Mount Gambier area.

TL Date
Sample Site Lithology sampled Mineral Y/N
MGth/3 Brmwnes Lake Spatter lava Quartz N
MG2t/I Blue Lake pump house Heated tuff deposits Quartz ¥
MG2S/| Blue Lake pump house Bridgewater Fm. sand below tuff and lava Quartz Y
MG2b/10 Rlue Lake pump house Hard ruff 10 cm below lava Quartz Y
Fine grains = N
MG2e/12 Blue Lake cliff behind pump house = Upper tuff (c¢. 20 m above level of MG2S/1) Quartz y
Fine grains oN
MG2id/l2 Blue Lake carpark Banded upper tuff (same level a5 MG2e/12) Quartz Y
Fine grains oN
MG4 Devil's Punchbow| Sediment/tull Quartz N
MG5S/U.1 Valley Lake, Nurses Landing Baked tuff below basalt (0.1 m) Quurty, ¥
MGSS/0.5 Valley Lake, Nurses Landing Baked tall below basalt (0.3 1) Onartz y
Glass N
MGSS/1.5 Valley Luke, Nurses Landing Baked tuff below basalt (1.5 mi) Quurtz Y
MG5 lava —- Valley Lake, Nurses Landing Basalt Quartz N
MO6S8/60 2 kro south af Mount Gambier Tulf, sunlight bleached Quartz, Y
MG? Potters Point Lookout Ropey lava from path below tank Quurtz N
SC10S/0.6 = Mount Schank Hard tuff layer Quartz Y
SCI2S/a Mount Schank Hard layer of bedded tuff Quartz Y
etal. 1993), leading to extraction of 90-125 pm quartz
grains, the yield amounting to | to 2% of the bulk dry
weight. Fine grains containing a mixture of minerals
were extracted from the 4-11 yam fraction. _
The location of the sites is shown in Fig. 1. { af | GARDLTIE PENT
The spatter lava at MGIb/3 is described by Sheard Ra\ So eS THE MEF | LT UASALT Lv
(1978) as representing the last evidence of volcanic NH \ SE |
activity, Unfortunately no material suitable for TL ‘ah Te \ he ‘|
daling was extracted from it. The same outcome Ns | chet :
resulted from attempts to extract quartz from the ropey ( ARP ALTA
lava at MG7, Yh a
The MG2 site at the Blue Lake was extensively SPO ote) ude |
sampled from the sides of the crater just below the Pe. ? scen u
pump house where there are heated tuff layers covering <9 ‘ =

sunds of the Bridgewater Formation and from the cliff
face above and behind the pump house where the tuff
layers were deposited as a result of fall out from the
eruptions.

The group of samples (MG4) collected near the

Devil's Punchbowl! contained terrestrial sediments of

the Wangerrip Group from below the Gambier
Limestone, dispersed in the volcanic tuff where it had
been carried by the eruption, They were found not to
have been sufficiently heated to make TL
measurements.

The MGS5S samples from Valley Luke were collected
from the baked tuff at 0.1, 0.3 and L5 m below the
base of the lava flow. It was not expected that the lowest
level would have been sufficiently heated but i! was
included to check this point. A small quantity of
volcanic glass Was extracted from the MG58/0.3
sample but this was highly magnetic and produced an
insignificant TL signal, Basalt was also collected [rom
this site, It contained very small quantities of extracted

— f q
e aw ‘ |
| my Part ‘Mar Tbiewet
Histor ~y yee
ae ed
| lage Nort Audives hue

1 : Fane tn Th!

HOSES
AMCHHLs

CENIENen
myers

— Mb

. HeyIUS
TOMO M EOL

marros

Fig. 1, Lower South-east of South Australia showing the
locations of the voleanic deposits and details of the sampling
sites. Adapted from Sheard (1978, 1983).

TL DATING AT MOUNT GAMBIER 9

quartz, and 300 jim sections cut from the bulk sample,
which might contain other minerals, yielded only low
levels of TL,

Other samples of tuff and sand were collected from
a site some distance to the south of the lake (MG6S/60)
and from sites in the Mount Schank area (SC series),
| km east of Mount Schank.

TL Analysis

The measurements required in order to caleulate an
age are the intensity of the ‘TL in the natural sample
and the TL sensitivity, as shown by the TL generated
by a known amount of radiation. These two values are
combined ta determine the amount of radiation dose
received by the natural sample, the equivalent dose,
ILis also necessary to measure the environmental dose
rate received by the sample in the local environment
The age 1s then given by
Equivalent dase (Gy)

Dose rale (Gy ka!)

where the unit of radiation dose ts the gray (Gy).
The TL, analysis protocal was selected accard|ng la
whether the samples were thought to have been zeroed
by beat (samples in the MG2 and MGS series) ur by
sunlight (samples from MG6 and the SC series). In
the case of the heated samples. the usual procedure for
pottery dating was followed (Aitken 1985). tn the case
of the sunlight bleached samples the selective bieach
method was used (Prescott & Mojarrabi 1993): the
analysis to give the equivalent dose followed the
“Australian slide" method (Prescott et a/. 1993).

Age tha) =

Dose Rate

The sample dose rates were determined by thick
source alpha counting (TSAC), by neutron activation
analysis (NAA), by X-ray fluorescenve spectrometry
(XRS) and/or by ganuiya-ray s¢intillometry (seint), as
appropriate lor (he esumation of the elements uranium,
thorium and potassium (Prescott& Hutton 1995), The
concentrations of these elements are given in Table Z
together with the calculated dose rates. The samples
are grouped according to type and location and it is
noticeable that the baked tuff below the layer of lava
has higher concentrations of all the elements than the
tuff found in other locations and also the Sand from
the Bridgewater Formation, Consequently the dose
rales in the baked tull-are about twice those in other
tuff samples and higher levels of TL might be expected
in these,

For each sample the quoted concentrations of U and
Th agree reasonably well (within 2 sigma limits for
most of the samples) among the various methods of

analysis used, implying that the members of the U amd
Th decay chains are in equilibrium in these samples.
The worst case ts MG2S/1 which shows evidence of
higher yalues for “*U obtained by TSAC and
scintillometry compared with the values obtained by
DNA and by XRS_ This situation has been shown ty
arise in cases of disequilibrium between the parent
“NU and its immediate decay products dows to 4U
and the remainder of the decay chain from 7°TIr ty
"Po occurring in samples collected jn a similar
environment at Mount Schank (Smith & Prescott
(987). The dose rates have been calculated to allow.
for the deficit in the early part of the chain, taking the
extreme possibility of a value of G80 pg g! for
AL24U and 1.49 jyrg! for Th onwards.

The dose rates were derived from the element
concentrations using the conversion factors of Nambi
and Aitken (1986), The values quoted in Table 2 include
a cosmic ray contribution of O10 to O12 Gy kat
depending on the site (Preseott & Hutton 1994). The
weighted means of the various estimates ot dose rate
were used in the calculation ol the aves of the samples.

Age Determinations

As is'usual for heat-zcroed samples. first- and
second-glow growth curves were obtamed and the two
infereepts on the dose axis combined to obtain the
equivalent dose (Aitken 1985), The growth curves were
fitted to a linear relationship and used to generate the
equivalent dose plateau which is shown superimposed
on the natural glow curve of MG2S/l in Fig, 2. Valid

Equivalent dose (Gy)

Temperature (°C)

Fig. 2, Equivalent dose plateau for the MG2S/) sample with
4 TL glow curve superimposed -

= 10

a

* ao

i

nal

= + +e eee

ra

vo

=

5. mie wu ee Ee yz
ir 275 300) W5 350 375 400

Temperature (®C}

Fig. 3. Equivalent dose plateau for the MG5§/0,3 sample with
its TL glow curve superimposed. The double plateau
indicates Iwo penods of heating.

10 G, B, ROBERTSON, J. R. PRESCOTT & J. T. HUTTON

TABLE 2, Concentrations of uranium, thorium and potassium in the samples, determined by thick-source alpha counting
(TSAC), X-ray fluorescence spectrometry (XRS), neutron activation analysis (NAA), delayed neutron analysis (DNA) and

Sample Method Uranium Thorium Potassium Dose Rate
we ee" % Gy ka!
Baked tuff below tava
MG2t/1 TSAC 2.344048 6.434161 2.45 +014
XRS 2.604 0.40 8.50 40.50 1.254001 2.67 £0.10
MG5S/0.1 TSAC 2.71 40,27 8.04 + 0,92 3.08201
XRS 3.00 + 0,40 10.20 +0.50 182+ 0.01 3.32+40.1
MGS5S8/0.3 TSAC 2.464 0.41 10,79 + 1,38 3,52 +012
DNA, NAA 2.2020.1 9.00 + 0,45 3.38 + 0.06
XRS 2.304040 10.40+ 0.50 2.19+ 001 3,51 40.08
Scint. 2.62+0,35 8.524005 2.03 + 005 3.47 +009
MGSS/1.5 TSAC 20403 6941.4 2.6) +0)
DNA 1.87 +0,08
XRS 1.594001
MG5/Lava TSAC 1.93+0,35 4.594115 2.52 + 0.08
DNA, NAA 1.10 + 0.40 570+ 0,50 240+ 0,08
XRS 1.50+0.01
MG2b/10 TSAC 2.58 40,38 8.49 + 1,05
DNA 2.4340,09
XRS 2.09 +002 3.26 + 0,22
Bridgewater Formation sand
MG2S/ TSAC (a) 1444 0.21 4,5940.69 1,.58+0.07
(b) 1,07 +0,20 6,06 + 0.65 164 40.07
DNA, NAA 0804010 5.50 + 0,40 1.564 0,05
XRS 0.90 +040 7.40 +£0.50 0.60 +001 1.48 + 0.08
Scint 1.494 0.18 577 40,24 0.78 +002 1.73 + 0.05
Tuff various locations
MG2¢/12 TSAC 141 +0.26 3.03 + 0.68
XRS 01.66 4 0.01 1.274008
Scint. 150+0.12 4.3240.20 071 40,02 151 +005
MG2d/I2 TSAC 1.05 + 0.40 6.3241.36
XRS 0.74+ 0,02 1.504014
MG6S/60 TSAC 1.904.0.46 4.70£1.70 1.66 +0.13
DNA, NAA 1.84+0.19 6.244019 1.76 + 0,07
XRS 074+ 0.05
SCIOS/06 TSAC 2,04 + 0.33 S.fl 41.15 1.934014
DNA, NAA 2.32 + 0.60 733 +4017 2154015
XRS 1.70+017 TWL077 0,9440,10 2.03 40.12
SCI2S8/a TSAC 1.8040,39 4.0641.33 171 0,12
DNA, NAA 0.89 + 0,42 3.264015 1584010
XRS 1.60 +: 0.16 6.704067 0.87+0,10 1.8640.09
dating requires that there should be a distinct plateau Discussion

over a range of temperature from about 300°C ta
400°C (Wintle & Huntley 1982). This conditron is well
met in the case of sample MG2S/1. Sample MGS5S/0.3
on the other hand, has two plateaux (Fig. 3). The
double plateau suggests that this sample experienced
two heating events, the second one of Which did not
reach a sufficiently high temperature to zero the TL
above 350°C, Il therefore yields two ages, for differing
events. Analysis of the phenomenon of the double
plateau is given in more detail in Robertson e7 al.
(1996). The calculated ages are given in Table 3
together with the equivalent doses and the dose rates.
The analysis of MG2b/10 was not completed because
there was not enough pure coarse-grained quartz.

The ages obtained show a spread of values which
are not easy to interpret but it is interesting to note that
in the MG samples there is evidence of events haying
occurred at about 4 ka and at about 7-8 ka, coincident
with other methods of dating. The figures for MG5S
show the single age for the 0.1 m sample and the twa
ages resulting from the double plateau for the 0.3 m
sample, H was expected that, as the last heating raised
the temperature of the 0.3 m level to no more than
350°C, the age of the 1.5 m level would be at least
7 ka, but the very much older age (98+15 ka) does
not fit in with the observations and the geological inter-
pretations (Sheard 1978, 1995). li suggests that at least

tl, DATING AT MOUNT GAMBIER i

TABLE 3, Age estimates of the samples maained from the equivalent doses and dose rates skown.

Saniple AdTL Analysis Equiyalent Dose Rate Age (ka)
Code Dose (Gy) (Gy ka!)
M21 95042 Heated 9.464047 2604003 3.64 + 0,25
MG25/1 95043 Heated 913 +0,32 1.454003 6.294025
MG2e/12 95044 Heated A45 435 1.394004 25) +30
MGid/12 95045 Hesiod 8210 1,50+0,14 55417
MG5S/U.4 95039 Hewiel 18.7 41.3 3.2040.17 4.9) +£0.48
MG5S/0,3 YS Heuted 4 at 3,4640,05 4.084038
95040 Heated 23.9425 3,4640,05 TAS 2075
MG5S/1.5 95041 Hewted 241 4 40 2.46+0.06 OR + 15
MG6S/60 95046 Selective nleach 14.16 +060 1.744006 §.14+0.44
SCIDS/O 98047 Selective bleach 3,21 40,20 2.032008 1.584012
SCI25/a FMB Selective bleach 2.234084 1.76 2006 1.274048

the quartz present inthis sample was actually in place
well before the eruptions occurred.

The 7 ka age found here is to be compared with that
reported by Leaney er al. (1995) for a major change
in the sedimentology of the floor of the Blue Lake.
However it is difficult to reconcile the time sevle of
the history of the Blue Lake as set by Leaney er al,
with the violent eruptive events traced by Sheard (1978,
1990). One possibility is that Leaney eral, have net
sufficiently allowed for the incorporation of old carbon
into their inorganic samples.

The very large ages for the tuff MG2c/l2 and
MG2d/12, collected from the cliff face at the Bluc Lake
pump house suggest that this material dud nol in fact
become sufficiently heated during the eruptions The
very recent ages for the Mount Schank samples suggest
thal there was some bleaching mechanism occurring
1-2 ka ago, another date which has previously been
associated with events at Mount Gambier, although it
ix now thoughe that the C-14 date on which this is based
may he due tn intrusive tree root charcoal (pers. comm.
Blackburn to Sheard 1995),

Because of Lhe problem in TL dating of determining
whether the event that is being dated removed all the
existing stored energy, all the dates should be carefully
considered with regard to other avidence. For example,
do the two apparent groups of ages obtained, ie. c.4
ka and c.7 ka really indicate two eruptions sepurated
by 3 ka? During this time interval there should have
been considerable weathering of any tuff ejected.
However, the chemistry of the tuff samples MGS5S/0.1,
MG5S/0.3 and MG28/1 tuff (grouped together in Table
2) shows that the ratio of the mote soluble and mobile
clements, sodium, magnesium and phosphorus to the
insoluble element, titanium, is the same as that found
for the solid Inve, Thus the lava must have proteeted
the tiff very soon after it was deposited. There are
also ny known palacosols developed within this tulf
or on top af the lava (Sheard 1990), The exposed tuff
MG6S/60 does show considerable weathering, wall the
loss of about 50% of tbe soluble elements in relation

to titanium. Incidently the loss of elements from
MG6S8/60 is about the same as the Joss of the sanic
elements from the similar site SCIOS/0.6 near Mount
Schank, The age of the Mount Schank eruption was
found to be 4.96.5 ka by Smith and Prescott (1987)
using well baked tuff/sand under the lava flow and the
age of MG6S/6l) should be similar. ‘These chemical
considerations support the suggestion that neither
healing nor exposure to sunlight has been sufficient
in some of the samples to remove all of the pre-existing
TL.

M, J. Sheard (pers. comm.) states, “The phreatic
style of eruption so evident at Mount Gambier (i.e.
associated wilt copious H,O) means that many
eruptive products incorporating exotic quariz may nol
have heen raised much above 100°C. . . . In addition,
under high volume tephra explosions some or most of
the exotic quartz could escape light bleaching (ie. re
setting) due lo ash cloud or surge cloud density and
subsequent rapid fallout and burial. Thus, it is possible
to have exotic quartz grains, only partially reset or left
with their “older” 'TL signature, incorporated into much
younger tephra.” Sheard's remarks seem to indicate the
possibility of finding samples containing partially
heated quartz grains mixed with thorougiiy bleached
tephra. und so possibly being able to detect different
dates for differem sized prain fractions, With this end
in view, un altempt was made to date the fine grain
fraction of the samples from which quartz prains had
been extracted, but it was found in the three samples
tested (MG2b/10, MG20/12 and MG2d/12) that either
there were insufficient fine wralos or that the fine prains
showed no TL and no sensitivily to radiation.

The TL dates in conjunction with other geochrono
logical evidence have reinforced the belief that then:
was volcanic activity at Mount Gambier at about 4 ki
ago, and that there may have been activity at 1.3 and
7 ka, but that the latter dates should he accepted with
caution. The investigations have glse illusjrited that

12 G. B. ROBERTSON, J. R. PRESCOTT & J. T, HUTTON

it is often difficult to select appropriate samples for
TL dating from a complex system like the one at Mount
Gambier.

Acknowledgments

The late John Hutton was associated with all of the
work described here. It is a matter of regret that he

did not see the final publication. The project was begun
in collaboration with CSTRO and the work was
supported by CSIRO and the Research Fund of the
University of Adelaide, and by AINSE. Kym Lawry.
Adrian Murphy and Phillip Stamatelopoulos assisted
with the work. M. Sheard and D. Leaney gave much
useful advice.

References

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(Academic Press, London)

Barnettl, M. & SHEARD, M, J, (1981) Palacomagnetic
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J, Geal. Soc. §. Aust. 28, 385-394,

Barton, C. E. & McELuinney, M. W, (1980) Ages and
ashes in lake floor sediment cores from Valley Lake, Mt
Gambier, South Australia. Trans. R. Soc. §. Aust. 104,
161-165,

BLACKBURN, G., Altison, G, B. & Leanpy, FW. J. (1982)
Further evidence on the age of tuff at Mt Gambier, South
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Prrausson, G. J, & Rarer, T. A. 957) New Zealand C4
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Quat. Sei. Rev, 12, 1-20,

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Nabil, K. 8. V, & AiTneN, M. 1b (1986) Annual dose
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Preseoi, J. R.. Husireey, BJ & Horton, J.T. (1993)
Estimation of equivalent dose in thermoluminescence dating
~ (he Australian slide method, Aneient TL UW, 1-5,

& Hurrron, J. T, (1994) Coamic ray contributions to

dose rates for luminescence and ESR dating: large depths

and long-term time variations, Rudiat. Meas, 23, 497-500.

& (1995) Environmental dose rates and

radioactive disequilibrium from some Australian

luminescence dating sites. Quar. Sci, Rev, (Quat. Geovhron)

439-448,

& Moijarrasi. B. (1993) Selective bleach: an
improved partial bleach technique for finding equivalent
doses for TL dating of quartz sediments. Ancient TL 1,
27-30.

Ropertson, G. B., Prescot, J, R. & Hutton, J. T. (1996)
Thermoluminescence date for lava. flow ut Mount Gambier,
South Australia, Proc. Fifth Aust. Archaeometry Conf,.
Armidale, 1994 in press.

Suearp, M. J. (1978) Geological history of the Mount
Gambier volcanic complex, southeast South Australia.
Trans. R. Sov, Aus}. 102, 125-139,

(1983) Volcanoes’ pp. 714 Jn Tyler, M. J., Twidale,
C.R., Ling, J. L,, & Holmes,.J, W. (Eds) “Natural History
of the South-East.” (Royal Society of South Australia,
Adelaide).

(1990) A guide to Quaternary volcanoes in the lower
South-east of South Australia pp. 40-50 fn Drexel, J. F.
(Ed.) “Mines and Energy. Review South Australia.” No. 157,
(Department of Mines and Energy).

(1995) Quaternary volcanic activity and volcanic
hazards pp. 264-268 /n Drexel. J, F & Preiss, W. V. (Eds)
“The geology of South Australia, Vol. 2° South Australia
Geological Survey Bulletin, 54.

SmivH, B. W, & Prescot, J, R. (1987) Thermnluminescence
dating of the eruption at MtSchank, South Australia, dase,
J, Earth Sei. 34, 935-342

Winthk, A, G & HUNTLEY, DLJ. (1982) Thermolumines-
cence dating of sediments, Quan Sei, Rev b, 34-53.

NEMATODES ASSOCIATED WITH DIPTERA IN
SOUTH AUSTRALIA: A NEW SPECIES OF FERGUSOBIA
CURRIE FROM A FERGUSONINID AND A NEW RECORD OF
SYRPHONEMA LAUMOND & LYON FROM A SYRPHID

By KERRIE A. DAVIES* & JANINE LLOYD*

Summary

Davies, K. A. & Lloyd, J. (1996) Nematodes associated with Diptera in South
Australia: a new species of Fergusobia Currie from a fergusoninid and a new record
of Syrphonema Laumond & Lyon from a syrphid. Trans. R. Soc. S. Aust. 120(1), 13-
20, 31 May, 1996.

Fergusobia fisheri sp. nov., associated with a fly Fergusonina sp., is described from
leaf galls on a hybrid of Eucalyptus leucoxylon. Like other species of Fergusobia,
these nematodes are semi-obese and have a generation parasitic in the fly followed by
a generation parasitic in the plant. The new species is distinguished from F.
tumifaciens by its smaller size, larger manubrium on the spicule of the male, and
smaller cephalic region and tail in the parthenogenetic female, with vulva and anus
opening into cuticular depressions. Syrphonema sp., a nematode parasite of syrphid
flies, is described and recorded for the first time outside France.

Key Words: Taxonomy, Nematoda, Fergusobia, Diptera, Syrphonema, new species,
new record.

Transactions of the Royal Society af S. Aust. (1996), 1200), 13-20.

NEMATODES ASSOCIATED WITH DIPTERA IN SOUTH AUSTRALIA: A NEW
SPECIES OF FERGUSOBIA CURRIE FROM A FERGUSONINID AND A NEW
RECORD OF SYRPHONEMA LAUMOND & LYON FROM A SYRPHID

by Kerrie A. Davies* & JANINE LLOYD*

Summary

Davies, K. A. & Luoyp, J, (1996) Nematodes assuciated with Diptera in South Australia! a new species of
Forgusobia Currie from a fergusoninid and a new record of Syrphanema Laumond & Lyon from a syrphid. Trans.

R. Soe. 8. dust. 1200), 13-20, 31 May, 1996.

Fergusobia fisher’ sp. nov., associated with a ly Fereusenina sp,. is described from Jeaf galls on a hybrid
of Bucalypruy lewcexylon. Like other species of F-rgusobia, these nematodes are semi-obese and have a generation
parasitic in the fly followed by a generation parasitic in the plant, The new species ix distinguished from J? ramifaciens
by its smaller size. larger manubrium on the spicule of the male, and smaller cephalic region and fail mn the
purthenogenetic female, with yulva and anus opening into culicular depressions, Syrphonema sp., 4 nematode
parasite of syrphid Nes, ts described and recorded for the first time outside France

Key Worps: Taxonomy, Nematoda, Fereusobia, Diptera, Syrphonema, new species, new record,

Introduction

This paper describes a new species of the tylenchid
nematode Fergusobia, the only zenus of nematode
known fo parasitize both a plant and an insect
(Maggenti 1981). Jt has a bipartite life cycle, with a
generation parasitic in galls on Myrtaceae followed hy
one parasitizing the fergusoninid fly Fergusenina
Malloch. Also reported ts the first collection outside
France of the rhabditid nematode genus, Syphonem.
Both nematodes were collected in Adelaide, South
Australis.

Materials and Methods

Nemutodes were collected from galls cut open in tap
water, relaxed and fixed in hot FA 4:1. Insects were
dissected in 0.85% saline, and nematodes tram these
were fixed as above. Nematodes wene transferred from
fixative to [1% pglyceral in 30% ethanol in glass blocks
and placed in a desiccator containing 96% ethanol for
2 days. For the following J4-days nematodes were kep|
ut 40°C and the Mocks were partially covered with
glass lids. During the first three days, one ar cwa drops
ofa solution of 5% glycerol in 95 ml ethanol were
added four times daily. Slow evaporation was continued
thereafler, For light microscopy, processed nemavodes
were mounted in glycerol on glass slides and exarntned
using, interfetence microscopy. Scanning electron
Tnicroscopy studies were made. Nematodes were taken
from glycerol, passed through a series of
ethanol/glyveral solutions with increasing proportions

* Deparment of Crop. Protection University of Adejanle
PMH J Glen Osmond S. Aust. S064

of ethanol and washed three times with 100% ethanol,
They were then dried using CO, in a critical point
drier, mounted on stubs, sputter coated with
approximately 30 nm of gold and viewed at 20 kV

Measurements aré in zm, Drawings and measure-
ments were made from material mounted in glycerol
using a camera lucida. Body width was measured at
mid-length, Spicules were messured along the mid-
line in lateral view, De Maa’s ratios, je, Y = anter
ior end to vulva as percentage of body length, T =
length of testis tram) cloaca to flexure as percentage
of body length, 4 = length divided by greatest body
width, b’ = length divided by distance trom anterior
end to base of oesophageal glands, c = length divided
by tail length, & = tail length divided by width at anus
were determined. Compuariscins were made with
described species using published descriptions.
specimens of parthenogenetic females of Fergusobia
widgha Siddiqi (Queensland Museum (QM)
G200512-200519) and specimens of all stages except
parasilic females of F lumitacieny Curne isolated by
the authors from flower bud galls on Eucalyptus
cumeldulensis Dehn, at Urrbrac, South Australia
(Waite Institute Nematode Collection (WINC) 943)
The holotype of the new species is deposited in the
South Australian Museum, Adelaide (SAM).

Taxonomic descriptions

Fergusobia fisher sp. nov.
(FIGS 1-2)

Holearype: Parthenogenetic Q, Black Forest, South
Australia (34°57 'S, 138°34'E), 3.vini.t993, W. Frost.
collected trom a leaf gall on a hybrid of Fucalyprie
leucaxvlon Fo Muell,, ARC207051 (SAM),

14

Paratypes: WINC 81S, slides [-28, including 20
parthenogenetic 9 9, 25 preparasitic infective 9 9,
39 parasitic 9 O, 32 ao, 59 juveniles, collected
from leaf galls on a hybrid of E. levcaxylon at Black
Forest between August and October 1993, by W. Frost.

Measurements: Table 1.

Description af parthenogenetic female (Figs 1A, H,
2A)

Occurs in leaf gall. Dorsally curved when relaxed
by heat, with ventral side convex. to form open. C-
shape, Smaller than amphimictic preparasitic female.
Cuticle weakly annulated, striated; lateral fields
obscure, Cephalic region small relative to width of
body atanterior end, off-set, lightly sclerotized; region
appears roughly circular, unstriated, with 8 sectors;
dorsal and ventral sectors less than half width of each
of others; in side view sectors with rounded outline
and no central elevation around stylet opening.
Amphidial openings pore-like, situated near dorsal
edge of lateral sector. Stylet slender; conus forming
half length; small fusiform basal knobs. Orifice of
dorsal oesophageal giand approximately 3 pm posterior
ip stylet knobs. Digestive tract with swollen anterior

TasLe 1. Measurements of Fergusobia fisheri sp. nov.

K, A, DAVIES & J, LLOYD

part with valve-like structure. followed by short narrow
“isthmus” which widens abruptly to broader part
associated with the oesophageal glands; oesophago-
intestinal junction obscure. Lumen of tract broadens
posterior to “valve” and again at level of secretory-
excretory pore. Oesophageal glands large, occupying
about three quarters of body width, extending over
intestine to about 55% of total body length; dorsal
gland with large nucleus. Secretory-excretory pore
50-92 ym from anterior end with short duct leading
directly to excretory cell. Nerve ring at base of swollen
anterior part of digestive tract, Hemizonid not seen.
Reproductive tract with simgle gonad, prodelphic,
extending to nerve ring; oviduct flexed in some
females; uterus with quadricolumellar, often contains
single egg, curves to join vagina, which js directed
slightly forwards. Vulva usually conspicuous depressed
transverse slit, but in some specimens, vulval lips
protrude slightly. Cuticle wrinkled on ventral side just
anterior to vulva in half specimens examined. Rectum
simple tube, without obvious musculature: anus
inconspicuous pore, usually associated with distinct
indentation in the cuticle. Tail conoid, narrowing
sharply w rounded tip, 1,0-1.4 times as long as anal
body width, Phasmids not seen.

Measurements in jim; n jor each measurement in brackets below mean.

Holotype Paratypes
(parthenogenetic Parthenogenetic females Males Pre-parasitic females —- Parasitic females
female) mean S.D. range mean §.D, range mean §.D. range mean S.D range
Length 250, 25300 «23.2 «6228 = 336 39.5 292- 349 358 24) TRH STR 690-
(12) 305 (16) 453 (25) 395 (20 905
Width 5) 3 21 3340 37 37) 0-32-4638 62 2849 U2 79 105-
(12) (16) (25) (20) 128
Styler length 9) 94 038 9-10 Q LO 8-1 8 L4 6-11 nil 7 =
(8) (13) (ti)
Anterior end to BO zh) 5.4 S092 7 122 61-93 75 0.6 66-91 -
SeC-EX, pore (9) (6) (5)
Tail length 5 la 13-16 A 23-42 30 21-49 nil -
(12) (16) (25)
Vv 87 85 14-83-88 - 80 30 76-88 = 83 27 TReBY
(12) (25) (18)
T - - - 75, 72 59-84 ~ - - -
(16)
Spicule length - - Ik 18 620 - - - - - -
(17)
a 5 68 O7 60-85 92 14 63-113 95 17 88125 70 06 5.98)
(12) (16) (25) (20)
b 18 19 IS LB-2T 2R OAL 2.334 32 042 28-40
(12) (10) (9)
c 15.6 Bl 44 1025 1229 15 I? 29 44 7-23
{12) (16) (25)
¢ 1.2 LZ O13) (9-13 ‘ - - -

NEMATODES ASSOCIATED WITH DIPTERA 1S

Description of infective pre-parasitic female (Fig. 1B,
F, G)

Occurs in leaf gall, infects mature larval stage of fly.
Anterior part of nematode straight when relaxed by
heat; posterior part curved dorsally. Maximum body
width at mid-length. Cuticle with inconspicuous annu-
lations; strongly striated; lateral fields arising about
one body width behind head, with irregular, broken
striae, obscure when viewed with light microscope.
Cephalic region continuous with body, weakly sclero-

tized. Stylet slender, weakly sclerotized with smaller
basal knobs than in parthenogenetic female. Amphids
not seen, Orifice of dorsal oesophageal gland
approximately 3 ym posterior to stylet knobs. Secret-
ory-excretory pore approximately half the distance
along the oesophageal glands (65-90 ym from anterior
end), Nerve ring at base of swollen anterior part of
digestive tract. Hemizonid not seen. Oesophageal
glands often obscure, elongate, occupying about half
body width, extend over intestine to about 30% total
body length. Anterior part of digestive tract swollen,

+——.

4

Fig, 1, Fergusobia fisheri sp. noy, A, Entire parthenogentic female. B. Head of pre-parasitic female, C. Entire male (bursa
nol seen due to angle at which tail viewed). D. Detail of tale tail, ventro-lateral view, showing bursa and angulated spicule
E, Entire parasitic female. F. Entire pre-parusilic female. G, H, 1. Stylets of pre-parasitic female, parthenogenetic female
and mule, respectively. J, K. Lateral and dorsal view, respectively, of spicule, Scale bars = 10 pm G HIS K, 20 pm

ABCDE, 80 pm E.

16 K. A, DAVIES & J. LLOYD

non-muscular with valve-like structure. Intestine may
contain many dense granules or lumen may be broad
and empty, possibly reflecting nutritional status. Cells
of intestinal walls have prominent nuclei with large
nucleoli. Uterus extends almost to base of oesophageal
glands, acting as spermatheca and packed with sperm:
no post-vulval sac; vagina directed anteriorly, plugged
with refractile material; oviduct short, often with
flexure; ovary extending to nerve ring. Vulva transverse
slit, inconspicuous; lips may be raised. Anus pore-like;
rectum very small, non-muscular. Tail almost
hemispherical. Numerous large nuclei scattered along
length of nematode in epidermis.

Description of parasitic female (Fig. VE)

Occurs in haemocoel in abdomen of fly. Epidermis
thickened. Cephalic region may or may not be offset.
Body swollen, sausage-shaped, obese, filled with
hypertrophied reproductive organs. No stylet seen.
Oesophagus and intestine degenerate, anus not seen.
Ovary convoluted. Several eggs in uterus at one time.
Vulva depressed transverse slit.

Description of male (Figs 1C, D, 1, J, K; 2B, C)

Occurs in leaf gall. Body shape variable when
relaxed by heat, posterior portion of body may be
arched dorsally, tail curved ventrally. Cuticle with
longitudinal striations, without annulations; lateral
fields indistinct under light microscope, appear to be
3.or 4 incisures or several irregular striae. Cephalic
region off-set, with lightly sclerotized framework.
Stylet, oesophagus, intestine and secretory-excretory
pore all as for parthenogenetic female. Oesophageal
glands extend over intestine to about 35% of total body
length. Reproductive tract with single testis, extending
to nerve ring; extensive vas deferens, with amoeboid
sperm. Bursa membranous, smooth, often difficult to
: — see; extends to tail tip, appears to be peloderan;
» “ collapsed in specimens prepared for SEM, seen as
wrinkled membrane lying on cuticle of tail region;
variable length, usually arises 1.5-2 tail lengths anterior
to cloaca, but in one specimen arose in anterior half
of nematode. No genital papillae seen. Spicules robust,
paired, angular near their middle so that manubrium
and shaft appear to be perpendicular to each other in
ventral view; manubrium large. No gubernaculum, Tail
bluntly rounded.

Fig. 2. Fergusobia fisheri sp. nov. A. Parthenogenetic female
head to show buccal region and stylet (arrow). B, Male
tail showing bursa (arrow). C. Amoeboid spermatozoa from
squashed male. Scale bars = 10 pm.

NEMATODES ASSOCIATED WITH DIPTERA i

Diagnosis and reletionships

F fishert sp. noy. is characterized by having a
panhenogenetic lemale with ibe cephalic region sal
relative to the body width, with 4 flat terminal profile,
the volval slit in a conspicuous depression oF the
cuticle, the anal opening in a similar but smmller
depression and a short tail (0.9-1.3 times anal hody
width) with a narmow cone shape, The male fas sasrular
spicules with a large manubrivm.,

F fishert sp. nov, differs Fram F jambaphila Siddiqi
in haying a flat cephalic region without a central conical
elevation and angular spicules and fron F inidica
(Jairaypuri) and F magna because the parthenogenetic
female has a short tail. The aous opens into an oby ious
cuticular depression in the parthenogenetic female ot
£ fisheri sp. nov, bul not in the other described species.
FE fisheri sp. now, assuines a similar shape to F
tumifacicay when heat-killed bul is smaller (average
length of parthenogenetic females 253 and 318 pm
respectively and of males 336 and 420) «p-
Measurements for & curried (=tumifuctens) from
Fisher and Nickle 1968), The parthenogenetic female
of F fivheri sp, nov. has a smaller cephalic region
relative to bocly width than FO tuwifaciens and may have
wrinkled cutiele on the ventral side of the body anterior
to the vulva, absent in A fantifaeiens, The vulval slit
is situated ina distinct cuticular depression in FE fishert
sp. oy. as in jambophila and F indica, but not in
F tumifaciens or in BE magna. The volume of the tail
of the parthenogenetic female is snaller in F fisheri
sp. nov. than in FE rantifaciens, having a narrower cone
shape. The point of origin of the caudal alae ts vartable
it A fishert sp. ney, males, in contrast to F nwpifaciens.

Frymolopy

Named afler Dr J. M. Fisher, formerly of the
Department of Crop Protection University of Adelaide.

Biology, life evele and general comments

The nemalode & fiskeri sp. noy. was found on leaf
galls ot the southern blue gum. &. leacoxylen, in
association with an unknown species of the
fergusoninid fly Ferevsoitia sp, Galls were first
collected ii carly August M993, followed by successive
collections until October 892 when no nemaludes (yt
fics were found. tn the early August collection, gulls
contained many nematodes (arveniles. parthenogenetic
und infective females aid inales) and fly larvae. OF
32 larvae dissected, the abdomen oF two contained i
toral of three ferpilized infective females and one male
nematode. A week Juter, 14 ily larvae andl 1S paiparia
were collected from valls and dissected. Six of the
larvae contained two to four fertilized inlicleve te rele
nematodes. Ne pentatodes were found in poparia, In
early September, galls contained # lew uclult rrogle: sane!
infective female nematodes, Puparia cemtamned pharaic

adult flies. Nineteen pharate adult flies were dissectedl.
eight of which were male and had no nematodes,
However, 10 of the I female flies contained from |-H
(averaye 6.4) mature parasitic female nematodes per
fly. In six of these, eggs had been laid in the
haemolymph and i most cases, the eggs were in the
carly stages of embryonic developmen. One fly,
however, contained eggs in which the juvenile
nematodes were Well-formed.,

These observations on the life cycle peucrally agree
with (hose reported by Fisher & Nickle (1968) for &
curici (=tunifirciens), No nematodes have been found
in mate flies. Eggs are Jaid in the haemolymph of the
abdomen of the tly bur at is not known bow the fly
deposits both insect eggs and nematode juveniles into
Eucalypiusy ussde. Presumably, these nematode
juveniles develop jnta parthenogenetic females. which
luy eggs. It is possible that juveniles developing from
the early eggs beconte miles, as in ramnifactens,
While the first collection of F fisheri sp. nay. Trons
leat galls yielded all plant parasitic stiges Of the
nematode, the fourth stage juveniles found were all
fernale {distinguished from males by a mure bluntly
rounded tail and development of the uterus), suggesting
that male development had occurred earlier, Fisher &
Nickle (1968) stated that only fertilized infective-stage
fétnales of F curriei (=tumifaciens) penetrated fly
jurvae but female and male & fisher? sp. nov, have been
found in fly larvae. F mmifacicas may deposit eges
in the haemolymph of the fly before it emerges as an
adult female (Currie 1937) or this may be delayed uncl
after emergence of the fly from the pall (Fisher &
Nickle 1968), Here, & fisheri sp. nov. had produced
eggs before the purasitised fly had emerged from the
puparium.

Currie (1937) described one species, F tarmifaciens,
associated with Fr. carter? Tonn.. from leaf galls on
£, Stuartiana (sie) Ev. M. He alsa observed minor
morphological differences. between nematodes
callccted from leaf bud, axi bud, stem tip and flower
Dud galls associated with a number of other fly species
on more than a dozen species of cucalypt. He suggested
“further work will show that the nematodes associated
with the diffrent Mies have differences in structure
which ¢otitle them to be considered as different species”
(Curt\e 1937), These differences inelikled turn£ot un
the point of origin of the caudal alae and the size and
position of the oesophageal glands. Observations of
material held in the WINC conlinn thal these
cbanicters will be important in species differeniahon
Fisher & Nickle (1968) redescribed Fo rvrnes
(=luniifaciens) from flower hud galls an b.
Camaldulensis Dehn. associnted wath An filled
Tonn. Of the other deseribed species (a fenguvensia,
Siddiqi (1986) descrihed A ruigna from Efecalypeus
stem galls associated with an unknywn Ny bost ape
from soil under host trees, & imdiew trom suil ii lndia

18

and F) jambophila from flower bud galls on Syzygium
cumini (L.) Skeels associated with Fr. syzygii Harris,

The Waite Institute Nematode Collection contains
specimens of Fergusobia collected from stem, bud and
Jeaf galls on Excalyptus fram Adelaide and Mt
Gambier, respectively, which appear to be undescribed
Species, Fisher (pers. comm,) found an undescribed
species of Fergusohia in leaf galls on &. camaldulensis
associated with #n, lockharti Tonn, lt is apparent, given
the different forms of galls, that each of these
undescribed species of nematode is associated with a
different species of Fergusonina. Bach, however, is not
Necessarily restricted to one species of Aucalyptas.
Presumably the nematode has evolyed mechanisms,
which are probably host-specific, to escape the fly's
immunological system during the generation i which
it is an insect parasite. Thus, fergusobia, like the wenus
Anguina Scopoli (Krall 1991) may have a high degree
of host specificity, in this case, for the insect host, Jf
so, Currie (1937) and Fisher & Nickle (1968) may not
have deseribed the same species of nematode, given
that they were associated with different species of
Fergusonina. Indeed, Currie refers to the margins of
the caudal alae of males of his specimens of F
tumnifaciens as being slightly crenate, a character not
observed by Fisher & Nickle (1968) nor in specimens
isolated from &, camaldulensis by the present authors.
Also, the oesophageal glands of the parthenogenetic
female of the specimens from £. camaldulensis are
longer than in Currie’s description and drawing of FL
rumufactens.

The structure of the digestive tract of Fereuvobia
remains unclear. Given the small size of these
nematodes, and their typical dark colouration, it is very
difficult to disuaguish the anterior parts of (he tract,
Fisher & Nickle (1968) described the anterior part of
the besuphagus as swollen, narrowing abruptly to form
a short tsthmus at the level of the nerve ring, then
broadening again to contain the large plinds, They
believed that the oesophago-intestinal junction occurred
at about the level of the secretory-excretory pore.
Siddig) (1986) interpreted the anterior swelling of the
digestive tract as a “pseudo-pharynx” and believed chat
ihe yalve-liKe #leuctnre it contuins marked the

TABLE 2. Measuremenis for urtulr femates uf Syrphonema ap.
All Theasurementy i jar

Length Width Aglerior Tul
end ti hase length
of bulb
mica a7U AY 0.5 6.0
n 12 2 0 0
Str 16 68 20.5 16.
ninge 1240-1890 29-5h W171 431u)

kK, A, DAVIES & IL LLOYD

junction of the oesophagus. and intestine. An electron
microscope study of the anterior part of the digestive
iract of Fergusobia is needed to decide which of these
interpretations Js correct

Syrphonema sp,
(FIG, 3h

Measurements: Table 2,
Descriprion of female (Pig. 3C, D, B, KG)

Nematodes straight or slightly C-shaped when
relaxed by gentle heat. Cuticle has longitudinal proaves
with many fine transverse markings; offen appears
loose at head and/or tail; lateral lines large (31 yan wide,
27-33 wim), with smooth ribbon-tike appearance and
single central ridge, Lip region often indistinct; six
separate lips, each wilh small papilla. Stoma cup-
shaped: cheilostom reduced; promesustom about 4 wnt
long, and about same width; thickening of cuticle an
ventral side of metastom, which makes stoma
asymmetrical, and may form a rhabdion; lelostom
present. Amphid openings at base of lateral lips.
Oesophagus rectilinear; no true bulb; vestigial valve
present. Nerve ring situated in postenur third of
oesophagus. Secretory-excretory pore opening just
behind perve ring: prominent excretory cell just behind
oesophago-intestinal junction. Deirids not seen,
Hemizonid just posterior to netve ring. Intestinal lumen
lined with refractive material from oesophago-intestinal
junction to rectal valve; lumen wide in young females
but narrower in older, Three rectal glands.
Reproductive tract with single gonad, amphidelphic,
will genital tube running anteriorly; ovoviviparaus.
long uterus, extending almost to point of Hexure of
genital tube just behind ocsuphago- intestinal junction,
oviduet extending back down dorsal side of body; small
posi-uterime sac present; no true spermatheca, sperm
not seen; vulva with iransverse opening. well-
developed associated musculature; posterior vulval lipr
often more praminent than anterior; vagina not direcred
ameriorly or posteriorly. Phasmids not secn. Tail
conical, terminus varizhle (Fig. 3)

isolated Jrom Adelaide.

Anterior Vv 4 b v
end 40
vulva.
(200 BLG 29:3 {0.6 38
12 12 12 ip iN}
22) 37 36 17 is
25-36 RX a Nay

YU-R20 -AES-RES

NEMATODES ASSOCIATED WITH DIPTERA 19
Description of juveniles (Fig. 3A, B)

Second stage juveniles 573 pm (473-624; n=8),
third stage juveniles 830 um (806-873; n=3): fourth
stage juveniles 1062 wm (920-1260; n=7).

As for adult females, except that the lateral lines
consist of a single central ridge only. Gonad primor-
dium well-developed in third and fourth stage juveniles,
developing uterus particularly obvious in fourth stage
nematodes, enabling rapid determination of the various
juvenile stages.

Collector, hest and localiry

The nematodes were dissected from the intestine of
two females of the syrphid fly Simosyrphus
grandicorinis (Macquart), collected on sow thistle,
Sonchus L, sp., atthe Waite Campus of the University
of Adelaide, Glen Osmond in January and December
1993 by Mr E. Soleyman, Nematode specimens are
held in the WINC 687.

Biology and general comments

A search of Helminthological Abstracts suggests that
this is only the second record of the genus Syrphonema,
erected by Laumond & Lyon (1971), and the first
outside France. Its occurrence in South Australia
suggests that the genus may have a cosmopolitan
distribution. Laumond & Lyon (1971) collected S,
intestinalis from the digestive tracts of 12 species of
syrphid flies. The infected flies found here were part
of collections made in a study of the biology of the
syrphid flies, 5. grandicorinis and Melangyna
viridiceps (Macquart). No nematodes were seen in
dissections of 305 M. viridiceps and only two of 105
8S. grandicoriniy dissected contained nematodes
(Soleyman pers. comm.) suggesting that the infection
rate is naturally low, It is not known what effect, il
any, infection has on the survival and reproductive
capacity of the fly.

The nematodes described here from South Australia
were classified as Syrphonema on the basis of the host
fly, rectilinear oesophagus without a bulb and with a
vestigial valve and because the female is ovoviviparous
arid has a posterior vulva. In the absence of males, it
is not possible to decide if the nematode is S.
intestinalis or a new species. The body lengths of the
South Australian and French forms suggest that the
former were smaller, but the De Man ratios are very

Fig, 3. Syrphonema sp. A, Entire fourth stage juvenile. B.
Anterior of fourth stage juvenile. C. Anterior of adult
female. D, Entire adult female. E. Variable tail shapes of
adult females. F. Vulva and tail of female, G, Stoma of adult
female. Scale bars = 10 um BCE FG, 20 um A, 50 pm D.

20 K. A. DAVIES & J. LLOYD

close (Table 2). Some apparent morphological
differences have been observed. Laumond & Lyon
described the stoma of their specimens as reduced and
“vestibule-formed”; in the nematodes described here
the stoma was cup-shaped but asymmetric with
cuticular thickening (possibly a rhabdion) of the ventral
metastom. The nerve ring seems to be located more
posteriorly in the South Australian than in the French
specimens. Again, the drawing of the female in
Laumond & Lyon (1971) does not show a post-uterine
sac Or a protuberant posterior vulval lip, both present
in the specimens examined here. While Laumond &
Lyon state that §. intestinalis does not have a
spermatheca, they have drawn a structure, also seen
in South Australian females, which could function as
a spermatheca. This is an apparent modification of the
reproductive tube, just on the uterine side of the flexure

of the oviduct; however, no sperm were seen. An
attempt to obtain material from France for comparative
studies was unsuccessful.

Acknowledgments

We thank Dr D. N, McAlpine, Australian Museum,
Sydney for identification of adult Fergusonina collected
from Black Forest, Dr J. Gardiner, University of
Adelaide, for indentification of FE. leucoxlyon, Dr W.
Frost for collecting leaf galls, Mr E. Soleyman for
specimens of Syrphonema and information on infection
rates and Mrs F. Reay and Mr G., Taylor for helpful
criticism of the manuscript. This work was supported
by a grant from the Australian Biological Resources
Study,

References

Curriz, G. A. (1937) Galls on Eucalyptus trees. A new type
of association between flies and nematodes. Proc. Linn.
Soc. N.S.W. 62, 147-174.

FisHer, J. M. & Nickie, W. R. (1968) On the classification
and life history of Fergusobia curriei (Sphaerulariidae:
Nematoda). Proc, Helminthol. Soc. Wash. 35, 40-46.

KRALL. E. L. (1991) Wheat and grass nematodes: Anguina,
Subanguina and related genera pp, 721-760 In Nickle, W.
R. (Ed.) “Manual of agricultural nematology” (Marcel
Dekker, New York).

Laumonp, C. & Lyon, J. P. (1971) Le parasitisme de
Syrphonema intestinalis n.g., n.sp., aux depens des

syrphides (insectes dipteres) et la nouvelle famille des
Syrphonematidae (Nematoda: Rhabditida). C. r hebd.
Sean. Acad. Sci., Paris, Ser. D 272(13), 1789-1792.

Maccent!, A. (1981) “General Nematology” (Springer-
Verlag, New York).

Sippigi, M. R. (1986) A review of the nematode genus
Fergusobia Currie (Hexatylina) with descriptions of F,
Jambophila n.sp. and FE) magna n.sp. pp. 264-278 In
Swarup, G. & Dasgupta, D. R. (Eds) “Plant parasitic
nematodes of India, problems and progress” (Indian
Agricultural Research Institute, New Delhi),

PRELIMINARY INVESTIGATIONS OF DUNES OF THE
GAWLER RANGES PROVINCE, SOUTH AUSTRALIA

By E. M. CAMPBELL*, C. R. TWIDALE*, J. T. HUTTON} & J. R. PRESCOTTE

Summary

Campbell, E. M., Twidale, C. R., Hutton, J. T. & Prescott, J. R. (1996) Preliminary
investigations of dunes of the Gawler Ranges province, South Australia. Trans. R.
Soc. S. Aust. 120(1), 21-36, 31 May, 1996.

Three fields of dunes have developed in the recent past within the Gawler Ranges in
the arid-semiarid interior of South Australia. The dunes (lunettes, parabolic dunes,
transverse dunes, linear dunes, climbing dunes and falling dunes) are essentially relic
forms, were active about 4000 years BP and are now stabilised by vegetation though
strong winds still cause occasional sand movement. Some of the dunes demonstrate
sand transport over distances of at least 25 km. The origin of the various
morphological dune types is discussed. Supply of sand, the moisture content of the
substrate, the vegetation cover and wind speed and direction are all important.
Topography influences the morphology of the dunes in various ways and is
fundamental to any explanation of climbing and falling dunes.

Key Words: Gawler Ranges, lunettes, parabolic dunes, transverse dunes, linear dunes,
climbing dunes, falling dunes, thermoluminescence dating.

Transactions of the Royal Sactety of §. Aust, (1996), LAO), 21-36,

PRELIMINARY INVESTIGATIONS OF DUNES OF THE GAWLER
RANGES PROVINCE, SOUTH AUSTRALIA

by E. M. CAMPBELL, C. R, TWIDALE* J. T. HuTTONT & JR, PRESCOTT

Summary

CAMPBELL, E, Mo, Twpalr, C. RL, Huron, J. 1 & Prescorr, J. R. (1996) Preliminary investigations of dunes
of the Gawler Ranges province, South Australia. Trans. R. Sov. S. Aust, 120(1), 21-36, 31 May. 1996,

Three fields of dunes have developed inthe recent past within the Gawler Ranges in the arid-semiarid interior
of South Australia, The dunes (lunettes, parabolic dunes, transverse dunes. linear dunes, climbing dunes and
(illing dunes) are essentially relie forms. were active about $000 years BP and are now stabilised by vegetation
though strong winds still cause occasional sand movernent. Some of the dunes denioastrate sand transport over
distances of ut least 25 km.,. The origin of the various morphological dune types is discussed, Supply of sand.
(he moisture content of the substrate, the vegetation cover and wind speed and direction are al! important. Topography
influences the morphology of the dunes in various ways and is fundamental to any explanation of climbing and

falling dunes.

Key Words: Gawler Ranges, lunettes, parabolic dunes, transverse dunes, linear dunes, climbing dines, falling

dunes, thermoluminescence dating,

Introduction

In the mid-latitude deserts extensive fields of sand
dunes are restricted to plains. Sand dunes have,
however, been reported from desert uplands where
topographic obstacles deflect or funnel ihe regional
airflow and produce depositional forms and patierns
different from the essentially regular and repeated
formations found in the dunefields of the adjacent
plains (Wilson 1973; Smith 1982). They include sand
shadows of various types, sand sheets, obstacle dunes
and climbing and falling dunes (Planhol & Rognon
1970; McKee 1979; Mainguet 1984; Greeley & Iverson
1985). The Gawler Ranges, located in the arid-semiarid
interior of South Australia, 1s a desert upland within
which three fields of sand dunes have penetrated the
valleys between the bornhardt massifs and in some
areas have overridden the low domical hills (Fig. lu, b).

Geologic Background

The bornhardts of the Gawler Runges are developed
ina Jayered sequence of silicic yoleanic rocks (mainly
rhyolites, rhyodacites and dacites) of Mesaproterozoic
age (1592 + 2 Ma- Fanning ef al. 1986). The volcanic
rocks consist predominantly of subaerially erupted
ignimbrites (nuées ardentes deposits), welded to
varying degrees, and with local occurrences of basaltic
lava. and agglomerate. They were intruded by granites
of the Hiltaba Suite (1485 + 16 Ma - Creaser 1989

* Department of Geology and Geophysics, University of
Adelaide S, Aust. 5005,

+ Deceased.

+ Department of Physics, University of Adelaide S, Aust.
5005

cited by Blissett e7 al. 1989; see also Flint 1993) which
now occur extensively in the western part oF the upland,
in the Kondoolka and Hiltaba areas, as well as in small
isolated Outerops near Kokatha Homestead (H.S.) and
Lake Everard H.S. They are also exposed to the W,
SW and 5 of the Ranges.

Where exposed, both the volcanic and granitic
crystalline rocks are massive and compact but a well
developed system of orthogonal fractures trending
NNW and NE, and including also latitudinal and

tae AGoraeera
@& can harms | ~ ~ =
hanno | .
at _ a
2b hatyet ain 4 é a )
es it cer
Quiles x 4 |
\sertake kvernidy ¢ |
Pte ape he
gi = £" S S 4, (daira
eh 4 ‘ 4) 5 ots i
"= | ATs mn
“Meant Nt |
esti
Heataull 1 > \
PALAU Hm are’ |
<b oe E taenrt . \ ,
ena) = =) tee
an — i Ai ots . ° | ie Frrelane
reiting J ; \
% /
hike t - (
Jona
4 ’ WN
woh Sy
. &
¢
r mnt
= sie e |
Wrwlla
tin (arn \
Oe M, tani
= bigitey - -
SJukey Blutl
u = Bul hin ti iy
LT oe ~

NW Stivy

Fig. la. The Gawler Ranges province, showing location of
the study site in South Australia (inset), localities mentioned
in the text and average annual isohyets (mm), The
Corrobinnie Depression dune samples analysed by Gostin
were collected from dunes adjacent to the road from
Wirulla to Hiltaba HS.

22 &, M. CAMPBELL, C, R, TWIDALE, J. T. HUTTON & J, R. PRESCOTT

meridional sets, has been explomted by weathering and
erosion to form the major valley systems of the Ranges
(Campbell & Twidale 1991), The summit and upper
slopes of the bornhardts are essentially devoid of any
Weathered materials, though isolated patches of regolith

T] Satin
Py] Gririte
es Gewler Range
Hd Vetor piles

i a,

nt
————

5

Fig. lb. The geology, including dunefields, of (he Gawler
Ranges province (after Williams 1994). A, Hier Dunefield,
B. Precadilly Dunefield. C, Moonaree Dunefield. B
Beacon Dunefield. E; Mkina Dunefield F. Serubby Peak
Dunefield,

iin ym

3 A

‘se aks | iukn
wo \) tote Farrer

ae) 1 %

ee % I
Ad “ne \ Lf 4 hee fa ‘
. Laer fs anae if” 4
, wQ Meraamare a 94) et us ~ fi

mL 5

Mee)
; ow al
at
\ “rit
4 oe
i ~ — bien
_ uf hinger dune
a) hardboll) dupe
= Traisverse bide
= 3 A L y tall
r Shem
_— 45 ™~. CC | Quneteld

Fig. Ic. Schematic diagram showing location of lunettes, linear
dunes and parabolic dunes of the Gawler Ranges province,
South Australia, The orientation of the linear dunes is also
indicated. Not all dunes are shawn.

"CAMPBELL, EB. M. (1990) Structure and surlace in the
Gawler Ranges, South Australia. PhD thesis, University
of Adelaide (Unpub.)

are preserved on the lower slopes, as for example 7
km east of Nonning H S., and in the valley floors drill
cores and dam sections indicate that the volcanic:
material is weathered to depths of up to 50 m.
Individual bornhardts attain heights of over 460 m in
the S, but they decline in elevation to the N (e.g. Nukey
Bluff ¢, 460 m, Mt Nott 430 nm, Bond Aill 336 m,
Chitanilga Hill 317 m, Nueckulla Aull 256m, Mortimer
Hill 232 m ~ see Fig. ta). The upland is hounded on
its SW and § sides by an escarpment which, though
dissected and tisiiy wooly 150-200 m above the
surrounding plain, is nevertheless, and viewed
renionally, linear and imposing, so much so that the
explorer BE. J. Eyre was Jed to describe the Gawler
Ranges us “a vast MOUNTAIN Mass rising abruptly ow
of the low scrubby country” (Eyre 1845). To the N_
W.and E the relict amplitude diminishes and the hills
peter oul as the plains become more and more
extensive.

The bornhardts. of the Gawler Range province have
evolved in (Wo major stages, The first involved
planation and fracture-controlled differential subsurface
weathering m Jurassic ot earher Gmes, the second, the
stripping of the regolith in the Early Cretaceous to
expose bevelled domical forms. Remnants of the etch
planation surface represented by the summit bevels are
particularly: Well preserved in the south, There has been
only minor erosion of jhe ranges sinve the Mesuzoiv
(Campbell 1990'; Campbell & Twidale 1991).

Numerous yalinas, including Jakes Gairdner, Harris,
Everard and Acraman and many small saline playis.
occupy low lying areas in valleys and plains. The
depression occupied by Lake Acraman 1s the site of
an ancient meieorite impact (Williams 1994). The large
lakes stand about 120 m above sea level, They are the
termini of elosed drainage basins. most of which are
not much larger than the lakes themselves.

Climate

At present the area is semiarid in the § grading to
arid in the N (Fig. Ja). Jn the south there is. a pro-
nounced winter rainfall maximum, produced predom-
inanuly by the easterly passage of cold fronts, but a
significant proportion of the rainfall is derived from
occasional influxes of moist tropical air. In the north
the rainfall is derived from. both these sources and the
annual distribution is more uniform (Fig. 2a; Bureau
of Meteorology 1993a). Sammers are hot and winters
cool. Annual evaporation is about 2700 mm, with a
mean monthly evaporation ranging from about 80 mm
in June to 390 nun in January.

The only records for modern wind regimes in the
area are from Nonning (Fig, 2b). In summer, sand-
moving winds (stronger than about 20 km per hour)
blow predominantly from SE, § and SW. In winter,
strong winds blow from several quarters but N. NW

DUNES OF THE GAWLER RANGES

and SW winds are important (Bureau of Meteorology
1993b). Woomera, located more than 100 km ‘to the
NE of the Ranges, experiences predominantly SE-S
(total) winds in summer although in winter, winds are
more variable, but with a strong N and NW-SW
component. At Ceduna, 100 km to the W, in surrumer
SE, S and SW winds predominate, whilst in winter
NE through NW-SW winds are most common (Bureau
of Meteorology 1988) Fig. 2c).

The Dunefields

The Australian dunefields form a huge whorl of
linear dunes around the centre of the continent (Brook-
field 1970; Wasson et al, 1988), though it is not estab-
lished whether all the sectors of the pattern were
formed by winds related to one and the same
atmospheric system or that they were ever active al
the same time. The Australian desert dunes are
characteristically long, parallel sand ridges extending
unbroketi over tens and even a few hundreds of
kilometres. Many are asymmetrical in cross-section
and display tuning fork or Y junctions. They are
generally restricted to the desert plains (Madigan 1936,
1946, Wopfner & Twidale 1967),

Although the precise mechanism of formation is
debated, and it is likely that the dunes originate in
various ways (McKee & Tibbetts 1964; Woptner &
Twidale 1967; Brookfield 1970; Tsvar 1989; Tseo
1993), some are apparently initiated in the lee of
lunettes or other accumulations of sand (Twidale 1972)
and extend in a downwind direction. There is a mild
controversy as to whether the sand of which the dunes
are built is of local derivation (Folk 1971; Wasson 1983)
or Whether it is essentially exotic and far-travelled
(Wopiner & Twidale 1967), The Gawler Ranges
dunefields yield evidence germane to this problem.

Immediately to the W of the Gawler Ranges the
linear sand ridges of the Great Victoria Desert, the
southern part of the great Australian dune pattern, trend
WNW to ESE and there are zones of parabolic dunes.
Within the Gawler Ranges province the sand dunes are
more varied and Junettes, transverse dunes, and
climbing and falling dunes, as well as linear and

MOONAHEE NONNING
ao 30
Mean munthl r Mean nirinthily
taintall ¥ Th tal tall
+ ¥, ¥
E40 | 30
5 -
2 | | Mean no
ai rain days
197) Mean no. ox Ti 1”
rain uays S
=

TEM AMT ASOND
Months:

Fig. 24. Mean monthly rainfall and mean number of rain days
for Moonaree and Nonning (Bureau of Meteorology 1993a).
Length of record: Moonaree - 108 years. Nonning - 90
years,

Months

ro
le

NONNING
January July

N

) 4-29 >20 km/hr

Fig. 2b. Nonning wind data (Bureau of Meteorology 1993b)
The percentage of calm observations is indicated in the
centre of the rose. Length of record 23 years.

WOOMERA

January

CEDUNA

January

N

i) 41011-30790 kiv/hr

Fig. 2c. Wind roses for Woomera and Ceduna (Bureau of
Meteorology 1988),. The percentage of calm observations
is indicated in the centre of the rose.

24 E, M, CAMPBELL, C. R. TWIDALE, L T. HUTTON & J. R. PRESCOTT

parabolic dunes, are developed. Some are stable, but
others are occasionally mobile.

Three fields of dunes penetrate the uplands (Fig. Ib).
In the N, the Hiern Dunefield extends WNW to ESE
between the Kokatha hills and Lake Everard to the
western shore of Lake Gairdner. The dunes are
predominantly linear forms. Dunes also occur N of
the Ranges and also on some of the islands within Lake
Gairdner. In the same latitude, and to the lee of a major
lunette developed on the E shore of Lake Gairdner,
the Piccadilly Dunefield extends eastwards for 35 km
across the plains located between Lake Gairdner and
Island Lagoon. Here linear sand ridges and parabolic
forms are well developed and some lunettes occur on
the eastern margin of small salinas.

The Moonaree Dunefield occupies the plain between
the volcanic Everard hills to the N and the granitic
Kondoolka hills to the § and extends eastwards to Lake
Acraman. In this part of the Moonaree Dunefield there
is a Sharp boundary between parabolic dunes to the
S and linear sand ridges to the N. Dunes occur on
islands within Lake Acraman and on its eastern shore.
To the NE of Lake Acraman the plain carries a veneer
of sand but dune forms are absent. Further to the E,
however, linear sand ridges are again developed and
extend as far as the shore of Lake Gairdner.
Immediately to the W of this salina, some of the dunes

“smitH, D. M. (1976) The denudation chronology of the
southern Gawler Ranges and adjacent areas, MA thesis,
University of Adelaide (Unpub.)

override the low volcanic hills forming climbing and
falling dunes. The Beacon Dunefield (the Black Oak
dunefield of Smith 1976") extends eastwards from the
E shore of Lake Gairdner, again in the lee of a major
lunette. This field consists mainly of linear sand ridges
but there are some lunettes and parabolic forms.

The most southerly dunefield, the [kina Dunefield,
is part of the Kododo Dunefield of Smith (19767).
Both the field and individual dunes trend NW to SE
between the Corrobinnie Depression (Bourne e7 al.
1974, Binks & Hooper 1984) and the SW margin of
the Gawler Ranges from near Yarranna Hill to the
vicinity of Mt Sturt. Within the Corrobinnie
Depression, complex parabolic forms are well
developed. In the vicinity of Mt Centre, lincar sand
ridges trom the Ilkina Dunefield diverge ESE and
extend across narrow plains and valleys between the
volcanic uplands and extend into the hilly areas to form
the Scrubby Peak Dunefield (Fig. 1b). In both the N
and S arms of this dunefield there are departures from
the general ESE trend as a result of topographic
interference with the airflow. Both crestal transverse
dunes and climbing and falling dunes result from such
topographic effects.

Dune morphology

Linear dunes
Linear sand ridges dominate the dunefields within

the Gawler Ranges (Fig. 3). These linear forms trend
WNW to ESE in the W and latitudinally further to the

Fig. 3. Linear sand ridges of the Scrubby Peak Dunefield funnelied along broad valleys between the bornhardts of the southern
Gawler Ranges, South Australia, Field of view in foreground approximately 5 km.

DUNES OF THE GAWLER RANGES 25

E, In places, e.g. near Mt Granite (Fig, 4). funnelling
of the Wind has produced dunes aligned at various
angles to the regional trend. ‘The linear dunes vary in
height, length and linear frequency, i.e, the number
of sand ridges per unit distance measured normal to
the dune trend, The maximunt height of the dunes
varies from 5-15 m above the interdane corridors. They
vary in length from a few tens of metres up to 20 km,
none extends unbroken for many scores or hundreds
of kilometres as do some of the sand ridges of desert
plains such as the Simpson Desert (Wopiner & Twidale
(967, 1990; Twidale |981). The lincar frequency of the
dunes varies between two and six per km The
interdune corridors are sand covered. Most of the dunes
are symmetrical, with smooth crests which rise and
fall to form peaks and saddles, The slopes are gentle,
considerably less than the angle of repose of the sand.
No deposinonal structures and no slip faces have been
noted. The dunes carry a covering of low shrubs and
amall trees, though little or no soil development is
apparent and there is, today; only occasional and minor
reworking of the sand by wind and water, The dunes
are relic according to the classification of Livingstone
& Thomas (1995),

Parabolic dunes

Groups of parabolic or U-dines occur within the
linear dunefields. Most of the parabolic dunes occur
outside the Ranges, and notably in. the Corrohianie
Depression (Bourne er a/. 1974), though there is a W-
E zone within the Moonaree Dunefield and patches

\
i
'
r
N \ wu
‘
"OME tinatite, !
} Se
Py
)
I 2 ‘
\ \
j
'
|
ra | |
iv. \, MN 7 2 yn ;
Playas "y \ of,
aN ae On| ' oS
> Duras ac i thoy
'0U. Com itours it |
wr (ey é
ae \ rl |
— fm \ : |
—_ H i} «| a |
kn a “\ _

Fig. 4. Schematie diagram of linear dunes oriented NW tu
SE, and irregular patterns of linear and transverse dunes
due to topographic interference to the wind near Mt Granite,
Gawler Ranges, South Australia (from aerial photographs,
Department of Lands, South Australia and 1100 000
National topographic map series). Not all dunes are shown

of parabolic forms occur within the Piccadilly and
Beacon dunefields. Although many of these dunes are
complex.in plan form, with transverse, rake-hke and
circular patterns well developed, the basic unit is a U-
shaped dune about § m high and with the open end
of the U pointing to the W (Fig. 5).

Climbing and falling dunes

In the Gawler Ranges most of the dunes are
developed on broad valley floors between the
bornhardts. In some areas. however, linear dunes
penetrate into the hilly terrain and suffer modification
asa result of funnelling and diversion of the wind (Fig-
4). In other areas, the dunes extend over the bormhardts
On the reasonable assumption thal the sand migrated
southeastwards, dunes piled against the windward
(northwestern) slope of a hill are termed climbing, or
rising, dunes; Where sand has overridden the crest ot
a hill and extended on to the leeward (southeastern)
slope, falling, or hanging, dunes are formed (Fig. 6a).

i) a Playas

~ Dunes

~~ _
== Roads

< & Contours my
3

{ }
Hy
ee

VLA

1,
/y

Fig, 5. Linear and parabolic dunes of the Moonaree Dunefield
15 km south of Lake Everard H.S. (from aerial photographs
Department of Lands, South Australia and 1100 000
National topegraphic map series),

Fig, 6a. Echo, climbing and falling dunes (after Mabbutt
1977). Arrow indicates direction of the wind, A. Linear
dune not anchored by topography. B. Linear dune rising
over topographic obstacle, C, Climbing (1) and falling (27
dune. D. Echo dune.

16 E, M. CAMPBELL, C, R. TWIDALE. J. T. HUTTON & J, R, PRESCOTT

These dunes have oot been studied in detail in the arid
mountains of Australia, although dunes which arguably
ascend cliffs have been studied in the coastal context
(Jennings 1957: Langford-Smith & Thom 1969).

Climbing and falling dunes are known from various
parts of the world, for example from periglacial Finnish
Lapland (Seppila 1993), coastal west Galicia, Spain
and NE Spain (Cros & Serra 1993), but mose
previously published reports pectain ta warm desert
environments, é.g, Califorma (Evans 1962; Smith 1967
cited by Bender 1982; Anders 1974 cited by Bender
1982; Lancaster 1994), Colorado (Johnson 1968),
Idaho (Koscielniak 1973°), Arizona (Greely & Iverson
1985) und Utah (Alhbrandt 1979), all in the United
States, where most are inactive, veneered by gravel and
dissveted by ephemeral streams (Smith 1982), nerthern
Mexico (Stone 1967), Brazil (Biyarella 1975, 1979),
Egypt and Jordan (McKee 1979), the Sinai Peringuls
(Ahibrandt (979), the Sahara (Smith 1954) and the
central Namib Desert (Goudie 1972). They are also
found in the eastern Flinders Ranges, South Australia
(Green 1994*), near Port Stephens and in the
Shoalhaven River area in New South Wales (Thorn e7
gl, 1994), on the Eridunda Range 160 km south of Alice
Springs and on the northern margin of the Simpson
Desert where dunes override some of the fatitudinal
ranges. Greeley (1985, Fig. 7.39) illustrates a field ot
climbing dunes drifting over the rim of @ 16 km
diameter crater on Mars.

In the Gawler Ranges climbing and falling dunes
occur in three areas. Firsi, examples were noted by
Smith in the Senubby Peak Dunefield (19767; Fig, 6b,
c), Second, Giles (980) remarked that sand dunes
encroach On to the slopes af Mt Sturt. Sand from the
kina Dunefield has accumulated on the NW slopes
of Mt Sturt (the western peak) and forms an irregular
mound along the base on its SE side. Third, climbing
and falling dunes aré common in the Moonaree
Dunefield E of Lake Acraman, Where stall dunes
trending W-E are essentially restricted to the plains,
though they partially override many of the bornhardts
(Fig. 6d)-

In the Scrubby Peak Dunefield, some linear dunes
have been diverted around the major volcanic bills (Figs
4, 6b,c) but elsewhere, especially where the relief is
lower, the dunes traverse hill and valley alike. The
dunes ascend the lower hills (in general terms those
that stand less than some 40 m above the adjacent valley
floors) without significant interruption of form and are

Roscigumiak, D. B. (1973) Eolian deposits an a voleanic

terrain pear Saint Anthony, tdahe. MA thesis, University
of New York (Unpub,)

4Greene, 8. J. (1994). A geomorphological and sedimento-
logical study of a climbing dune, northern Flinders Ranges,
South Australia. BA (Hons) thesis, University of Adelaide
(Unpub.)

AGiLes, C. W. (1980) Spring Hill, southern Gawler Ranges.
Geol. Soc. Aust. SA. Div Geological Monument Ill, File
E 20 (Unpub,)

classed as climbing dunes. In some instances the dune
is diverted around the flanks of the hill and continues
downwind (Fig. 6b), Elsewhere, the dune forni 1s
interrupted, for although there are many grains and
even small pockets of sand in fissures and shallow rock
basing on the crests and upper slopes of the hills, there
is no dune form; a short distance downslope fram the
crest, however, the dune form is resumed an falling
dunes 3-4 m high (Fig. 6c).

Transverse dunes

In the Serubby Peak Dunefield funnelling of the wind
has produced dunes of varied orientation, In this part
of the Gawler Ranges elongate bornhardts are aligned
exsentially N-S. Sand ridges also aligned N-S are
located in the valleys between the bornhurdts. There
are some W-B dunes which override the bornhardts
and, in addition. N-S trending elongate dunes are
located just below the crest on the lee side of these
hills (Pigs 4, 7). These crestal dunes ure tentatively
classified a5 OF Iransverse type,

Lunettes

Lunettes are developed along at least part of the E
side of most of the lange Salinas and many of the smaller
playas in the region (Fig. ic). Lunettes are transverse
dunes located on the lee shores of lake basins. The

name “lunectie™ was first applied to the form by Hills

sh “AN Ss ; ‘|
oN Re oA Divertaa

“ON, WW \ nn, tow (1) |
A. Si Lay x
_ — ae, .
te eR! =F
ee
Climbing Loe
ue (C) \ .
%
1? Sthnly ote :
Cd Hil tiewior Bani vetttorier:
[J Sand dune y
Tailing «
chirws (F}
Oe ae
en aes
ee ncomer this ~—e _
Ww Pa
cress]
an
ps Can
a Saf
¥ z

Fig. 6b, Scrubby Peak area, Gawler Ranges, South Australia.
showing diverted dune (D), climbing dune (C) and falling
dune (F) (from 100 000 National topographic map series).
The dune-forming wirids were from the northwest sector:

DUNES OF THE GAWLER RANGES

Fig. 6c. Diverted dune (D), climbing dune (C) and falling dune (F), Scrubby Peak area, Gawler Ranges, South Australia.
View to the north. The hill stands about 25 m above the surrounding plain.

Fig. 6d. Climbing and falling dunes, Moonaree Dunefield, Gawler Ranges. South Australia, Note that the falling dune,
on the near side of the bornhardt, resumes in a topographic embayment. Field of view approximately 2 km.

2s E. M, CAMPBELL, CR. TWIDALE, J.T. HUTTON & I. R. PRESCOTT

(1940) who deseribed lunettes of silty-clay
compositions from NW Victoria. Subsequently,
Juneties OF various sizes und mineralogies have heen
reported from all states of Australia. They range in
composition from quartz-rich to clay-rich to almost
pure gypsuin. The sandy guartz-rich dunes were
formed hy deflation from beaches on the lake margin,
The clay-rich diines were derived by. deflation of clay
ageregates fram {he saline lake floors. The gypsum.
dunes are composed either of rounded crystals deflated
from the dry jake bed or of fine ‘knpi, some of Which
may be due to Weathering of saliated particles since
depositiin (Stephens & Crocker 1946; Campbell 1968;
Bowler 1968, 1983; Chen ef al, 1991).

In the Gawler Ranges area lunenes of gypseaus
composition oeeur an the eastern margin of lukes
hyverard, Hatris, Aeraman and of many of the smailer
salinas. Both gypseoug and sihceous lunettes are found
om (he eastern side of Lake Gairdner. The most
prominent silictous lunettes are located opposite the
dunefields which impinge on the W side of the lake.
Both dunefields found E of Lake Gairdner, the
Picvadilly and the Beacon, are developed in (he lee
of these prominent siliceous lunetles, The lunette on
the NE margin of Lake Gairdner rises about 35 m
above the Jake bed. Much of the surface is bare and

erosion by wind and water hus created a series ol
darnical remnants. standing 3-4 m above gentle swales,
ln addition. tunettes consisting predoniinantly of
fragments of Gawler Range Volcanics of sand sive
occur discontinously along the margin of Lake
Gairdner (Fig. te).

Sedimentology

A total of 16 sand saniples, each from the crest of”
a dune, and including at least one from each of the
dunefields in the Gawler Ranges province, was
examined to determine cumipasition and grain
morphology (Table 1) and grain size and related
parameters were determined using 0,5 phi standard
sieves,

The saril samples are all various shades of yellow-
red (2.5 to 10 YR Munsell Suil Colours). All samples
consist of at least 90%, and most more than ¥8%,
quartz grains. The biiner constituents are quartz tock.
feldspar, Gawler Range Voleanies fraginents. ?iron
oxide and organic matter, In most samples the grains
arc predominantly frosted, but sume are polished
Saniples from two dunes in the Serubhy Peak Dunefield
show higher percentages of polished prams. Grains
{rom all samples show lerruginous coatings of yellow,

Fig. 7. Transverse crestal dune, Scrubby Penk Dunefield. Gawler Ranges, South Australia. View to the south to the Corrohinnie

Depression. The crestal dune is abour & m high.

DUNES OF THE GAWLER RANGES

Sample Dune type — Colour! Composition? — Surface Surface Roundness> —_—-Sphericity
texture? coating®
Beacon Dunefield
! linear 2SYR 5/6 quartz 95% 85% ME Y,O,R,B. SRSA, high,
(1,2,3,4,5) Is% P few R same eclongute
Piccadilly Dunefield
2 linear SYR 4/6 quartz, 99% 98% MF YO SA-SR, high,
(1,2,4.5) few P few R some elongate
3 parabolic SYR 5/6 quartz 99% Ono ME ,O SA-SR, high.
(4,5) few P few R, WR some elongate
4 linear 25YR 4/8 quartz 99% 95%. MF Y,O,R SA-SR, high,
(4,5) few LF fow R some clongate
few P
5 linear 2.5YR 4/8 — quartz 98% OR% MF YR. B SA-SR, high,
(1,5) 2% P few R some elongate
Hiern: Duinesiets |
6 linear SYR 5/6 quartz 99% 98% F Y, 0 SA-SR, high,
(1.4.5) 2% P few WR some elongate
7 irregular SYR 5/6 quartz. 99% 95% F Y,0 SA-SR. high,
(4.5) 5% P few R some elongate
8 linear TSYR 6/6 = quartz 999), 95% MF Y,O SA-SR, high,
(1,4,5) 5%P few R, WR some elongate
9 linear SYR 5/8 quartz 90% 90% F Y,O A-SR moderate,
(1,3.4,5) 10% P some R some high
Moonaree Dunefield
10 linear TSYR 5/6 quartz 98% 90% LF y, O SA-SR, high,
(1.4,5) 10% P some A, R some elongate
ul irregular 7SYR 5/6 = quartz 99% 95% F Y, 0 SA-R, high,
(4,5) 5% P few WR some elongate
Scrubby Peak Dunefield
12 linear 7SYR 5/6 quartz 95% 95% MPF YR, B SA-SR. high,
(1,4,5) 5% P few R, WR some elongate
3 linear 1OYR 5/4 quartz 99 % W% MP Y,O,B SA-SR, high-moderate,
(1,4,5) 30% SF some R some elongate
I4 linear 1OYR 6/4 quartz 99% 50% P Y, 0 SA-SR, high-moderate,
(1,4,5) 50% F few R some elongate
15 linear TSYR 4/6 quartz, 9% % 98% F Y, 0 SA-SR, high-moderate,
(1,4,5) 2% P few ALR some elongate
Ikina Dunefield
16 linear TSYR 6/6 — quartz 98% 70% LF Y,O.R SA-SR, moderate-high,
(13,4) 30% P few R sume elongate

‘Munsell Soil colours,

“Minor constituents in brackets: [: quartz rock. 2: feldspar. 3

‘F: frosted. P: polished. M: moderately. L: lightly.
*Y: yellow. O: orange. R: red. B: brown.

> Gawler Range Volcanics. 4: ?iran oxide. 4; organic material.

“SA: subangular. SR; subrounded, A; angular, R: rounded, WR: well rounded

and less commonly orange, ted and brown, material,
The grains in all samples are predominantly subangular
to subrounded, with small amounts of angular and well-
rounded grains. High to moderate sphericity is char-
acterisuc, with most samples containing same elongate
grains.

The dune samples are all fine to medium grained
sands (mainly 0.125 to 0.5 mm diameter - Folk 1968).
They are well-sorted to poorly sorted, with most
samples moderately well-sorted.

Age of the dunes

The sand of the Gawler Ranges dunes is typically
a yellow-red colour (Table 1), suggesting sufficient time
for initial weathering of clay, release of iron, and
development of a faint ferniginous patina. The sand
is not the brilliant red of the deserts of central Australia,
nor the dusky red (IOR 3/4 Munsell Soil Colour) of
the sand derived from the local Gawler Range
Volcanics. Some authors would attribute the contrasting

40 EE. M. CAMPBELL, C. R. TWIDALE, J. T. HUTTON & J. R. PRESCOTT

colour to different source materials (Wasson 1983:
Nanson eta/. 1992). Others. e.g, Wopiner & Twidale
(1967) and Walker (1979) consider that the intensity of
the red colour increases with time and hence is an
indication of the age of the dunce.

In ain attempt to attain a more precise estimate of
age, sand from the Scrubby Peak Dunefield was lested
for thermoluminescence (TL). Samples were taken
from a falling dune on an unnamed hill (National
Topographic Map Series Minnipa 5932, 12100 000,
Grid Reference NE3I6018) 1.5 km E of Scrubby Peak
(Fig. 6b). The method wasa variation on the “partial
bleach” method developed for the TL daling of
sediments by Wintle and Huntley (1982). ‘The age a4
estimated by measuring the TL energy stored in the
lative of @ suitable mineral, in this case, quartz. The
time interval measured is he time since the stored
energy Was last resel to zero Or near Zero by exposure
to solar ulfra-violet radiation. After such a re-setting,
energy accuinulates again ala known rate by exposure
fo midiation in the environment from the naturally
radioactive elements K, U, Thand from cosmic rtys
The aye ts found from the so-called age equation:

age = natural TL

TL per unil dose * dose rate.

Samples were recovered from depths of 35 and 70
cm within the dune by means of an aliger, taking care
to shicld the sample from light during and after
collection. After digestion with 20% HCI 1a remove
carbonates and NaOH to remove clay, the 90-125 am
Iruction Was recoverd hy sieving. A 40 minute etch with
40% HP removed feddspars and a $urlace layer of the
yuitrly, Plolation on aquenus sodium polytungstate al
w relative density of 2.67 followed; the end product
was very pure quarts and it is an this sample chat the
measurements were carried our

One of ie problems wath TL dating of sediments
is Unceralty obeur the degree to which the TIL wis
feset al the beyianing of the time of interest. 11 is mere
for the TL to be removed completely, even by
prolonged exposute to sunlight. Morewver, the amount
Of rele TL varies trom sample to sample and muy vary
with the age of the sample(BeTger 1990), In the present
investigation, i was land that the wecumulated TL
was smull so (hal any uncertainty in the degree uf
reseiting would result in signifewnt uncertuiary om the
age. The level ot resetting Was found from 2 surtace
sumple collected hy pressing packing type against the
exposed dine Surface This showed that the TL clack
had not been coinplerely reset ty zero in spite of Lhe
long time likely to have been spent in the sun by the
sample in reaching WS present posilion. Under these
circumslances Special procedures are nevessary. ws
deseribed by Preseott and Mojarnabs (1893). They make

use Of the tact that many quartz samples have a sc-
called “rapidly bleaching” peak (RBP) at 325°C in the
thermoluminescence glow curves, which bleaches le
near zero Within a wiatter of Minutes When exposed
to light of wavelength longer than about 500 nm
(Spoaner ef al, 198%), This means that exposures uf
the order of minutes fo fatural sunlight in the
environment will have ensured that the trap concerned
had been emptied completely and that, at least-so far
as the 325°C peak is concerned, the TL clock af the
sediment was completely reset at the time of
deposition. Inaddition, this peak emits ina wavelengts
band centred near 420 nm, so that an oplical filter
transmitting this band will be selective for the peak
in question (Prescot & Fux 1990; Scholefield ef ai,
1994), The 325°C peak rides on an unbleached
background, hich is measured and allowed for by ue
procedures, The surface samople menuoned above hat
zero TL when measured with the revised procedures.

The TLuis expressed in terms of an equivalent dose
measured in grays (Gy). The equivalent doses are: fir
the 35 cm sample 1,24 + (1.20 Gy; and far |he 70 cm
sample 1.53 4+ 0.25 Gy, The dose rate has been
measured by three essentially independent methods
(Hutton & Prescott 1992). They are, with the relevant
dose rates in brackets: in sitn gamma ray spectrometry
(0.153 + 0.028 Gy ka!); thick source alpha counting
for U and Th with X-ray spectrometry (XRS) for K
(U.142 + 0,029 Gy ka"): and delayed neutron analysis
(DNA) for U. neutron activation analysis (NAA) for
Th with XRS for K (0.168 + 0.041 Gy ka!) The
weighted average is 0.152 + 0.010 Gy kat for both the
35 em and 70 cin saaiples.

Contributions for cosmic rays must be included.
These are 0,21 + O02 and (M18 + (02 Gy kw? for
the 35 and 70 cm samples respectively (Prescoit &
Hulton 1988, 1994). fis worth noting that cosmic rays
dominate the dose rate because the levels a! K, U and
Th are so extremely low (K- 0.04 + 001%; LE O22
© 006 npn, Th- 1.0 + O04 ppm). Over ihe time in
question, changes in Cosme ray antensilicy are
negligible (Prescott & Hatton 1994),

The dose rates are (135 + (103 Gy ka! at 34 cm
and 0.33 + 093 Gy ka! ar 70 cm, A contribanen
from systematic errory fas been added, Hence, the age
of the 35 em sample is 3.7 + 07 ka and of the 70 em
sample 4,6 + 0.9 ka. Although the deeper sample has
the preater TL age, the (we ages aro nor statistically
difterent and probably all that can be concluded is thue
the dune has beenin place for abont 4 ku

This ave is based on a single series of dates frum
ane dune, Obviously, more age determuiiations are
required, Neverttteless, the pale colour of the sund,
to Whith previous reference hus been made, and the
lack of any carbonnte accumulations in the dunes
(despite its avadlab|lipy) ate sugeestive of a youthful age,

DUNES OF THE GAWLER RANGES uM

The general appearances are consistent with the TL
dating in Suggesting that the Gawler Ranges dunes are
younger than the putative Late Pleistocene relic forms
of NW Eyre Peninsula (Twidale er al, 1976) and are
comparable to the Holocene forms of that area (Rankin
& Flint 199)) and of the Simpsou Desert (Woplner &
Twidale 1988, 1990). Nevertheless, whe age
determination obtained is for the uppermost layers of
a dune and there is no evidence of the age ol the sand
at the base ot the dune,

Origin of the dune sand

As mentioned previously, the provenance of the sand
in dunefields, whether it is of local derivation or far-
travelled, 18 controversial. The question can be clantied
by a consideration of the sedimentologic characteristics
of the dune and other sands, In the Gawler Ranges
provinee, a local origi of the dune sands ts preeluded
by their composition and granulometry as set out in
Table | For exarnple, the Scrubhy Peak Dunetield
overlies outcrops of Bucarro Dacite, and Yardea Dacite,
with small areas of Yannabie Rhyodacite, Paney
Rhyolits and, at the base ot the Yardea Dacite, “black”
dacite (Blissert er al. 1988, Pig, 8). The microscopic
groundmass of the volcanic rocks is rich in quartz, but
the grains are much smaller than those of the dune
sands. There ure no quartz phennerysts in the dacite
(Blissett 1986), No quartz of a size equal la, or preater
than, that of which the Scrubby Peak dunes are
composed (and hence susceptible to attritian to produce
sund-sized grains) could be derived from the Yardea
and Bucarro dacites which are the country racks overt
which, overwhelmingly, the dunefields have extended.
No lakes or streams which might constitute 4 possible
source of sand in the dunes are known trom within
the province,

If it is accepted that the dunes of the Scrubby Peak
and other dunefields of the western Gawler Ranges
extended trom the W or NW, then there are three other
possible sources of the quartz sand.

First, there are outcrops inwthe western Gawler
Ranges of Yannabte Rhyodacite and of Paney Rhyoliie
(Fig. 8), both of which contain phenocrysts of quartz
of a size equal to, or greater than, the dune sand (0.2
to 2.0 mm in diainéter - Blissett (986). Similarly, and
second, granite with abundant coarse quartz crystals
crops out to the west of the Ranges (Blissett et al.
1988). But difficulties attach to these outcrops us
sources of the dune sand: they are of limited extent
(about 60 km? compared with the 300 km? of the
Scrubby Peak Dunefield}; and it can be questioned
whether they could produce a volume of quartz. sand
compatible with that represented by the total of the
dunes, Also the outcrops da not extend across the width
of the dunefieldl, so that the spread of sand from them

calls for varied strong winds, and lor distribution in
topographically difficult terrain. Moreover, the
dunefield extends westwards, i.e, windwards, of the
outcrops in question (Fig. 8). Against these arguments,
the Gawler Range Volcanics form a tegional basin
structure so that before erosion to their present
occurrences, the quartz-bearimg members could have
extended further to the W, In addition, the former shape
and size of these members could have been very.
different from their present representatives. But, on
the evidence, the rhyodacite, rhyolite and granite
outcrops of the Western ranges and adjacent areas do
not seem likely sources of suitable quartz sand.
The third possibility ts that the dune sand has been
derived from the Corrobinnie Depression, This runs
in a NW-SE direction west, and therefore windward.
of the dunefield and contains detritus derived from the
pranite dreas to the S. W and N as well as from the
Gawler Ranges. It contains quartz. comparable in size
and character to the dune sand of the Scrubby Peak
Dunefield (fine-grained, moderately well sorted,
typically subrounded, frosted und coated with iron
oxide - Gostin pers. comm, 1993). [tis concluded that
the dune sands of the Scrubby Peak Dunefield cannat
have been derived from the disintegration of the Gawler
Range Voleanies, but rather have been transported an
the wind from the Corrobinnie Depression. 3 distance
of at least 30 kot. Even if derived from the thyolite,
rhyodacite and granitic ouicrops, the sand must have
travelled 25 knv to crogs the zone uf dacitic bedrock.
Given the wind regime, the northern ann of the
Scrubby Peak Duneficld (X in Pig. 8) could only have

Fig. 8, Scrubby Peak Dunetield, western Gawler Ranges,
South Australia, illustrating bedrock type and possible
sources of the dune sant, X, Northem arm of the
Dunefteld, Y Nearest upwind wutcmp of rhyolne/rhyo-
dacrte, ED. HKuearto Dacite, YD. Yardea Dacite R-
Rhyohle, RD, Rhyodacite, See text for explanation. After
Blisseth.ef al. 1908,

52 OM, CAMPBELL, CR, TWIDALE, J ly BUTTON & JR. PRESCOTT

onginated in the rhyolitic/granitic outcrops indicated
by Y in Pig. 8, of from qutcrops further westward

Whether this would be considered far-travelled {sa
matter of definition, bul the sand is certamly not of
local denvation,

Mechanism of dune formation

Prevailing winds

It is suggested (see below) that the dunefields of the
Gawler Ranges have been shaped hy winds fron) the
western sector, This is consistent with |e pulusive
source of the sand of which the dunes are constructed
(see below). The relic linear dunes to the south of the
Gawler Ranges, which possibly formed jn are
Pleistocene times, extended fram NW to SE across lhe
northern. base: of Byre Peninsula, for they extend an
to the western shores of salinas such as Lake Awars,
but not on jo the eastern shores (Twidale & Campbell
1985). These NW-SE dunes also extend well below
low tide level between Cowell and Whiyalia (Van Deur
1983") but only in minor degree on the eastern side
of the Guifon northweslem Yorke Peninsula, where
the aeolian forns. were deposited duriny, a phase of
rising sea level and Where the dunes sre truncated by
wave action at the coast (Jessup 1967, 1968). This is
consistent with a wind regime domynaled by north-
westerlies.

In addition. at Lake Gaindner, the Junettes of the
eastern shore are much more substandal chan those of
the western and, as the lunettes are comparable to
coastal foredunes (Campbell 1968), this supports a
westerly Wind regime. Also, the huge Late Pleistovene
calcareous aeolianite toredunes of weat-facing shores
in South Australia (e.y. on Eyre Peninsula) dwarf their
east coast coumerparts. Thus, there is evidence ofa
predominantly westerly wind regime in the Gawler
Ranges and surrounding arcas during the pend, or
periods, of dune formation,

There is, however, an anomaly berween the presear
wind regime, as illustrated by the wind nse for
Nonning (Fig. 2b) and the presumed westerly wind
of dune formation, since the winter sandareoving winds
blow from the westerly sector, whereas the summer
sand-moving Winds are [rom the SW, S and SE, It is
presumed that most of the sand moyement would jake
place in Summer under hot and dry conditions. with
only minor transport in the matst. coal and vegetated
winter conditions. But, if there were only a slight
jaurudina! migration of clunatic zones during the period
of formation of the dines, a8 siggested for example
by Mabbutt (1977) and Sprigg, (1979), then the region

® Van Deur, W. 7, (1983) Submerged dunes of northeastern
Eyre Peninsula, MA thesis, University of Adelie
(Unpub,)

would have heen influenced by sumer rainfall
maxima Which would reduce sand movement during
thal season, On the other hand. dry winter conditions
would be saitable for the evidenced transport of sani
by westerly winds. The lack of compatibility between
Uuine orientation and wind direction remains a problem.
But assuming a Westerly wind regime, what factors
are important in the formation of linear dunes? Why
do parabolic dunes develop? How do the climbing and
falling dunes forin, and why do some of these forms
comlinue across the crests of the hills, whereas others
terminate on the upwind side only to resume on the
lee slope? How are the transverse dunes formed?

Linear dienes

The anigin of linear dunes as sGll debated (Cooke
e! af. 1993), Where the denes have been. closely
caamitned, as inthe Simpson Desert. these sand ridges
appear to display the same range of marphology, and
jnternal Structures and temporal variations in
asymmetry, indicstive of formation under a
bidirectional wind regime (McKee & Tibbetts 1964;
Wopfrer & Twidale |967; Brookfield 1970; Tsen 1990,
1993) The lincar dunes of the Gawler Ranges,
however, developed in an upland setting rather than
on desert plains. The vanfined valleys ought, in theary,
to funnel the wind and hence to be conducive to #
unidirectional Wind regime, buat bidirectional winds
could be either dominant or be superimposed on
unidirectional effects. No structures have been observer!
Within the dunes and, though this may reflect absence
of deep expositres as much as any diagnostic factor,
it *< not possible to state whether the dunes have been
shaped under a unidirectiunal or 2 bidirectional wind
regime.

Judging [oor the ofientation of the linear dunes in
the Gawler Ranges, the airflow was apparently
disturbed by the hills of ihe province and was funnelled
along valleys. The hills induce zones of increased and
of decreased sir flow and of enhanced turbulence. The
dunes chat are diverted around flanks of bills alse reflect
topographic control of the wind. The changes in dune
oniemation and morphology between, on the one hand,
the Great Victoria Desert. and, on the other, the
Gawler Ranges, are due to several tactors, First, the
westerly winds are diverted along the valleys. The
linear dunes are not everywhere parallel with the
regional air flow, a6 is characteristic of dunefields on
plains but their oriemtation is, 10 purt. determined by
the local wind regime. Second, sand supply decreases
within che upland where jhe silicic voleanic rocks
weather Jess readily than do the granites to the wes!
and, im particular, the supply of quartz grains is
reduced. Third, sand movement is impeded as a result
of the preserice of near surface moisture, held either
in valley alluviano or in rock fractures, and consequent
Vegetation growtls.

DUNES OF THE GAWLER RANGES x

Because of the lack of observed struvmres in the
dunes, the uncenainty aboul the relationship between
dune morphology and wind reginie ancl ihe tact that
the dunes aré relic and possibly related co different
wind velocities, wind directions jini! miinftll amounts
and distributions, the classification of the dunes as
linear, ic. elongate forms aligned in the direction ot
the dominant sand moving winds, is tenbtive.

Parabolic dunes

‘The occurrence of parnsolic rather chan linear dimes,
can be explained as follows, In the Curtobinnie
Depression to the § of the Gawler Kariwes (Bourne et
al. 1974) and elsewhere (McKee 166; Wasson ef al
1983) parabolic dunes are locaied in low lying areiss
characterised by an abundant supply of sand and by
proximity to proundwaters, which leads to the luwet
parts of the dune being stabilised by moisture and
vegetation. This allows the higher zones of sand to be
transported downwind to give blowouts or L-dunes,
In the Gawler Ranges area the peraholic dunes ocour
only in wide open yalleys and on plains. for example
in the western Moonaree Dunefield and in patches in
the Piceadilly Dunefield. However, they are no}
necessarily restricted to ihe towesl parts of these
valleys, On the available evidence and as indintied on
the 1.100 000 topographic map with a contow tsierval
of 20 m, the W-E belt of parabolic dupes in the
Moonaree Dunefield is sharply delimited on the
northern side by a belt of linear dunes and, less sharply,
on the southern side by dune-free plains, The panehole
dunes override low N-S tises in the valley and linear
dunes occupy some low-|¥ing areas in the mortherit part
of the dunefield. Thus, in addition bo stabilisation by
vegetation and an abundant supply of sarml, parabolic
dune. formation may require a eritical wind velocily
such as is attained only in wide valleys and on plains.

Climbing and falling dienes

‘The climbing and falling dunes are a parmicabar
variety of linear dune which mise arn descent
topeyraphic obstructions wher: the local wind is strong
enough to carry Uie available sand grins up and over
the topographic rises. The wind velociry is apparently
reduced on approaching the pdstacle and depositin
of sand occurs, Many of the bounding slopes of the
bornhardts are gentle (abut 512") and reverse eddy
flow is generally not developed, and hence celur duties
(Tsoar 1983; see also Fix. 6a) are not found winkbward
of cliffed obstacles. Where lie supply is sufficient,
sand accumulates until the dune reaches the height of
the obstruction, Where the bormmlardt ¢s low (in the
Gawler Ranges <40 mj the dune extends on co and
over the crest as a climbing arc! Gilling dune. Where
the bormhardt is high (> 40) m), the dune form tnay
be discontinous though sand is carried into the orest,
as evidenced by grains trapped in basins and crevices.

oa

Downwind of the obstucte, however, there ts a zone
of reduced wind velocity and sand deposition and dune
formation occurs. There may be further tunnelling ot
ihe sand to the lee of Ihe obstacle where the falling
dune is resumed in a topographic embayment (Fig. 6d).
The contrast between those linear forms that continue
unbroken over crests of bedrock hills and those m
which the climbing and falling components are
separated, evidently reflects the Bernouilli effect (Pye
& Tsoar 199).

Transwerse dunes

The wansverse dunes of the Scrubby Peak Dunefield
occur immediately downwind of the Corrobinnie
Depression, the presumed source of the sand, and
where the sand supply is abundant, The bornhardts
in this area stand about 100 m above the level of the
plain, form N-S trending ridges and the bounding
slopes are generally 5-10° with same as steep as 18°.
‘The plain is sand covered, with linear dunes of varied
orientation, hut generally NW-SE where there are nu
topographic obstacles, N-S in the valleys and W-E on
the bornhardts rises (Fig, 4). Some of the N-S linear
dunes override topographic obstacles and hence are
classed as climbing and falling dunes. The W-E
transport of sand across the bornhardt rises alse
explains the presence of sand in thé valleys. However,
some of the dunes are limiled to the upper slopes of
the barnhardts and ure located immediately to the lee
of the crest of the bornhardts (Fig. 7). Although they
inay be linear dunes formed by winds from a northerly
or southerly direction, in which case they do not
conform to the pattern of dunes throughout the region,
itis more likely that these crestal dunes are transverse
lo the originating wind. Ir is suggested than the sand
in these transverse dunes was driven up the windward
slope of the bornbardts and lithe or no deposition
occurred here due to acceleration of the air flow.
However, immediatly downwind of the crest,
separation of the air flow and deceleration occurred
so that deposition of sand sventuated. However, further
downwind, air flow accelerated and no dune formed.

lt is suggested that in this area the dunes are a resul)
of two different wind regimes; one a NW-SE wind that
was deflected by the tspography and one a NW-SE.
wind that was of sufficient strength to transport sand
aver the obstacles.

Significance of lanettes and salinas in sand supply

The Juneties located on the eastern shore of Lake
Gairdner evidently spawn fields of linear dunes in their
lee in a manner similar t that described from the
Simpson and other deserts (Twidale 1972, 1981). The
iransport of sand to the salinas by rivers and the
formation of the luneites are important influences on
sand supply and dune formation, Whether sand is
carried by the wind froa the W to the E shore of Lake

34 E, M, CAMPBELL, C. R. TWIDALE. § T. HUTTON & JR, PRESCOTT

Gairdner (some 30 km) fas not been determined. No
dunes have been observed an the bed of the lake (J.
Andrews, pers, comm. 1994), though small burchanoid
lorias have been reported on the bed of Lake Harris
(R. Major, pers. comm. 1992). Sand could be carried
by sultation across the salina Given the hygroscopic
character of the halite crust this may be difficull te
conceive, though Clarke (1994) deseribed saltation on
some salinas in Western Australia and 8. Wells (pers.
comm, 1994) has observed grains saltsting across a
salt surface in Califia. Alternatively, sand reaching
the W shore of the wind could be carried by wave
action to the E shore during the occasional periods
when there is water in the lake (Campbell 1968),
though not from the lake bed unless the salt crust is
dissolved or otherwise removed. Small! ephemeral salt
dunes, noted on the eastern shore of Lake Gairdner,
indicate the temporary redistribution of some of the
salt by deflation.

Conclusion

More data on the dunes of the Gawler Ranges
province are required before firm conclusions can be
drawn concerning the origin and age of the various
dune forms. The available information suggests that
the variations in marphology depend, at least in part,
on supply of sand, morsture content of the substrate,
vegetation coyer, wind speed and direcrion, and
topographic interference to the wind. The suggestion

that the lormation of parabolic as opposed ro linear
dunes 15 dependent on an abundant supply of sand and
fixing of the dune by vegetation only partially explains
the distribution of these dune types in the Gawler
Ranges province; other factors are apparently involved.
Climbing dunes are a vanant of linear dunes and torm
inthe zone of reduced wind velocity upwind of an
obstacle where the slope of the obstacle is gentle and
does not generate reverse eddy flow, Falling dones are
associated with climbing dunes provided the sand
supply 1s sufficient They develop in the zone of
reduced wind velocity to the lee of the obstacle, The
Cresial transverse dunes are also due Ww deposition in
the zone of reduced wind velocity. Though the dunes
of the Gawler Ranges area are essemially relic and ure
now stabilised by vegetation, there is sand movement
during very high winds. The duncs were active about
400) years BP The dunes of the Serubby Peak
Donefield in the southern Gawler Ranges demonstrate
that here the sand has been transported by the wind
al least 25 km from its source.

Acknowledgments

The authors thank J. A. Bourne for assistance in the
field, V. A. Gostin for advice on sedimentology, K.
Moxham for advice concerning air flow around
obstacles and R. Rice, R. Barrett and S Proferes for
technical assistance. Two referees gaye helptal
Suggestions On an earlier draft of the paper.

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A REVISED SYSTEMATIC PLACEMENT FOR
AUSTROTROMBELLA SOUTHCOTT
(ACARINA: HYDRYPHANTIDAE)

By MarK S. HARVEY*

Summary

Harvey, M. S. (1996) A revised systematic placement for Austrotrombella Southcott
(Acarina: Hydryphantidae). Trans. R. Soc. S. Aust. 120(1), 37-40, 31 May, 1996.
Austrotrombella leprosa Southcott, 1991, is transferred from the Trombellidae
(Trombidioidea) to the Hydryphantidae (Hydryphantoidea) and compared with other
thyasines of the Panisellus group.

Key Words: Taxonomy, Acarina, Hydryphantidae, Austrotrombella, Trombellidae,
South Australia.

Transactions of the Raval Sociery af S Aust. (1996), 1201). 37-40.

A REVISED SYSTEMATIC PLACEMENT FOR AUSTROTROMBELLA
SOUTHCOTT (ACARINA: HYDRYPHANTIDAE)

by Mark 8S, HARVEY*

Summary

Harvey.M. S. (1996) A revised systematic placement for Austrotrombella Southeou (Acarina: Aydryphantidac),

Trans, R. Soc, 8. Aust, 120K), 37-40, 31 May, 1996.

Austrotrambella leprase Southeott, 1991, is transferred from the Trombellidae (frombidiwidea) w the
Hydryphantidae (Hydryphantoidea) and compared with other thyasines of the Panisellus group.

Key Worps: Taxonomy, Acarina, Hydryphuntidae, Austretrombella. Trombellidae. South Australia,

Introduction

The monotypic genus Austrerrombella Southcott,
1991, was recently described from four unusual
specimens collected from wet shellgrit and soil beside
the cdye of a swamp neur Robe, South Australia. The
sole species, 4. /eprosa, was extensively described and
illustrated by Southcott (199]) and placed in the
trombidioid family Trombellidae. However,
exanunation of the type specimens lodged in the South
Australian Museum (SAM), reveals that the genus Is
misplaced and more closely resembles water mites of
the family Hydryphantidae than mites of the family
Trombellidac. A redescription of the genus is presented
here, along with an examination of its systematic
position within the Hydryphantidae,

Terminology mostly follows Cook (1974).

Family Hydryphantidae Piersig, [896
Genus Austrotrombella Southcott, 1991

Austrotrembella Suutheolt, 991; 207-208,
Type species: Austrotrombella leprosa Southeott, (91,
by monotypy.
Diagnosis

Differs from all other mites by the lollowing
combination of characters: pedipalpal tibia with distal
Sela; swimaming hairs absent: lateral eyes m capsitles.
idiosoma with numerous large plates: median eye
present and Situated near posterior margin of
prefrontalia, three pairs of acetabula in anterior group:

Remarks
Although regarded by Southcott (1991) asa member
of the trembidioid family Trombellidae, Arstrorram-
bella has more in common with the water mite family
Hydryphantidae, [n. particular, the chelate morphulogy
of the pedipalp, with a prominent dorso-distal libial
seta and a subdistally positioned tarsus, is virtually

* Western Australian Museum Francis Street Perth W. Aust.

diagnostic for the family (Cook 1974) and is completely
unlike trombellids and other trombidioids which haye
the tarsus inserted subbasally on the tibia (eg
Womersicy 1934). In addition, the idiosoma lacks the
dense vestiture of setae characteristic of most adult and
nymphal trombidioids which is, instead. represented
by longitudinal series of glandularia [termed ‘cupolae”
by Suuthcott (199))].

The presence of lateral eyes in capsules and the lack
of swimming hairs places the genus within the
Thyasinae (Cook 1974) and the large dorsalia and
ventralia Suggest a strong similarity with the Panisellus
group as detined by Bader (1985). This group contains
Fanisellus K. Viets (with P. thiertnemarni (K. Viets)
from northern Europe), Placothyas Lundblad (with P.
octopora (K, Viets) from South Africa), Octethyas
Lindblad (with Q. hewiritaé Lundblad from South
Africa), Porathyas Lundblad (with P theracata Piersig
and P primitive Lundblad from Europe and North
Africa) and Thyavella K. Viets (with T. mandibalaris
(Lundblad) from northern Europe). Therefore,
Austratrombella and its sole species, A. leprosa, is here
transferred to the hydryphantid subfamily Thyasinae.

Austrotrombella leprasa differs trom these other
penera in a number of small but significant ways: It
closely resembles Panisellus and Placethyas in the
location of the postocularia within the prefrontalia and
it differs from alt members of the group by the
possession of three pairs of acetabula in the anterior
group {1-2 pair$ in all others) and by the inclusion of
the acetabula on to the genital flaps (published illustra
tions of all other genera appear to indicate that they
are separate). it further differs from Panisellus by the
presence of a median eye (absent in Panisellus) and
from Placarhyas by the posterior position of the median
eye on the prefrontalia (situated near anterior margin
in Placothyas) and the presence of 6-8 pairs of
acetabula in the posterior group (2 pairs in Placorlyas).

This species is only the sécond thyasine reported
from Australia. The first, Netopanisus vinnulus Harvey
from Tasmania, differs by the lack of large dorsalia
and yentralia (Harvey 1988).

38 M, S. HARVEY

Austratrombella leprosa Southcotl, 1991
(FIGS 1-8)

Austrotrombella leprosa Southcott, 1991: 208-211, Figs
1, 2, 3a-e, 4a-c.
Material Examined
Holotype: 9, map reference (Penola I: 250 000)
283411, Robe district, S. Aust, [37°12'S 139°47’E],
in wet, alkaline, shellgrit-containing soil near swamp
edge, under a stand of Leptospermum lanigerum
(Aiton) Smith, 22.11.1990, R. V. Southcott (SAM
N1991112)_
Paratypes: 1 9, 1 o, 1 deutonymph, same data as
holotype (SAM N1991113-115).
Diagnosis

As for genus.

ag) |pro
|

Ne

—— << /
Iga /
4 ae / |

Description of adult

Integument slightly papillate. Lateral eyes on ocular
capsules; anterior-lateral eye (not visible in Fig. 1)
slightly larger than posterior-lateral eye; postocularia
slightly posterior to median eye, situated near posterior
margin of prefrontalia (Fig. 1). Idiosoma with
numerous porose platelets arranged as follows: large
prefrontalia; 1 pair of postfrontalia; 4 pairs of
dorsocentralia, posterior pair larger than others; 4 pairs
of dorsolateralia; 4 pairs of auxiliary platelets: 9 ventral
platelets, 1 between coxal plates, 2 behind genital
region, 2 pairs flanking anus, 1 pair situated
posteriorly. Six pairs of dorsoglandularia, 5 pairs of
lateroglandularia, 5 pairs of ventroglandularia (Figs
1, 2); sclerites associated with glandularia not forming
full circle (Figs 1, 2); vg2 situated near postero-lateral
margin of genital flaps and directed posterior-laterally;

Figs 1-2, Austrotrombella leprosa Southcott, holotype 9. 1. Idiosoma, dorsal. 2. Idiosoma, ventral (setae omitted from
one side). Abbreviations: dgl-6, dorsoglandularia; |g1-5, lateroglandularia; me, median eye; poo, postocularia; pro, preocularia;

vgl-5, ventroglandularia. Scale bar = 500 um.

REVISED PLACEMENT FOR AUSTROTROMBELLA 49

vg3 situated on level mid-way between genital flaps
and anus; vg4 situated on same level as anus; vg5
situated much closer to anus than to posterior margin
of body (Fig. 2), Genital region (Fig. 3): genital flaps
with setae on mesa] edge and scattered over posterior
third; 9-LL pairs of acetabula, 3 pairs situated in anterior
third, remainder (varying from 6-8 per side) situated.
on posterior third, all acetabula circular. Chelicera
(Fig. 7) of normal proportions, cheliceral claw curved,
with several teeth; cheliceral lamella about two-thirds
as long as claw, serrate. Capitulum without long, down-
turned anterior extension. Pedipalp (Fig. 6): tibia with
a thickened sub-medial seta on medial surface and with
stoul distal seta. Pedal coxae covered with long, thick
setae (Fig. 2). Legs (Figs 4, 5) without swimming setae
but most segments with numerous thick setae. Pedal
claws completely smooth (Figs 4, 5). Anus surrounded
by thick sclerotized ring (Fig. 2).

Ditnensions (um): holotype 9: body length 1464,

i!
A Nf
wy

op yey

L } } f ee
4 pane f- {

A — it

wa \ —— a ae _ \
2 —_-

width 1098; capitulum length 390; chelicera length
367, genital field length 333, width 314, pedipalp:
trochanter 63, femur 139, patella 112, tibia 195, tarsus
51; leg I: trochanter U1, femur 250, patella 182, tibia
236, metatarsus 269, tarsus length 255, width 64; leg
IV: trochanter 250, femur 276, patella 179, tibia 378,
nietatarsus 380, tarsus length 300, width 52.

Paratype 9: body 1488/1104; capitulum length 435,
chelicera length 385; genital field 381/346; pedipalp:
not measurable; leg I: trochanter 109, femur 287,
patella 173, tibia 262, metatarsus 302, tarsus 280/72;
leg IV: trochanter 303, femur 321, patella 210, tibia 443,
metatarsus 443, tarsus 350/58.

Paratype O°: body 1408/1024; capitulum length 358;
chelicera length 318; genital field 288/276; pedipalp-
not measurable; leg I: trochanter 106, femur 251, patella
140, tibia 218, metatarsus 255, tarsus 266/59; leg IV:
trochanter 230, femur 243, patella 163, tibia 336,
metatarsus 362, tarsus 288/45.

\

l : |
_ 7 | | \ { |
Sf ry I

\. TWiT Os, a {
oe y ‘< df “
YY Ho

Figs 3-8, Austrotrombella leprosa Southcott, 3-7, holotype 9. 3. Genital field. 4. Right leg I. 5. Right leg IV. 6. Right
pedipalp. 7. Left chelicera. 8 Provisional genital field, paratype deutonymph. Scale bars = 200 ym 3, 6, 7; 500 pm
4, 5; 100 um 8.

40 M. S. HARVEY

Description of deutonymph

Much as in adult except as follows: genital flaps with
2 pairs of acetabula situated at anterior and posterior
ends of flaps (Fig. 8).

Dimensions (um): body length 582, width 406;
genital field length 102, width 83.

Acknowledgments

I wish to thank David Hirst (South Australian
Museum) for the opportunity to examine the type
specimens of Austrotrombella leprosa.

References

Baber, C. (1985) Panisus-Studien: 6. Die Gattungen der
Panisellus-Gruppe (Acari, Actinedida, Hydrachnellae). Ent.
Basil. 10, 7-17.

Cook, D. R. (1974) Water mite genera and subgenera. Mem.
Am, Ent. Inst. 21, 1-860.

Harvey, M. S. (1988) Three new unusual water mites from
Australia (Chelicerata: Acarina: Hydryphantidae,

Hygrobatidae and Athienemanniidae). Mem. Mus. Vict. 49,
355-361.

SoutucortT, R. V. (1991) A new trombellid mite (Acarina:
Trombellidae) from South Australia. Trans. R. Soc. S. Aust,
115, 207-212.

WoMERSLEY, H. (1934) A revision of the trombiid and
erthyraeid mites of Australia with descriptions of new
genera and species. Rec. S. Aust. Mus. 5, 179-254.

TRANSACTIONS OF THE

ROYAL SOCIETY
OF SOUTH AUSTRALIA

INCORPORATED

VOL. 120, PART 2

SEVENTEEN NEW SPECIES OF CASTIARINA
(COLEOPTERA: BUPRESTIDAE)

By §. BARKER*

Summary

Barker, S. (1996) Seventeen new species of Castiarina (Coleoptera: Buprestidae).
Trans. R. Soc. S. Aust. 120(2), 41-59, 31 May, 1996.

Seventeen new species of Castiarina namely C. adusta sp. nov., C. antarctica sp. nov.,
C. aura sp. nov., C. azurea sp. nov., C. charientessa sp. nov., C. daranj sp. nov., C.
demarzi sp. nov., C. enigma sp. nov., C. ferruginea sp. nov., C. hemizostera sp. nov.,
C. jackhasenpuschi sp. nov., C. nonyma sp. nov., C. nullarborica sp. nov., C.
paulhasenpuschi sp. nov., C. phaeopus sp. nov., C. subcincta sp. nov., C. ustulata sp.
nov., are described and three established species namely C. cincta (Blackburn), C.
femorata (LaPorte & Gory), C. octospilota (LaPorte & Gory) are redescribed.

Key Words: Coleoptera, Buprestidae, new species, Castiarina.

Transactions of the Royal Saciety of 8, Aust. (1996), 120(2),. 41-59,
SEVENTEEN NEW SPECIES OF CASTIARINA (COLEOPTERA: BUPRESTIDAE),

by S. BARKER“

Summary
Barker, S, (1996) Seventeen new species of Castiarina (Coleoptera: Buprestidae), Trans, R, Soc, 8. Aust, 120(2),

41-59, 31 May, 1996.

Seventeen new species of Castiarina namely C. udusta sp, noy.. C. antarclica sp. nov., C aura sp. nov., C.
azureasp, noy,, C. charientessa sp. noy., C, darany sp. noy,, ©. demarzi sp. nov., C. enigena sp. now, C. ferruginea
sp. fov., © hemizostera sp. nov., C. jackhasenpuschi sp, nov,, C. nenyma sp, noy., C nullarborica sp. nov, ,
C paulhasenpusehi sp, nov., C phacupus sp. noy., C. sdheineta sp. nov. €. ustilar sp. nov., ure deseribed
and three established species namely C. citer (Blagkburm), C, /emoraia (LaPorte & Gory), C. avtoypilota (LaPorte

é& Gory) are redeseribed.

Key Worps; Coleoptera, Buprestidae. new species, Custiarina.

Introduction

The genus Cayriarina (LaPorie & Gory)
(Coleoptera: Buprestidae) 1s widely distributed in
Australia and also occurs in New Guinea where tis
distribution and abundance are virtually unknown. The
udults are often found on the flowers of native
Myrtaceae species and the larvae which are root and
stem feeders are largely unknown, A]though twenty
two new Australian species have beer identified
recently (Barker 1993, 1995) a further close
exumination of material collected over many years has
revealed even more new species. Ten of these are
associated with the Cudtiarina parallela (White)
complex and all occur only in WA; two are associated
wilh C. uctesprlota (LaPorte & Gory) and one euch
of these occurs only i. NSW and WA, The specific
status of a further species in this complex C. femorata
(LaPorte & Gory) is restored from synonymy: this
species also occurs in NSW Five new species
belonging to neither of these complexes ure described
from recently collected material

The complex to WA. has previously been mis-
iWentified as C parallela (White) but in fact this species
oecurs only in the eastern states. All members of the
group are elongate and have dark pronotini and elytral
volours, most offen rusty-bpown but dark blue in Lwo
species, a yellow margin-and a single raw of yellow
spoly along the middle of each elytran with a maximum
of four in each row and minimally one, when all of
the spots are fused. All species in the complex have
a dense layer of silver, flattened. feathery hair lining
the entire ventral surface and encroaching on to the
lateral surfaces nf the pronotum, This distinguishes
them from all other Castiarina which haye thin, round
hair on the yenlral surface. Most of the new. species

* Department al Zoology, University of Adelaide S AusI:
SOK)S:

also have sculptured proctivers in both sexes and this
feature is unique among Custiarina as all species
outside this complex have rounded, unsculptured
proctigers, The lwo species previously desenbed trom
the complex in WA are C. cracicolar (LaPorte & Gory)
and C. veropanctara (Barker 1995). Because this is the
most difficult group of Custiarina to identify, 1 have
included a key to the twelve known WA species.

©. oetaspilora (LaPorte & Gory) has.a dark head
with a yellow frontal spot, dark pronatum with yellow
Tateral margins and dark elytral markings with yellow
spots. The ventral surface is yellow with blue sutural
murkings and blue legs. A species occurring on the
eastern edge of the Darling Scarp, WA and on the
coastal plains has long been misidentified as C
vctospilota, [t resembles C cincta (Blackburn) which
occurs inland but is-casily distinguished by differences
in colour, being blue and yellow with blue legs whilst
C. cineta has some red markings on the elytra and red
legs with red sutural markings. The aedewgi are
different, There appear lo be twa species, ©, cincia
whieh is redeseribed und a new spevies which is
deseribed,

None of the remaining five species js close fn each
other and all ate distinctive. They have all been
collected recently, one of jhem by use of a Ture, a
technique not used before for the capture of Castiarina.

Materials and Methods

Male genitalia were prepared and jilustrated by the
method described by Barker (1987). The holotype is
illustrated in all new species except ane in which the
allotype is illustrated. Measurements given are mesn
total boxy Jength and width with standard error, except
where there dre insufficient specimens to make the last
calculation. Codens used jn the text for museum and
private collections followin the four letter system of

42 8. BARKER

Watt (1979) and Arnett et al, (1993) are: ANIC: Castiarina femorata (LaPorte & Gory) 1838
Australian National Insect Collection, Canberra; (FIGS 1B, 2B)

BMNH: British Museum (Natural History), London;

MNHN: Museum Nationale Histoire Naturelle, Paris; — Stigmodera femorata LaPorte & Gory 1838: 37. PI.
NMVA: National Museum of Victoria, Melbourne, 8, Fig. 42.

SAMA: South Australian Museum, Adelaide; WAMA: Stigmodera (Castiarina) octospilota var. roseipes
Western Australian Museum, Perth; HDWA Mr H, Deuquet, 1956 (new syn.).

Demarz, Guilderton; JHQA: Mr J Hasenpusch,

Innisfail; MHSA: Mr T. M. S. Hanlon, Sydney; Halarype: Sex unknown, S. femarata LaPorte & Gory,
MPWA: Mr M. Powell, Melville. Aust. MNHN (not seen)

Fig. |.. Photomicrographs of aedeagi and male and female proctigers of the following Castiarina species. A. Castiarina
daranj sp. noy. B. €. femorata (L & G), C. C. cincta (Blackburn), D. C. subcincta sp, noy, BE. C. actaspilota (L & G)-
EC, pawhasenpuschi sp. nov, G, C. bucolica (Kerremans). H. C. demarzi sp. nov. |. C. placida (Thomson), J. C. nullarborica
sp. nov. K. C. eyelistee (Rainbow). L. C. jackhasenpuschi sp. nov. M_ C aglaia Barker. N.C. hémizostera sp. nov. O.
© antarctica sp. noy. |. proctiger male. 2. proctiger female. P. C. octepunctata Barker. |. proctuiger male. 2. proctiger
female. Q. C._ferruginea sp. nov. 1. proctiger male. 2. proctiger fernale, R. C. adusia sp, nov, |. proctiger male. 2. proctiger
female. S.C. aura sp. noy, |, proctiger male 2, proctiger female. T. C. charientessa sp. noy. |. proctiger male, 2. proctiger
female, U. C. azurea sp. nov. |. proctiger male. 2. proctiger female, V. C. ustulara sp. nov. |, proctiger male 2. proctiger
female. W, C. phaeopus sp. nov. 1. proctiger male. 2. proctiger female. X. C. wonyma sp. nov. |. proctiger male. 2. proctiger
female. Y. C. enigma sp. nov. |. proctiger male. 2. proctiger female. Z. C crocievlor (L & G), |, proctiger male. 2.
proctiger female.

SEVENTEEN NEW SPECIES OF CASTLARINA 43

Colour

Head dark blue with green and purple reflections,
elongate yellow frontal spot, muzzle blue. Antennae
blue. Pronotum dark blue with yellow lateral margins
width increasing basally. Scutellum dark blue. Elytra
yellow with following dark blue markings; narrow
basal margin, elongate sinuous villa oo each side
meeting basal margin over humeral callus, meeting
post-medial fascia at margin enclosing spot on margin.
apical mark enclosing elongate yellow mark between
it and post-inedial fascia, srnall yellow apical spot on
each side of suture, marks all connected along sulure.
Ventral surface yellow with testaceous-ted sutures and
testaceous-red edges to abdominal segments, except
S_, Legs: coxae and trochanters testaccous-red and
dark blue: femora mainly testaceaus-red, apically dark
blue; tibiae a dark blue with ventral testacequs-red
mark near ventral apex; tarsi blue, Hairs silver.

Shape and sculpiure:

Head shallowly punctured, median sulcuy narrow,
muzale short, Antennomeres 1-3 obeonic, 4 halt:
toothed, 5-1] toothed, Pronoturm shallawly punctured,
apical margin straight, basal margin barely bisinuate,

Fig.2. Habitus illustrations of the following Castiarina species.
A. Custiarina actospilota (L.& GQ), B,C. femorata (L &
G). C. © subcincia sp. nov. holotype. D, C, darany sp.
nov. holotype. E. C cincta (Blackburn).

median basal fovea projecting almost to middle as
glabrous line; laterally parallel-sided at base, rounded
to apex. Scutellum scutiform, unpunctured, Elytra
punctate-striate, intervals convex, more so apically than
basally, lightly punctured; laterally angled outwards
from base, rounded at humeral callus, rounded post-
medially and narrowed to trispinose apex, marginal
spine small, interval to small median spine straight,
margin rounded and indented to small sutural spine;
apices slightly divergent. Ventral surface shallowly
punctured, edges. of abdominal segments glabrous,
elsewhere with sparse lorig hair, S,° male rounded,
female round, indented medially.

Size
Males, 124 + 0.35x4.9 4 0.14 mm (20). Females,
13.4 + 0.22 x 55 + 0.09 mm (5),

Aedeagus (Fig. \B)

Parameres angled outwards from basal piece.
rounded premedially then narrowed, rounded apivally,
Penis sharp, sides acutely angled away. Hypophysis of
basal piece medium width, apices rounded,

Distribution
NSW: Armidale district. central coastal.

Remarks

This species was synonymised with C. eetaspilora
by Saunders (1868) wha was followed by all subsequent
authors, Deuquet (1956) gave the varietal name roseipes
joa specimen he Wentified as 8. netespiluta. A single
male specimen in the South Australian Museum iden:
ified as Srig.8-spilora L. & G yar, roseipex. Deug. in
Deuquet’s handwriting, 1s clearly # separale specics
from €. octaspilora. The holotypes of C. acrospilate
and C. femorata are both sodged in the MNHN but
cannot be identified because their labels, along with
those of all other LaPorte & Gory type labels, have
heen removed, Deuquer’s description of the red femorit
and ted vermiculation on the ventral surface of his 5,
vetospilma vat, reseipes concurs with the origimal
description of §. femorala. The figure of S. femorata
(LaPorte & Gory 1838, P|.8, Fig. 42) also conforms
in general, except that the illustration shows the base
of the head to be yellow instead of the yellow frontal
spot in the Deuquet specimen. J assume that this is
artistic licence on the part of the illustrator because
none of the species in this species group has a head
with a yellow base. In the figure the pre-medial fascia
is complete. A similar pattern is presemt in only two
of the fourteen specimens examined. All other
specimens have the pre-medial fascia incomplete, thus
twa spots on each side of the suture coalesce to form
a sinuous yellow vitta. S. ectospilota vat. roseipes
Deuquet is undoubtedly a synonym of C. femorata (L
& G).

ad S. BARKER

Castiarina actospilota (LaPorte & Gory)
(FIGS 1B, 2A)

Stigmodera ocrespilora LaPorte & Gory 1838: 28, Fig,
29, PL fs

Holotype: Sex wniknown, Australie, MNHN (not seen)

Colour

Head; muzzle blue: base dull green-purple; yellow
frontal mark, Antennae blue. Pronotum: medially dull
#reen-purple; laterally yellow, base wider than apex,
Seutellum black with dark blue reflections. Elywa
yellow with following dark blue markings: narrow
basal margin, small and narrow pre-medial faseia
absent in many specimens, connected to lang oblique
vita from lower end reaching lateral margin, enclosing
very large basal yellow. spot and smaller ong.ontnargin;
broad post-medial fascia reaching lateral margin,
shighily angled posteriorly, enclosing large yellow mark
between i and first fascia; mark covering whole apex.
cnelosing small yellow mark between it and second
fuscia. Ventral surface yellow with blue sutures. Legs
blue. Fairs silver

Shape and seulmure

Head closely punctured, median sulcus small,
muzzle short, Antennomeres 123 obcunic, 4 half-
toothed, 5-IL wathed. Pronotum closely popetured,
minute basal fovea, extending forwards to middle as
glabrous line, basul notches represented by glabrous
areas, more marginal than medial; apicul margin
straight, basal margin bisinuate; laterally rounded out
from base, widest pre-medially, rounded and narrowed
to apex. Scutellum scutiform, glabrous, excavate at
busal edge. Elytra punctate-striate, intervals convex and
punethred, less sa tiedially than elsewhere, laterally
angled out from base, rounded at humeral callus,
concave, rounded post-medially and parrowed to
trispinose apex; small marginal and medial spines,
margin between straight, widely separated, small
sutural spine, close to medial spine, apives divergent,
Ventral surface with shallow punctures. edges of
abdominal segments glabrous, elsewhere moderately
hairy, hairs medium length. S.- truncate and indented
medially jn both sexes,

Size
Males, 12.5 + 0.28% 51 + 0.20: mm (33). Females,
13.6 + 0.23 x 56 + O.U mm (40).

Aedeagus (Fig. 1E)

Parameres angled outwards from basal piece,
rounded pre-medially, parallel-sided, rounded at apex
Penis sharp, sides obtusely angled away. Apophysis of
basal piece medium width, apically rounded.

Distribution

NSW: Blue Mis, Sydney, central to north coastal,
Qld: Southern and central coastal. Blaekdowi
Tableland, Shrove Is,

Castiarina eincia (Blackburn) 1890
(FIGS IC, 2)

Stigmedera cincta Blackburn t890: 13, 187
(replacement name for 3, eubrocineta Kerremans
1890: 46, primary homonyin §. rubrecincia Gehin
1855),

Holotype: 9, S milrecinera Kerromans, Bouvard
Australie. BMNH.

Colour

Head: muzzle blue-green; base. purple-green; large
yellow frontal mark, Antennae blue-green, Pronotum
laterally yellow, medially purple-green, Scutellum blue.
Elytra yellow with the following dark blue markings:
basal margin, pre-medial fascia not reaching margin
with ends projecting anteriorly to basal margin as vitta
enclosing large yellow basal spot, clongate yellow mark
on margin at humeral callus, narrow red apical margin:
post-medial fascia not reaching margin, enclosing
yellow band between it and first fasvia; pre-apical mark
in shape of short fascia enclosing elongate yellow band
between it and second fascia, all marks Connected alony
suture. Lateral red margin on ihe two intervals from
humeral callus, broader at pre-apex and apex, Ventral
surface yellow; sternum lateral blue-green sutural
marks, ted medially and along edges of abdominal
segments. Legs: femorg blue-green apically, red
medially, tibiae and tarsomeres blue-green, Hairs
silver,

Shape and sculpture

Head with shallow punctures, flat, muzzle short.
Antennomeres 1-3 obcanic, 4 half-toothed, 5-11
wothed, Pronotum with shallow punctures, basal fovea
represented by shallow depression, very small basal
notches more marginal than medial; apical margin
straight, basal margin almost straight, Scutellum
scutiform, without punctures, excavate along basal
edge, Elytra punctate-striate, intervals flal medially,
convex apically and laterally, punctured, less so
medially; laterally angled out slightly from base.
rounded a( humeral callus. concave, rounded post-
medially and narrowed to Laspmnose apex; marginal
Spine small and sharp, median spine larger and blunr,
sutural spine small and sharp, margin rounded between
spines. Ventral surface with shallow punctures, sparse
short hair, $); truncate both sexes.

SEVENTEEN NEW SPECIES OF CASTIARINA 45

Size
Males, 1.3 + 0.22.x.4.3 + 0.10 mm (24). Females,
12,2 40.34 5 47 + O15 mm (20).

Acideagus (Fig. (2)

Parameres angled outwards from basal piece.
rounded. pre-medially then angled outwards, rounded
apically. Penis sharp, sides. acutely angled away.
Hypophysis of basal piece narrow, apically rounded.

Disirthuttor
Occurs in intund squth-western WA.

Castiarina daranj sp. nev.
(FIGS 1A, 2D)

Holotype: or, 4km NE Rocky Glen, NSW, 3.411981,
S. Barker. SAMA J 21 300.

Allorype: Q.. same data as holotype. SAMA 12} 301.

Paratypes: NSW: 19, 43-km 8 Narrabri, 27.x,1975,
S. Barker, SAMA; 14 oo, 829, Binnaway,
2.x) 198], S. Barker, SAMA, 19. 6 km SW Rocky
Glen, 3,xi,1981, S. Barker, SAMA; 20° oO", 29 G4
kin SW Rocky Glen, 3.47.1981, 8, Barker, SAMA, 19,
3 km SW Rocky Glen, 3.¥7.1981, S, Barker, SAMA;
lor.) 9,2 km SW Rocky Glen, 3.¥i,1981, S, Barker,
SAMA; Soo. 49 @ , same data as holotype, SAMA;
Zo, 29 9. Garrawilla 10, 6 km NE Rocky Glen.
3,x1,1981, 5, Barker, SAMA; 29 @, Garrawilla T.0.,
8.«j.198], R. Anderson, SAMA; | 9, Garrawilla T.O.,
17.x1.1981. S. Barker, SAMA: Lot. 4@ ko NE
Coonabarrabran, 6.x1.1983, A. M. Sundholm, MHSA:
300, 19, 40 km & Coonabarrabran, 9.xi.1990, T.
M.S Hanlon, MHSA; 10°, 60 kin N Coonabarmbran,
9.xi.1990, T. M. S. Hanlon, MASA, 200,299,
40 km E Coonabarrabran, 8.xi-199], T. M.S, Hanlon,
MHSA; lo", Round Hill, 21. xi.1991, T. M. 8, Hanlon,
MHSA; 10°, 30 km E Parkes, 29.xi.1993, TM. &.
Hanlon, MHSA,

Colour

Head black with green reflections, muzzle blue, large
orange-yellow ltontal spot, Antennae blue. Pronotum
medially dark blue, laterally orange-yellow increasing
in width basally, Scutellum dark blue. Elytra orange-
yellow with the following black markings wath blue
reflections: narrow basal margin. sinuous yitta from
basal margin over humeral callus mecting margin-and
sutural mark enclosing yellow-orange elongate mark
an margin and basal spot; broad post-medial fascia
reaching margin enclosing large orange-yellow mark
between it and firstmark; mark covering apex meting
post-medial fascia on margin and enclosing a large
vrange-yellow mark; small orange-yellow apical spot

on each side, Ventral surface orange-yellow with blue
marks along sutures: and along edges of abdominal
segments. Legs: fernora and tibiae blue, tarsi bright
blue. Hairs silver.

Shape and sculpture

Head shallowly punctured, median sulcus shallow.
muzzle short Antennomeres 1-3 obconic, 4 hall-
toothed, 5-11 toothed. Pronotam shallowly punctured.
uarrow basal foyea extending forwards to middle as
glabrous line; apical margin projecting medially, basal
inargin barely bisinuate; laterally parallel-sided al base,
rounded to apex. Seutellum seuuform, glabrous.
excavale, Elytra punetate-striate, intervals convex,
lightly wrinkled and punctured: laterally angled out
from base, rounded at humeral callus, concave,
rounded post-medially and narrowed to tispinose
apex; spines small, margin straight between marginal
and median spine, rounded between median and sutural
spines, apices divergent, Ventral surface with shallow
punctures, edges of abdominal segments glabrous,
elsewhere moderately hairy. sparse medium length
hair S,! male truncate, slightly indented medially;
female trincate, deeply indented medially, margin
overhanging apex which is covered with bristles.

Size
Males, 10.4 + 0.15 % 4.0 + 0,06 mimi (34), Females,
hl + @20 x 44 4 0,09 mm (26),

Aedeagus (Pig. 1A)

Short. Parameres angled outwards fram basal piece,
rounded pre-medially, parallel-sided, apically rounded,
Penis sharp, sides acutely angled away. Hypophysis of
basal piece medium width, apically rounded.

Remarks

The basal colour of this species fades rapidly. in death
from orange-yellow to pale yellow, Both C octospilou
and C. femorata have a yellow basal colour in life, Alsu
it is smaller than the other rwo species and the male
genilalia are smaller and a different shape (Figs 1A.
1B, LE),

Etymology

Name derived from Arabic daranj, orange.

Castiarina subcincta sp. nov.
(Figs 1D, 2C)

Holorype: ct, Bold Park, City Beach, WA, 3.0 -1976,
RP. McMillan, SAMA [ 21 302-

Allotype; Q, City Beach, WA, 3.xii-1955, J. A. L.
Watson, SAMA I 21 303.

46 5. BARKER

Pararypes! WA: 19, Cannington, 12,x1i.1954, S,
Barker, SAMA; 30°", City Beach, 24.x,1954, S.
Barker, SAMA; I2oo, 399, City Beach,
26.x1.-25,xi1.1955, J. A. L, Watson, SAMA; 19, 9.5
km SW Jarrahdale. 11 xi 1956, S. Barker, SAMA; Lor.
City Beach, 6.97.1957, 5S. Barker, SAMA; Lo, City
Beach, 2.1964, S. Barker, SAMA; 30° co, Wembly,
3.%.J970, S, Barker, SAMA; 4a°@, 19, same data
as holotype, SAMA, 2o°o, 19, Glen Eagles,
7,1,1983, S, Barker & K. T. Richards. SAMA; 20° 0”,
3.9 9, Walyunga N.P., 4.xi.1984, T. M.S. Hanlon,
MASA; Sara, 19, Wembly, 4.xi.1985, T. M. S.
Hanlon, MHSA; Jo, Swanbourne, 23.4,1991, T. M,
S. Hanlon, MHSA; 6ccr, 19, Swan R., H.W.
Brown, SAMA.

Colour

Head basally dark blue with green and purpic
reflecuions, muzzle blue, large yellow frontal sper,
Antennae dark blue, Pronotum medially dark blue with
green and purple reflections, laterally yellow, Scutellum
dark blue, Elytra yellow with following black markings
with blue reflections: narrow basal margin; sinugus
vitta from basal margin over humeral callus mecting
narrow pre-medial fascia close to margin enclosing a
yellow spot on margin and latge yellow basal spot;
broad post-medial fascia reaching margin enclosing
yellow spot between it and pre-medial faseia. mark
covering apex enclosing elongate yellow mark between
it und post-medial faseia and variable apical yellow
spot, all marks connected along suture and along
margin except at humeral callus; outer margin of apical
spot variably red, Ventral surface yellow, sutures blue
and lateral blue spots on S,, S,, S.- Legs blue. Hairs
silver,

Shape and sculpture

Head shallowly punctured, median sulcus small and
shallow, muzzle short. Anfennomereé 1-3 obeonic, 4
half-toothed, 5-l1 toothed. Pronotum shallowly
punctured, narrow basal fovea extending forwards to
middle as glabrous line, basal notches represented by
glabrous area on cach side closer lo marvin than
middle, apical margin projecting medially, basal
margin almost straight, laterally parallel-sided at base,
rounded from base to apex. Scutellum scutiform,
glabrous, Nat. Elytra punctate-striate, intervals convex,
wrinkled; laterally angled out from base, rounded at
humeral callus, concaye, rounded post-medially,
narrowed {to trispinose apex; small marginal spine,
larger medial spine, smaller sutural spine, margin
rounded between spines, apices diverging. Ventral
surface with shallow punctures, edges af abdominal
segments glabrous, elsewhere sparse medium Jength
hairs. S$: truncate both sexes.

Size
Males, 12.8 + 0.13% 4.9 + 0,06 mm (41), Females,
3.5 + 0,22 x 5.2 + 009 mm (24).

Aedeagus (Fig. |D)

Parameres angled outwards from basal piece,
rounded pre-medially. parallel-sided, rounded apically.
Penis sharp, sides acutely angled away. Hypophysis of
basal piece narrow, apically rounded_

Remarks

This species, previously confused with ©
octospllota, forms a species pair with CL cincta
(Blackburn). [t occurs on the coastal plam of WA and
on the western edge of the Darling Searp whereas C°
cincta occurs in the more arid inland areas.of the south-
west. It differs from that species having only very small
red markings on the elytra and not op the legs or
abdominal segments. C cincta has red markings on
the elytra, red femora and red sutures on the ventral
surface. Also the elytral spines are mure obvious in
C. subeiner than in ©, cinera and the male genitalia
are a different shape (Figs IC, 1D).

Etymolesry
The name is derived from CF osaé, under, 1 cirectarn,
girdle.

Castiarina adusta sp. noy,
(Figs JR, | RI, | R2, 3)

Holotype: Oo, 5 km W Mt Dale, WA, 13.%.1980_ S.
Barker, SAMA [ 21 304.

Allorype’ 9, Lake Grace, WA. 19.*.1970, K, & E,
Carnaby, ANIC

Pararypes: WA: Sora, 49 9, same data as allotype,
ANIC, 19, 80 kin B Hyden, 29.%.1984, M_ Powell,
MPWA, Lov! km WNW Bonnie Rock, 20.1% 1990,
S$. Barker, SAMA

Colour

Head, antennae and pronotum bronze, Seutelluiy
dark blue. Elytra yellow with the following brown
markings! marks coalesced leaving a continuous yellow
margin from base lo near apex, a row of four elongate
spots down each elyiron, the first two variably
connected. Ventral surface and legs bronze. Hairs
silver

Shape and sculpture

Head shallowly punctured, median sulcus shallow,
sides variably glabrous basally, muzzle short. Antenno-

SEVENTEEN NEW SPECIES OF CASTIARINA 47

Fig, 3. Habitus illustrations of the following Castiarina
species. A. Castiarina nonyma sp. nov, holotype. B. C.
cracicolor (L & G), C. C. enigma sp. noy. holotype_ D,
C. antarctica sp. nov, holotype. E. C, phaeepus sp. nov,
holotype. F.C. ustulata sp. nov. holotype. G. C. azurea
sp. nov. holotype. H. C. ferruginea sp. nov. holotype. .
C. aura sp. nov. holotype. J. C. adusta sp, nov, holotype.
K. C. charientessa sp. nov. holotype.

meres 1-3 obconic, 4-11 toothed. Pronotum shallowly
punctured medially, larger and deeper punctures
Jaterally, narrow basal fovea extending anteriorly to
middle as glabrous line, basal notches represented by
glabrous area on each side closer to margin than
middle; apical margin projecting medially, basal
margin barely bisinuate; laterally parallel-sided at base,
rounded and narrowed to apex, laterally hairy.
Scutellum scutiform, punctured, flat. Elytra punctate-
striate, mtervals convex, wrinkled and punctured,
Jaterally angled out from base, rounded at humeral
callus, concave, rounded post-medially and narrowed
1 bispinose apex; spines small and blunt, margin
yariably rounded and indented or straight between
spines, apices hardly diverging. Ventral surface with
shallow punctures, edges of abdominal segments
glabrous, elsewhere hairy, hairs flattened and feathery.
Legs: femora hairy with flattened hair, S,: males
truncate; females rounded,

Nee
Males, 14.5 + 012 x 50 + 0.06 mm (7), Females,
59 + 045 . 54 + 0.12 mm (5),

Aedeagus (Fig. IR)

Parameres angled outwards from basal piece,
rounded pre-apically then parallel-sided, rounded
apically. Penis sharp. sides acutely angled away,
Hypophysis of basal piece medium width, rounded
apically, Proctiger, medial apical edge shallowly
concave, rounded Juterally (Fig. 1 Rl).

female terminalia (Fig. 1 R2)

Proctiger with apical edge flattened, rounded
Jaterally.

Remarks

C. adusta sp. nov. is the largest member of this group
in WA, It can be separated from C._ferruginea sp. nov.
the next largest brownish species, by its size, the
conformation of the elytral markings - there are our
spots in C. ferruginea and three in C. adusta, its
relatively unsculptured proctiger in both male and
female. whereas both sexes of C\ ferruginea haye
bilobed proctigers and in females they are spined. The
aedeagus in C. ferruginea is broader at the apex than
that of C. adusra (Figs JQ, IR).

Erymalogy
The name is derived from L adustus, brown.

Castiarina azurea sp.. Nov,
(FIGS 1U, 1 Ul, 1 U2, 3G)

Holotype: & , 2 km E Tallering Station, Pindar, WA,
22.ix.1989, S. Barker, SAMA | 21 305.

48 5. BARKER

Allotype: 9 , same data as holotype, SAMA [21 306.

Paratypes’ WA: lor, Goomalling, .ix1953, RB
McMillan, WAMA; Lo, 22 9, Moorine Roek,
16,x, 1953, FH. Uther Baker, SAMA; 19, Wialka,
Yun 1957, §. Barker, SAMA; 10°, Toohbin. 18.» 1958,
F_H. Uther Baker, SAMA; 20-9", Burracoppin-
16,*,1963, F H, Uther Baker, WAMA;: Jota", 29 O.
78 km NE Wubi, 17.1%.1970, 8. Barker, SAMA: 1,
9% km NE Wubin, |7ix 1970, §, Barker, SAMA;
tom, 19,55 ki S Payne's Find, 18.14, 1970, SAMA;
[ct 19,57 km S$ Payne's Find. 187.1970, S. Barker,
SAMA; 2¢ 9. 10 km B Blachburting Rock, Wialki,
2Uax.1970, 3. Barker, SAMA: Scere, 29, Wiulki,
2Lix,197), S. Barker, SAMA; 2am, 299,
Walgoolan, Six 1871, FH. Uther Baker, SAMA, 19,
Tallering Station, Pindar. 3,i2,]976, R. PR McMillan.
SAMA, Sara, 39 9, 18.9 kor WSW Coolgardie,
Ik rx 1976, RJ. Chinnock, SAMA! Lo, 50 km N
Kalbarri, 20viii-1978, M. Powell, WAMA; 1c, Balline
Station, 24/25.yi979, A, M. & M, J. Douglas.
WAMA; 1a. 22 2. Muckinbadin, (0.1979, R. P
MeMillan, WAMA; cr, J6 km E Mt Magnet,
20.ix. 1980, S. Barker & D. J. Williams, SAMA: lor.
1% kot N Carnarvon, 22.ix 1980, S. Barker & DL J,
Williams, SAMA; 10, 22 G, 89 km N Carnarvon.
27. ix 980, & Barker & DJ. Williams, SAMA: 1 cr.
29, 44 4m 6 Kalbarri. 2.1%. 1980, S. Barker & BD.
!. Wiliams, SAMA; 3a 07 46 kin E Kalbarry,
261% 1980, 8 Barker & D, J, Williams, SAMA) 20°C,
12, 8 Rem paddock, Taliering Station, Pindar,
27x 1980, S, Barker & D. F Williams, SAMA; 20 &,
LQ. I7 km W Mullewa, 29.7%.1980, S. Barker & D,
4. Williains, SAMA, 29 2, Mt Walker, 25.«.1980, R.
FP MeMillan, WAMA;: Io, Gubhin, 29.6198], R, P
MeMillan, WAMA; 29 9. Southern Orass, «1981,
R, FP. MoMillany WAMA: Ic, 394, 2 km N
Evanston, 237%,.1982. B. Hanich & T. P. Houston,
WAMA, 19, 64 km NE Esperance, 18,1982, S,
Barker, SAMA: Lor, Bullfinch, 2.x 1983, EB Jones,
MPWA; 19. 35 lon W Salmon Gums, 8.x%.1983, G.
Rrawning & G. Mutze, SAMA, 3° 3, 1 9, Southern
Cross, §,4,1983, R, P. MeMillan, WAMA, 6orc,
Q¢ @, Esperance to Norseman Hwy, 35 km W TG
to Peake Charles, 95 1983, G. Browning & G. Mute,
SAMA. 29 9. 35 km E Merredin. 24.1983, G.
Browning, SAMA; lor, 229, 50 kin E Merredin,
24.%.1983. SAMA, Lor, Uberin Rock, I6.ix. 1984, R-
P McMillan, WAMA,; Ler, 136 km NE Payne's Pind.
30.in O84, M_ Powell, MPWA; lo. 3 km W
Dowerin, 22,x, 1984, R, P McMillan, WAMA,; Ter,
Eneabbu, 4,x,1985, R. PB. McMillan, WAMA; lo, N
Tarin Rock teserve, 15/l6.x.!985. T. F Houston.
WAMA: 1o. 75 km E Hyden, 24/27.4.1985, T.
Houston. WAMA; 60° co’, W of Coorow, 2.x. 1986, A.
G. Wells, WAMA; 1 cr, 16 km NE Merredin, 9.x 1986,
R. P. MeMillun, WAMA; 2er co, Encabba, x./986, R.

P. McMillan, WAMA; |o. Bindoo Hill reserve.
12.1%.1987. TF. Houston, WAMA; 20°, 1.9, 7 km
SSW Jingemarra Station, 24/26iii,1988, R. P.
MeMillan & T, RK Houston, WAMA; 1 2, Shark Bay,
291.1988, A, Hay, MASA: Io, HO km N
Carnarvon, (8 ix 1989. 8. Barker, SAMA: 20° 0", 19),
Pindar paddock, Tallering Station, Pindar, 21.1x.(989,
S Barker, SAMA; 7o'o", same data as holotype,
SAMA; Lo, 19, 3 km N Tallering Station, Pindar,
22, 1x. 1989, 8. Barker, SAMA; 2c ot, & km N Tallering
Stalion. Pindar, 22.ix 1989, S. Barker. SAMA: Soo,
19 kro N Tallering Station, Pindar, 22,ix1989, S,
Barker, SAMA; |, pravel bay, Bonnie Rock,
20.ix 1990, S. Barker, SAMA, 26°, Merredin,
21,x.199], TM, §. Hanlon. MHSA; BC &, Ghooli,
21,1991. T M, 8, Hanlon, MHSA;, 2a o_ 3 km S$
Yellodine_21.%.1991, 'T. M_S. Hanlon, MHSA: 3.7 o,
N7T, 32 km E Southern Cross, 21-199). T) M.S
Hanlon, MHSA, Lo, Quees Victoria Springs,
21/22.41,1992, D. Knowles, MHSA; loro. 19.
Northam, SAMA; 20° co. Ankertell H. W. Brown,
SAMA.

Colour

Head dark blue with purple reflections Antennae
bronze. Preonotum dark blue with purple reflections.
Seutelum purple. Elytra with yellow background
colour and dark blue elytral markings, coalesced
farming four yellow spats an cachelytron. basal more
or less rounded, pre-medial elongate, pust- medial
founded, pre-apical elongate, basal and pre-medial
opalesced in about half specimens examined forming
an elongate mark, post-medial and pre-apical coalesced
in only ohe specimen examined, yellow margin from
base (6 near apex. Ventral surfice bronze with coppery-
purple reflections, Hairs silver

Shape and sculpiare

Head closely punctured, medtan. sulcus present,
oluzele short. Antennomeres 1-3 obcorie, 4-1 toothed.
Pronolum closely punctured, narrow basal foven
extending forwards to middle as glabrous line. basal
notches represented by glabrous area on each Side
closer to margin than middle; apical margin projecting
medially, basal margin almoxt straight; laterally angled
inwards from base for short distance then angled
outwards and rounded to widest part post-medially,
rounded and narrowed Io apex Seutellum scutitonn,
punctured. flat Elytra punciate-stnate, intervals
convex, wrinkled and punctured, more heavily laterally
than medially; jaterally angled out from base, roundes!
al humeral callus, concaye, rounded. post-medially,
narrowed to bispinese apex; marginal spine small and
sharp. sutural spine minure, margin indented and
rounded between spines, apices hardly diverging.
Ventral surface with shallow punctures, edges ul
abdominal segments glabrous, elsewhere hairy, hairs
medium length, flaltened and feathery. 85: truncate in
rnales, rounded in females,

SEVENTEEN NEW SPECIES OF CASTIARINA 49

Size
Males, 10,1 + 0.15 x 3.2 + 0.05 mm (110). Females.
10.7 + 0.24 x 3.4 + 0.08 mm (36),

Aedeagus (Fig. 1)

Parameres angled outwards from basal piece,
rounded at upex- Penis sharp, sides obtusely angled
away. Apophysis of hasal piece medium width, apex
rounded. Proctiger faintly bilobed, lobes rounded (Fig.
bub.

Female terminalia (Fig, 1 U2)
Proctiger faintly bilabed, lobes firintly rounded,

Remarks

C azurea can be distingdished from al] other
members of the C. parallela species group except C.
octapunctara by the dark blue colour of the elytra. C.
gelupunctaia has round yellow elytral markings
whereas they are elongate in C. azurea. In ©, azured
the aedeagus is short and broad and in C. ectopunctaia
iLts elongate (Figs IP, 1U). The proetigers in both sexes
of C. ectopunctata have pointed lobes whereas in both
sexes af C. azurea the proctiger lobes are small and
rounded (Figs | Pl, 1 P2, 1 Ul, | U2). There appears
to be a cline insize within © azurea, Specimens trom
north of Carnarvon and fram ihe NE wheatbelt areas
of WA are larger than thase fram further east and south.
A minority of specimens has the first two yellow marks
on the elytra fused (6 form an elongate basal mark.

Enology
The species name is derived from F azur. blue.

Castiarina charientessa sp. nov.
(FIGS IT, | Tl, | T2, 3K)

Holotype: o , 10 km $ Dongara, WA, 44x.1995, 5.
Barker. SAMA J 21 307.

Allorype: 9. 20 kin S Lancelin, WA, 4.51990, 3
Barker, SAMA [ 21 308.

Pararypes: WAc 2a, Cervantes, 23.1*%.1977. M.
Powell. MPWA; 2o°.c°, 69 9, 45 km N Eneabha,
20,ix.1980, S, Barker & D: J Williams, SAMA; 2° 9,
200 m N Ledge Pi T.0,, Lancelin Rd, 8.x.1980, 5,
Barker, SAMA, 10%, 299, McDermid Rock,
11.1981, G.I. Keighery, WAMA; 20° ot, | 9, Green
Head, 27.viit. 1981, Ro P- MeMillan, WAMA: lor, 19,
2 km N Badgingarra, M. Powell, 15.ix.1984, MPWA,;
Lor, Greenough, 26/29.viii 1989, R. P. McMillan,
SAMA; 20°", sume data as holotype SAMA; Ic,
10 kin S Dongara, 4.ix 1995, S. Barker, SAMA,

Colour

Head coppery, Antennae bronze. Pronotuin coppery.
Scutellum bronze with blue-green reflections, Elytra
yellow with coppery markings coalesced forining a
yellow margin two interstices wide; a yellow vita on
each side from base to pre-apical area, Ventral surface
cuppery. Legs: femora and tibiae cappery; tarsomeres
bronze. Hairs silver.

Shape and seulprure

Head closely punctured, median sulcus present,
muzzle-short. Antennomeres 1-3 obconic, 4-11 toothed,
Pronotum closely punctured, stnall basal fovea
extending anteriorly to middle as glabrous line, apical
margin projecting medially, basal margin almost
straight; laterally parallel-sided at base, angled
oatwards, rounded to widest at middle. rounded and
narrowed to apex, Scutellum seutiform, glabrous, flat.
flyrra punctate-striate, mtervals convex, wrinkled and
punctured more heavily laterally than medially;
laterally angled out (rom base, rounded at humeral
callus, concave, rounded post-medially and narrowed
to bispinose apex; bath spines minute, margin rounded
and indented between spines, apives diverging. Ventral
surface with shallow punctures, edges of abdominal
segmemts glabrous, elsewhere dense, flat, feathery
hairs. S,; males truncate: temales rounded.

Size

Males 12.9 +: 0:24.x,4,3 + 0.08 jam (12), Females,
135 + 017 x 4.3 + 0.13 mm (14).
Aedeagus (Fig, | TU

Parainetes angled outwards from basal piece,
rounded apically. Penis sharp, sides acutely angled
away, Apophysis of basal piece mediurn Width, rounded
apically. Proctiger bilobed, lobes near mid-line, blunt.
apical edge straight, rounded laterally (Fig. | Tl).

Female terminalia (Fig. 1 T2)
Proctiger as in male. lobes more pronounced,

Remarks

This species can be distiaguashed from any others
in the group by the elytral markings. ft is the only
species occurring in WA which has all of the yellow
elytral spots fused to form an elongate yellow yitta on
each elytron. Some specimens have darker red
murkings on the elytra than others and these tend tr
fade to dark brown in old specimens. Six specimens
of the type series have a long, separate pre-apical
yellow mark, All specimens except those collected at
McDertnid Recks were taken on the flowers of
Chamaelaucium sp.

Ervmalogy

The specific name is derived from Gk charientas.
beautiful.

50 S. BARKER

Castiarina ferruginea sp. nov.
(FIGS 10, 1 Ql, 1 Q2, 3H)

Holorypes o, Wialki, WA, 18.ix.1957, S. Barker,
SAMA ] 21 309,

Allatype. 9, same data as holotype, SAMA If 21 310.

Paratypes: WA. 20 ov, Wialki, ix.1959, FH. Uther
Baker, WAMA; | c*, 88 km NE Wubin, 17.ix.1970, S.
Barker, SAMA, 10", Walyahoioning Rock (30° 38'S
18° 45'B), 9.41972, A. Baynes & R. Humphries.
WAMA. 3 cror 29 9, Muckinbudin, I0.%1979, R-
P. McMillan, WAMA; 29 ©, 8 km E Woolgangic,
22.x,1980, 5. Barker & PG. Kernpster, SAMA, 2c7 5,
Southern Cross, .J981, R. PB. McMillan, WAMA; 1.
Johnson Lake, 8,41, 1981, D. Knowles, MP WA; Joo.
19, Southern Cross, 8.x1983. Ro PB McMillan,
WAMA; 2cr 0", Eneabba, 15.%,1985, R. P. McMillan,
WAMA; Lor, 19, Dedari, 20.%,1986, M, Powell,
MPWA, Ler, 30 km E Lake King, I8xi,1988, M.
Powell, MPWA,; 2070", 29 9, N7T Transinitter, 32
km E Southern Cross, 71.%.1991, 'T. M.S. Hanlon,
MHSA; 200,49 9, Dedari, 21.4./991, T M.S
Hanlon, MHSA; 19, Karlgarin, Bessy Tolland,
WAMA.

Colour

Head dark coppery with blue-green rellections,
Antennae bronze, Pronotunr dark coppery with hlue-
ween reflections. Scutelluntdark purple. Elytra yellow
with dark maroon markings coalesced farming a
narrow yellow margin fram base to pre-apieal area,
tne intersticé wide fram base, two interstices wide at
humeral callus. two wide pre-medially continuously
two wide from post-nedially: row of four yellow spots
om each side, basal. round the remaining three elongate
Ventral surface coppery. Legs: femora and tibiac dark
copper, tarsi bronze. Hairs silver.

Shape and sculpture

Head punctured, median sulcus present, muzzle
short. Antennomeres 1-3 obconic, 4-11 toothed,
Prangtum closely punctured, small basal fovex
extending anteriorly to middle as glabrous line: apical
margin projecting medially, basal margin almose
straight. laterally parallel-sided at base, angled
outwards then ratinded to widest medially, rounded and
narrowed to apex, Scutellum scutiform, clongate,
glabrous, excavate. Elytra punctate-striate, intervals
convex, puhetuced and wrinkled, more so laterally than
medially; laterally angled out fram base, rounded al
humeral callus, concave, rounded post-medially and
narrowed fo unispinose apex; blunt marginal spine,
margin indented und straight to suture, apices hardly
diverging. Ventral surface with shallaw punctures,
edges of abdominal segments glabrous, elsewhere with
dense, flattened, feathery hair. S,: males, truncate;
females rounded,

Size
Males, 12,3, + 0.2% 4.0 + 0.07 mm (21). Females.
13.2 + 0.22 x 4.3 4 0.09 mm (15).

Aedéagus (Fig. 1Q)

Parameres angled outwards from basal piece, pre-
medially rounded then shallowly concave, rounded
apically. Penis sharp, sides obtusely angled away,
Hypophysis of basal piece broad, rounded apicilly.
Proctiger bilobed, bluntly pointed near mid-line,
laterally straight (Pig. 1 Q)).

Female terminalia (Fig. | Q2)

Proctiger bilobed, strongly pointed near (nid-line,
jaterally straight.

Remarks

The range of this species overlaps with thal of C
watulata sp, nov. which is approximately the same
colour but smaller. They can be distinguished on the
basis of the yellow elytral markings: in © ferneginea
there are four yellow spots on each elyinan and in CG
usiuiata three, as the first two ure fused forming an
elongate basal spot and the last two are separate. C.
ferruginea has a single, large marginal spine at the apex
olthe elytra and C. ustndata has (wo small spines. ‘The
sedeapi are different (Figs IQ, IV), both male and
female proctigers in both species are strongly lobed,
(hose of males are quile similarin size and shape bus
proctigers of female C ferruginea are more pointed
than in C. astalera which have black pigment spats
al the up ef each lobe; these spots are absent in the
other species (Pigs | Q2, 1 V2).

Etymelary

The specific name os derived from L ferniinens,
rust-coloured_

Castiaring ustulata sp. nov
(FIGS IV, 1 Vt. | V2, 3F)

Holatype. & , 8 km E Woolgangie, WA, 22-x. 980, S.
Barker & P G Kempster, SAMA § 21 Hi

Allotype: 9, same dala as holotype, SAMA 1 21 312,

Puratypes: WA; (ot, Dumbleyung, 5.x.1963, H, Udell,
WAMA; ic, 19. same data as holotype, SAMA;
3.9 9, Wiualki, 21.ix.1970, §. Barker, SAMA) 2c cr,
9 km SW Walyahmoning Rock, 9.x.1972, A. Baynes
& R. Humpbries, WAMA; 1a, 18.9 km WSW
Coolgardie, 18.ix.1976, R. J. Chinnock, SAMA; 19,
Dedan, 8.x.1978, T. M. 8. Hanlon, WAMA, Ic,
Muckinbudin, 10.x.1979, R. P. McMillan, WAMA;

SEVENTEEN NEW SPECIES OF CASTTARING Sl

19, Southern Cross, x.1981, R, P. MeMillan, WAMA;
lo, Bullfinch, 2.x.1983, B. Jones, MPWA; 1c’,
Southern Cross, 8.x.1983, R, P- McMillan, WAMA;
19. 28.km NE Peak Charles, 9.x,1983, G. Browning
& G. Mutze, SAMA; Io. 1.9, 45 km SW McDermid
Rock, 24.x.1985, T. F Houston & R. W. Thorp,
WAMA; 299, Dedari, 20.x.1986, M. Powell,
MPWA: 1c’, Bindoo Hill Nature Resetve, 27 km W
Mullewa, 12.1x.1987, T, F. Houston, WAMA, 4c cr,
299. N7T Transmitter, 37 km E Yellowiline,
21.x.1991, T. M. S. Hanlon, MHSA; 20. 49 >.
Dedari 22.x 1991, T. M. S. Hanlon, MHSA; 3cr ct,
19, Karlgann, Bessy Tolland, WAMA.

Colour

Head coppery. Antennae bronze. Pronotum coppery
Scutellum blue. Elytra yellow with brown markings
with coppery reflections coalesced and forming a
yellow margin from base to pre-apical area, one
interval thick medially, two intervals thick elsewhere;
three medial yellow marks on each side with an
clongate basal mark formed from the fusion of the basal
and pre-medial marks, round post-medial mark and
elongate pre-apical mark. Ventral surface either all
coppery or with coppery sternum and coppery-brawn
abdomen with blue reflections. Legs: femora coppery ;
libiae and tarsi bronze. Hairs silver.

Shape and sculpture

Head closely punctured, median sulcus present,
muzzle short. Antennomeres 1-3 obconic, 4-1L toothed.
Pronotum closely punctured, small basal fovea
extending anteriorly to middle as impressed line; apical
margin projecting medially, basal margin almost
straight; Jaterally parallel-sided at base, rounded to
widest pre-medially, rounded and narrowed to upen,
Seutellum seutiform, glabrous, excavate. Elytra
punctate-striate, intervals convex, punctured and
wrinkled. more so laterally than medially; laterally
angled out from base, rounded at humeral callus,
concave, rounded poast-medially and narrowed to
bispiiose apex: sharp marginal spine, minote sutural
sping, margin rounded and indested between spines,
apices hardly diverging. Ventral surface with shallow
punctures, edges of abdominal segments glabrous,
elsewhere dense flattened, feathery hairs. S_. inales
tnineate; females rounded,

Size
Males, 12.1 + 0.15 x 3.9 + 0.05 min (20). Females,
12.7.+ 0.19 x 4.0 + 0.07 tum (18).

Aedeagus (Fig. IV)

Purameres parallel-sided from basal piece, angled
outwards pre-medially, rounded apically, Penis sharp,
sides acutely angled away, Hypophysis of basal piece

medium width, apically rounded. Proctiger bilobed,
lobes near mid-line, blunt, apical edge straight.
laterally rounded (Fig. | V1).

Female terminalia (Fig. L ¥2)

Proctiger bilobed, lobes near mid-line, blunt, cacti
with a dark pigment spot at tip, apical edge straight,
jJaterally rounded, hairless.

Remarks.

See remarks under C. férruginea. Elyiral markings
ure the same in this species as in C. antarctica sp. noy
but they have non-overlapping ranges. The aedeagus
in C. aniurctica is shorter and broader than that in C
ustalata (Figs. 10, 1V). The proctigers of both sexes
in C. pstulata are bilobed with more highly developed
Jobes than those in C. attarctica (Figs | Ol, 1 O2,
} VI, - V2).

Erymology

The specific name is derived from L ustulares,
scorched,

Castiarina phagopus sp. nov.
(FIGS IW, | WL, 1 W2, 3B)

Holotxpe: o, 3 km E Gusnells, WA, 4.x1.1956, S,
Barker, SAMA [ 2) 313.

Allotype; 2 , Red Hill, WA, 2.ix, 1949, R. P. McMillan,
SAMA I 21 314.

Pararnpes: WA: 30° oO, 1, na data. SAMA; 20°",
Swan R., Lea, SAMA; 1 9, Bunbury. W. M. Mack,
1.3898, SAMA: 29 9, Perth, xi.1906, SAMA; 20°C".
LQ, Perth, .1913, SAMA; 40°, Perth, xi-1920, J.
W. Mellor, SAMA: 1o", 19, same data as allotype,
SAMA; |or, Mimmegarra, Dandaragan, 30.x.1955, 5.
Barker, SAMA; 20° c*, 2 km E Gosnells, 4.xi.1956.
§. Barker, SAMA; 15a", same data as holotype,
SAMA, 20°, summil Mt Cooke, 10.xi.1956, S
Barker, SAMA; Lor, 19. 70 km SE Perth on Albany
Hwy, 10.%1.3956, S. Barker, SAMA; Lo, foothills
Kelmscott, 21.«.1958, J. Baldwin, SAMA; 34c¢c,
29 9, Wilga, 26.x.1972, K. & E. Carnaby. SAMA;
2oo, Lesmurdie, 28.ix1955, 1 A. Watson, SAMA;
19, Julimar Forest, 24-x.J971, FH. Uther Baker.
SAMA: Goct, 39 9, Cataby Bk, 18«.1983, G.
Browning & G. Muize, SAMA; Lo, Gosnells,
7.x.1980, S. Barker. SAMA; 1o@, Mundaring Weir,
30.ix. 1980, T. M. S. Hanlon, MHSA;1¢, ML Dale,
29.1. 1980, T. M. S. Hanlon, MBSA,

C ‘vlour

Head brown with coppery reflections. Antennac
bronze. Pronoiunt brown with coppery reflections.

52 S. BARKER

Scutellum coppery With blue reflections. Elytra yellow
with the following markings: nartow blue basal
margin, other markings blue with coppery reflections
coalesced leaving a yellow margin frotit base to apex
from one to two intervals wide and two yellow marks
in the middle of each elytron in the form of an clongate
basal vitta tormed by the Fusion of the first three spots
and an elongate pre-apical mark. Ventral surface
coppery. Legs: femora coppery; tibiae and tarsi bronze,
Hairs silver.

Shape and sculpture

Head closely punctured, median sulcus present,
muzzle short. Antennomeres 1-3 obconic, 4-L toothed.
Pronotui clagely punctured basal fovea extending
forwards to jmddle as impressed line: apical margin
Projecting medially, basal margin almost straight;
laterally parallel-sided at base, angled outwards.
rounded medially at widest part, rounded and narrowed
to apex, Scutellum scutiform, glabrous, flat, Elytra
Punctate-striate, intervals convex, punctured and
wrinkled laterally, punctured and smooth medially:
laterally angled out. from base, rounded at humeral
callus, concave, rounded post-medially, uarrowed to
bispinose apex; very stnall sharp spines, margin
rounded and indented between spines, apives hardly
diverging. Ventral surface with shallow punctures,
edges of abdominal segments glabrous, elsewhere
dense flattened, feathery hairs. S,: males truncate;
females rounded.

Size
Males, Wt + O11 3.7 + 0.05 mm (Sl), Females,
ILS + O35 x38 + 012 pam (12).

Aedeagws (Fig. WW)

Parameres angled outwards from basal piece,
apically rounded. Penis sharp, sides achtely angled
away. Hypophysis of basal piece medium width,
apicully rounded Progtiger with small medial notch
in apical edge, laterally rounded (Fig. | WI).

Female terminalia (Fig, 1 W2)

Proctizger bilobed, apical edge straight, laterally
rounded.

Remarks

C phaeopus sp. Qov, cat be distinguished franz all
others in this complex By being the only species which
hag the fest three elytral yellow spots fused to form
an elongate basal mark with the fourth au clangate pre-
apical yellow mark

Etymalagy:
The speeitic name ws derived from Gk pbaes,
brown

Castiarina antarctica sp. nov.
(FIGS 10, 1 GI, 1 02, 3D)

Halorype: & , 64 km NE Esperance, WA, 18.x.1982,
5. Barker, P. G. Kempster & H. Vanderwoude, SAMA
12) 315,

Allotype! Q , same data as holotype, SAMA 1 21 316.

Paratypes: WA: Lo, Mt Ragged, 24.x, 1980, S. Barker
& PG. Kempster, SAMA; Lot, 13 km N Israelite Bay,
24.x%.|980, S, Barker & P. G. Kempster. SAMA. | or.
29 9, 24 km N Israelite Bay, 24.x, 1980, S. Barker
& PG. Kempster, SAMA; 1c, same data as holotype,
SAMA; Soa, 7 km N Dempster Rd Scadden Rd
crossing, Esperance district, 18.x.1982, S, Barker. P
G, Kempster & H. Vanderwoude, SAMA: 20° cr,
29 2. Parmangoes Rd 2 km NE Clyde Hill TO,
Esperance district, 8. Barker, PG. Kempster & H-
Vanderwoude, SAMA; | @, Israclite Bay, 21.x.1982,
§. Barker, PG Kempster & A. Vanderwoude, SAMA-
270, 19, 17 km NW Israelite Bay. 21.56.1982. S.
Barker, PG, Kempster & H. Vanderwoude, SAMA

Colaur

Head coppery. Antennae bronze, Pronotum dark
bronze medially, with coppery reflections laterally:
Scutellum coppery-purple. Elytra yellow with the
following dark brown markings with coppery
reflections coalesced leaving yellow margin, twe
intervals wide at apex and at humeral callus, one
interval wide medially; a row of yellow spats medially
on each elytron. basal and pre-medial cowesced
forming an elongate mark, post-medial more or less
reund, apical smaller and elongate. Ventral surtace
coppery. Legs: femora dull purple with coppery
reflections; tibiae and tarsi bronze. Hailes silver.

Shape and sculpture

Head closely punctured, median sulcus nurrow,
muzzle short. Antenndmeres 1-3 obconie, 4-1 toothed,
Pronotum closely punctured, ‘small basal fovea
extending forward to middle as glabrous line; apical
Inarein projecting medivlly, basal margin almost
Straight; faterally parallel-sided at base, rounded to
widest pre-medially, narrowed to apex. Seutellum
sculiform, tlat, glabrous. Elytra punetate-striate,
intervals convex, punctured and wrinkled laterally and
upically, smooth medially: laterally soyled out from
base. rounded at humeral callus, concave, rounded
postanedially and narrowed to bispingse apex, very
small marginal spine, minate medial spine. roargin
rounded and thdented between spines. apioes divergent
Ventral sutlace with slullow punciures, edges of
abdominal segments glabrous. elsewhere densely huiry,
hairs flat and feathery. S- males troneste: femules
rounded -

SEVENTEEN NEW SPECIES OF CASTIARINA 53

a ize
Males, 12.0 + 0.19 x 3.9 + 0.06 mm (14). Females,
12.0 + 0.22 5 4.0 + 0.09 mm (7),

Aedeauus (Fig. 10)

Parameres angled outwards from basal piece,
rounded pre-medially, parallel-sided post-medially,
rounded at apex. Penis sharp, sides acutely angled
away. Hypophysis of basal piece medium width.
apically rounded. Proctiger bilobed, lobes bluntly
pointed near miid-line. laterally straight (Fig. 1 Ol).

Female terminalia (Fig. | 02)

Proctiger bilobed, lobes blunt near mid-line, laterally
straight.

Remarks

The remarks. made under those tor C. ustulare sp.
noy. apply equally ty this species as these are the only
two species in this complex which have this elytral
pattern. They can be easily distinguished by differences
ih aedeagi (Figs 10, 1V) and in male and female
proctigers (Figs | Ol, 1 02, 1 VI, 1 V2). Alsa they
are allopatne,

Etymalogy
The name is derived from Gk anturktikos, southern

Castiarina nonyma sp. nov.
(FIGS IX, | Xi, 1 X2, 3A)

Holarype: o . Summit Mt Cooke, WA, 10,x(.1956, 5
Barker, SAMA. I 21 Al7.

Allolype> 9, Julimar Forest, WA, 24.%.1971, fH.
Uther Baker, SAMA I 21 Sib.

Pararypes’ WA: Jorg, 29 9, Beverley, E. F du
Boulay, SAMA) 2c o, Perth, SAMA: Ic’, Swan R..
SAMA, | @, xii.1913, SAMA; Zo" cr, same data as
holotype, SAMA; Lor, Mi Walker (32°05S' § 18°45"
fe) 16-x.1979, R, PR McMillan, WAMA, 1, Gosnells,
7.x 1980. S. Barker, SAMA, 19. Forresifield,
271.1978, T M.S. Hanlon, WAMA: 1, Eneabba,
17.*.1985, Ro P McMillan, WAMA,

Colour

Head and anlennae dark maroon, Pronotum dark
maroon with blue reflections medially, Scutetlum dark
maroon. Elytra yellow with maroon markings
coalesced to form yellow margins. wo intervals wide
al humeral callus and apically, one interval wide
medially; a medial row of four yellow spots on cach
elytron, basal and post-medial more or less round, pre-
inedial and pre-apical elongate, m about a quarter of the

specimens examined the first two coalesced forming
an elongate basal yellow mark. Ventral surface maroon.
Legs: femora maroon; tihiae maroon proximally,
bronze medially; tarsi bronze. Hairs silver.

Shape and sculpiure

Head closely punctured, median sulcus shallow,
muzzle short. Antennomieres 1-3 obconic, 41) toothed.
Pronotum closely punctured, small basal fovea
extending forwards to middle as impressed line; apical
margin projecting medially, basal margin almost
straight; laterally parallel-sided at base, rounded to
widest medially, rounded and narrowed to apex.
Seutellum scutiform, glabrous, excavate. Elytri
punctate-striate, intervals convex punctured and
wrinkled laterally and apically, smooth medially,
laterally angled out from base, rounded at humeral
callus, concayeé, rounded post-medially and narrowed
to unispinose apex; small, blunt marginal spine, margin
straight and indented ta suture, apices diverging.
Ventral surface with shallow punctures, edges of
abdominal segments glabrous, elsewhere with dense
flat. feathery hair. S,: males. truncate; femiules
rounded.

Size

Males, 1.0 4 0.28 x 3.5 + 0.09 min (10). Females,
Ut + 0.24 x 3.7 + 0.07 mm (4),

Aedeagus (Fig, UX)

Parameres angled outwards from basal piece, slightly
rounded post-medially then angled outwards, apically
rounded. Penis sharp, sides uculely angled away.
Hypophysis of basal piece medium width. apically
rounded. Proctiger with medial apical edge straight.
then angled forming (wo small broadly pointed lobes,
laterally rounded (Fig. | X41),

Female terminalia (Fig, 1 X2)

Proctiger bilobed, medial apical margin faintly
concave, lobes small and broadly pointed, laterally
rounded.

Remarks

No female speciniens associated with males at the
same collection locality were available This and the
following species © enigma sp. nov, have elytni
predominently eight spotted, althougt.a small number
of’ each has the first two spots coalesced, The two
species can be distinguished by differences in aedeays
which are short and broad 1 C. enigma and elungale
in & nertynid (Figs 1X, 1V) and in male and female
procigers. which are virtually unsculptured in C.
cnigma and bilobed with pointed |ohes in C. nomynue
(Fips | XU, 1X2, 1 YL, 1 ¥2), The distribution af
nunynid appears to be mainty to lhe east of the Darling
Searp fault line, while that of © enigma is to the west
of the Darling Scarp on the coastal Plain,

54 5. BARKER

Erymolozy
The name is derived from Gk anenymes, unknown

Castiarina enigma sp. nov.
(FIGS 1Y, 1 YL, 1 ¥2, 3C)

Holotype: o& , Regans Ford, WA, 9.x.1970, K. & E,
Carnaby, SAMA | 2) 319.

Allorype: 9. 6 knit S Gin Gin, WA, 30.ix.1956, 8.
Barker, SAMA J 21 320.

Paratypes: WA: lo, 19, no data, SAMA; 10, E
Ashby, SAMA; | 2, Perth, SAMA; 19, Perth s7.1905,
SAMA; 2a o, Perth, x.1913, SAMA: 20°, 19,
same dala as allatype, SAMA, 80° or, 1.9, same data
us holotype. SAMA.

Colour

Head and antennae dark maroon with blue
reflections. Pronotum dark maroon, with blue
reflections medially. Scutellum dark maroon with blac
reflections. Elytra yellow with brown markings with
coppery reflections coalesced leaving a yellow margin
two intervals Wide at humeral callus and apically, one
interval wide medially; row of four medial yellow spots
on each elytron, basal and post-medial more or less
round, pre-medial and pre-apical elongate. Ventral
surface and legs coppery. Hairs silver.

Shape and sculpture

Head closely punctured, median sulcus shallow,
muzzle short, Antennomeres J-3 obconic, 4-11 toothed
Pronotum closely punctured, narrow basal fovea
extending forwards to middle as glabrous line: apici!
margin projecting medially, basal margin rounded from
base to widest medially, rounded and narrowed to apex.
Seutellum scutiform, glabrous, lat, Elytra punctate-
striate, intervals convex, punctured and wrnkled
laterally smooth medially; laterally angled ouc from
base, rounded at humeral callus, concave, rounded
post- medially and narrowed to bispinose apex; small,
blunt marginal spine, minute sutural spine. margin
rounded and indented between spines, apives diverging.
Ventral surface with shallow punctures, edges ol
abdominal segments glabrous, clsewhere densely hairy,
hairs flat and feathery. 8,: males truncate; females
rounded,

Size
Males, 10.7 + 0.17 x 3.4 + 0.06 mm (15), Females.
8 +4 0.37 x 4.0 + 0.13 mm (6)

Aedeavus (Fig. 1V)

Broad. Parameres angled outwards from basal piece,
rounded apically, Penis sharp, sides obtusely angled

away. Apophysis of basal piece medium width, apically
rounded. Proctiger broadly rounded at apex. sides
rounded (Fig. 1 YI),

female terminalia (Fig. | ¥2)
Proctiger rounded.

Remarks

The remarks under C. nonyma apply equally to this
species.

Etymology
The name is derived from L. aenigma, mystery.

Castiarina aura sp. nov,
(FIGS IS. 1 S1, | $2. 31)

Holotype: Oo, 131 kn S Exmouth, WA, 12.1x,1984, M,
Powell, WAMA.

Allatype: 9 , same data as holotype. WAMA.

Paratypes: WA. 12, 50 km N Kalbarri ‘T.0.,
20.viii.1978, T. M.S. Hanlon, WAMA; 10", Yardie Ck,
I8.vili. 1983, M, Powell. MPWA: |ar_ Coral Bay,
10,1x.]984, M. Powell, MPWA; Lor, 1°, Carnarvon,
28viii, 1987, A. Hay, SAMA; 1 2, 94 km S Learmonth,
2.1x.1995, Powell & Kershaw, MPWA: 19, 62 km S
Learmonth, 4.ix,1995, MPWA; Io’, 19, 26 km §
Learmonth, 3.ix.1995, Powell & Kershaw, MPWA,

Cnlour

Head coppery: Antennae bronze with coppery
reflections. Pronotum coppery, with medial blue-green
reflections. Scutellum blue-green. Elytra yellow with
the following elytral markings: markings coalesced.
coppery apically, with blue-green reflections along the
suture and over the humeral callus forming a yellow
margin two intervals wide, medial row of four yellow
spots on each elyiron, basal and post-medial more or
less round, pre-medial and pre-apiea! elongate. Ventral
surface and legs coppery: Hairs silver.

Shape and sculpture

Head closely punctured, median sulcus shallow,
muzzle short, Antennomeres 1-3 obconic, 4-1] toothed.
Pronotum closely punctured. basal fovea extending
forwards to middle as glabrous line; apical margin
projecting medially, basal margin almost straight:
laterally parallel-sided at base, rounded to widest before
middle, rounded and narrowed to apex. Seuteilum
sculilorm, wrinkled, excavate. Elytra puncutte- striate,
intervals convex, punctured and wrinkled laterally,
smooth medially; laterally angled outwards from base

SEVENTEEN NEW SPECIES OF CASTIARINA

rounded at humeral callus, concaye, rounded after
middle, tapered to unispinose apex; marginal spine
rounded, margin rounded and indented to suture, apices
diverging. Ventral surface with shallow punctures,
edges of abdominal segments glabrous, elsewhere
dense, flat, feathery hairs, S,: males truncate; females
rounded.

Size
Male, 1.9 + 0.42 x 4.0 + 0,15 mm (5). Females,
12.9 + 044 44.3 4 0.18 mnm (6).

dcdeagus (Fiz. 1S)

Parameres angled outwards from basal piece,
rounded post-medially, parallel-sided, rounded at
apices. Penis sharp, sides acutely angled away.
Apophysis of basal piece wide, apically rounded.
Proctiger bilobed, lobes blunt (Fig. | Sl),

Female terminalia (Fig. 1 $2)

Proctiger bilobed,
mid-line.

lobes strongly pointed near

Remarks

C. aura sp. nov, and C. ferruginea sp. noy. arc
similar in that both have four separate yellow spots on
each elytron and a large single marginal spine on the
apices of the elytra although the elytral colour is
different, C, aura being red with green reflections
while C. ferruginea is brownish. Aedeagi differ as they
are shorter and narrower in C. aura than they are in
C. ferruginea (Figs 1Q, 1S). The proctigers of C
Jerruginea males are strongly bilobed while those of
C. aura are faintly bilobed (Figs | QI, | SI). The
proctigers of females of both species are bilobed and
pointed but the lobes are further apart in C. aura than
they are in C. ferruginea (Figs 1 Q2, 1 S82).

Etymology
The name is derived from L aura, glow.

Key to WA species of C. parallela complex

|, Elytra background colour dark blue 2
Ulytea background colour brown,
red or green 3
2, Elytra with 8 round, yelluw marks oelopunctala
Barker

Elytra with 2

yellow marks
3. Bilytra bright red or partially

or wholly brassy green 4

Elytra brown ar red-brown 6
4. Elyta bright red, elytral apices: witty (wo

amall spines, elyinal spots coalesced into

single elongate yellow mark

on each side

Elytra bright ced or partially or wholly

brassy green, elyira with 4 yellow spots 4

found. 6 elongate
azure SP. NOY.

chariemessa Sp. Noy

nm
in

5. Elytral apices with | large spine aura sp, n0v.
Elytral apices with 2 small,
blunt spines cracivalar (L & G)
6, Some elytral marks coalesced forming
6 or fewer yellow marks 7
Elytra with 8 yellow marks 10

7. First 3 yellow marks on each elytron
coulesced, forming | elongate anterior
mark and } small elongale mark
posteriorly on each side
First 2 yellow marks on cach elytron
coalesced, forming | clongate anterior
mark and 2 small elongate marks
posteriorly on each side by

4. Pronotum bronze, elytra dull brown,
male procliger slightly sculptured,
female proctiger unsculptured, Largest
member of group.

Pronotum,, elytra with coppery reflections

9. Proctiger bilobed, lobes pointed, in

females With pigment spot al up

Procliger medially notched, lobes blunt

without pigment in females

Elyiral apices with single large spine

Elytral apices bispinose

Il, Aedeagus broad, proctiger unsculptured
in both sexes
Aedeagus narrow, proctiger sculptured
in both sexes

phacopus sp, nov.

adusia sp. nov.

ustulaia sp, nav,

antarvticd sp, NOV,

10, Jerruginea sp. nov.
I

eHigM Sp. NOY,
nonyma 8p. nov.

Castiarina nullarborica sp. nov.
(FIGS U, 4D)

Holotype: o, 5 km E Eucla, WA, 28.x,1989, K. L.
Walker, NMVA.-

Allotype: 2, Nullarbor Plain, SA, SAMA J 2i 321,

—s5 TA B =
fe y ’ 4 . f) *
! ' rit \ e if
= 4 ; —
es

qh ha)
: th f Jem pe \

Fig. 4. Habitus illustrations of the following Castiarina
species. A, Castiarina Jackhasenpusch sp. nov, holotype.
B.C hemizostera sp. nov. holotype. C. C. paulhasenpuscht
sp.nov. holotype. D. CG aullarbarica sp. nov. holotype.
B.C. demtarci sp. nov. allotype.

56 5. BARKER

Pararypes: SA: (o>, 32 km E Eucla, 1).xii.1984, M.
Powell, MPWA. WA; Ic, same data as holotype,
SAMA.

Colour

Head dark blue. Antennae blue-green. Pranotum
hronze medially, dark blue laterally. Seutellum dark
blue Elytra yellow with red margin and the following
black markings. with blue reflections: narrow basal
margin: broad pre-medial fascia not reaching margin,
distally angled anteriorly: broad post-medial fascia
reaching margin, spade-shaped apical mark coverins
apex and spines, last Iwo marks connected broadly
along suture, Ventral surface bronze. Legs dark bluc-
Hairs silver,

Shape and sculpiuré

Head closely punctured, median sulcus broad,
muzzle short. Amennomeres 1-4 obconic, 5-11 toothed
Pranotont closely punctured, ginal! basal fovea; apical
mately straight, basal margin histuare, laterally
parallel-sided al base, rounded to widest pre-medially,
counded and narrowed to apex, Scutellum scutiform,
glabrous, flat. Elytra punctate-stnate. intervals convex,
punctured; laterally angled out from base, rounded al
humeral callus, concave, rounded. post-medially and
narrowed (0 bispinose apex. sharp marginal spine,
small, sharp sutural spine, margin rounded and deeply
indented between spines, apical margin subsernite,
Ventral surface with shallow punctures. edges of
abdominal segments glabrous, elsewhere moderately
hairy, hairs medium length. S,: truncate in both sexes,
Male legs 2-3; pulvilli absent on tarsomeres 1-3
replaced with a small double, median spine,

Size
Males, 9.2 x.3.5 mm (3), Female, 10.6 x 4.5 mm (1).

Aedeagus (Fiz WU)
Wedge-shaped.

Remarks

This species is closest to the morph of © placida
(Thomson) which has a red margin and occurs on (he
west cous of WA and on Rottnest Is.. WA. C.
nullarborica is a smaller species than ©. placida and
the male genitalia differ in size and shape (Figs IL, LI)

Etymology

The name is derived from Nullarbor Plain, the area
where this species occurs.

Castiarina demarzi sp. nv.
(FIGS 1H, 46)

Holorype;> o , Burardy HS (27°34'S, 4°40'E) WA.
19.vii.1980, C. A. Howard & T. F. Houston. WAMA.

Allorvype. 9, 36 km NE Tamala Station, Shark Bay,
WA, 28.1x.1988, D. Knowles, WAMA.

Paratypes; WA, 19, same data as allotype, MPWA,
19, 26 km NE Tamala Station, 6.x.1988, D. Knowles,
SAMA: 19, found in seed collection fram NW coast.
29.vili, 1986, H. Demarz, HDWA.

Colour

Head, untennae and pronotum bronze with or
without coppery reflections. Scutellum blue or bronze
Elytra yellow with the following black markings with
blue reflections: narrow basal iuargin; pre-med
fascia not reaching margin, distally angled anteriorly,
post-medial fascia reaching margin, projecting
anteriorly in middle of cach elytron, mark covering
whole apex, marks connected along suture in holotype
but not allotype. Ventral surface and legs eoppery-
Hairs silver.

Shape und sculpture

Head closely punetured, median sulcus broad,
muzzle very short. Antennomeres compressed, 13
obeonic, 4-11 toothed, Pronotum closely punctured,
basal fovea extending anterlorly to middle as glabrous
line, basal notches represented by glabrous area on eayh
side closer to margin than middle: apical margin
projecting medially, basal margin barely bisinuale;
laterally angled outwards from base, rounded to widest
after pmddle, rounded and narrowed to apex. Scutellum
scutiform, glabrous, flat. Elytra punetate-striate,
intervals convex, punctured: laterally angled out from
base, roanded at humeral callus, coneaye. rounded
post-mediully and narrowed to bispinose apex; sharp
marginal spine, small, sharp sutural spine, apices
diverging, apical margin sabserrate. Ventral surface
with shallow punctures, edges af abdominal segments
glabrous, clsewhere short sparse hair $,: male
truncate, females rounded.

Size
Male. 10.0 % 3.7 mm (1). Females, 1L7 + 0.22 x 4.6
+ ON mm 14).

Aedeagus (Pig. 1H)

Parameres parallel-sided from basal piece, roundes
pre-nedially then narrowed to apex. Penis blunt, sides
acutely angled away. Hypophysis of basal piece
medium width, apically rounded.

Remarks

The Structure and. elytral markings of this species
resemble C. buecolica (Kerremans).. However C.
bucelica has head, pronotum. and ventral surface green
and there are size differences between aedeagi, that
ot C, demurzi being smaller than that of C buedlica
(Figs IG, 1H)

SEVENTEEN NEW SPECIES OF CAST/ARINA 7

Erymoloey

The name honours Mr Herbert Demarz, Guilderton.
who has generously assisted my research by loaning
specimens for many years.

Castiarina jackhasenpuschi sp. wv.
(PIGS IL, 4A)

Holotype: ot, Cardwell Ra., Qld. 22.x01995) J.
Hasenpuseh, SAMA L 21 322.

Allowpe: , Cardwell Ra,, Qld, 22.x17.1995, P.
Hasenpusch, SAMA T 21 323,

Colour

Head reddish-hronze, muzzle green-bronze,
Antennae green. Pronotum reddish-bronze with a
curved blue bar. concave inwards, on each side of the
mid-line fron base to apex, Seutellum green wath
yellow reflections. Blytra yellow with the following
black markings: broad basal margin; broad pre-medial
fascia with ends expanded anteriorly reaching basal
nlargin and posteriorly reaching margin enclosing a
yellow basal spot and a yellow spot on margin at
humeral callus; broad post-medial fascia reaching
Margit and mark covering whole apex: yellow medial
fascit not reaching margin: yellow post-medial fascia
Hol reaching suture or margin; ventral surface green
with yellow reflections, legs blue-green. Hairs silver,

Shape and sculpture

Head closely punctured, median sulcus present,
murzle short, Antennomeres 1-3 obconic, 4 half-
toothed, 5-11 toothed, Pronotum closely punctured,
small basal fovea; apical margin straight, basal margin
hisinuate: laterally angled inwards from base. rounded,
widest before middle, rounded and narrowed to apex,
Scutellum scutiform, punctured, flat, Elytra punciate-
striate, intervals convex, punctured; laterally anvled
out from base, rounded at humeral callus, concave,
rounded post-medially, narrowed to bispinose apex;
large sharp marginal spine, minute sutural spine,
thargin indented and straight between spines. apices
hardly diverging. Ventral surface with shallow
punctures, edges of abdorainal segments glabrous,
elsewhere sparse very short hairs. §,: male rounded:
female slightly rounded and turned under.

Size
Male, 6.4 x 2.5 mm (1). Female, 69 x 2.6 mm fl).

Aedeagus (Fig. 1L)

Narrow and elongate. Parameres angled inwards
from basal piece, parallel-sided, rounded at apex. Penis
sharp. sides acutely angled away. Apophysis of basal
piece wide, apically rounded.

Remarks

This species superficially resembles C. cvdista
(Rainbow). [t js however, smaller, the structure of the
aniennomeres differs as the 41h antennomere of C.
ceydista is fully toothed and male genitalia are dissimilar
(Figs (kK, IL).

Etymology

This species is named to honour Mr J, Hasenpusch,
Innisfail, who has generously supported my research
by loguning specimens and providing information.

Castiarina paulhasenpuschi sp. nov.
(FIGS IF, 4C)

Holotype; &, Marsupial Ck vear Croydon. Qld,
241.1995. P. Hasenpusch, SAMA [ 21 324.

Allotype: ©, sate data as holotype, SAMA 1 21 325,

Faratypes: QW: 19, same data as holotype JAQA,
20°70, Marsupial Ck, 1-15 iv.1995, PB Hasenpuseh,
JHQA,; Io, 19, Marsupial Ck, 2.yi.1995, J,
Hasenpusch, JHQA,

Colour

Head, bronze, Antennae bronze with green
reflections. Pronoiuin bronze. laterally with green
reflections. Scutellam. green. Elytra yellow with the
following black markings with blue reflections: narrow
basal margin, in the holotype a mark covering most
of apical half in torm of a post-medial fascia connected
to the apical mark leaving a pre-apical yellow spot on
each margin; in one specimen the fuscia is reduced
to two small black spots on the margin. Ventral surface
and legs green. Hairs silver.

Shape and sculpture

Head closely punctured, glabrous, median sulcus
present, muzzle yery short. Antennomeres
compressed. 1-3 obconic, 4-IL toothed. Pronotum
closely punctured, glabrous, basal fovea extending
forwards but not reaching middle, basal notches on
each side closer to margin than middle; apical margin
projecting medially, basal margin bisinuate; laterally
parallel-sided at base, rounded to widest at middle,
rounded to apex. Sculellum scutiform, glabrous,
excavate. Elytra punctate-stnate, intervals conyex, more
so apically; laterally angled out from base, rounded
alt humeral callus, concave, rounded post-mediually,
narrowed to bispinose apex; sharp marginal spine,

small sharp sutural spine, margin indented and rounded

between spines, apices diverging, apical margin
strongly sub-serrate. Ventral surface with shallow
punctures, edges of abdominal segments glabrous,

58 5. BARKER

elsewhere moderately hairy, hairs medium length. 3:
males truncate; females bilohed, each lobe with four
claws.

Size
Males, 13.2 + 0.16x 4.5 + 0.07 mm (4). Females,
13.5 4 0.45 x 4,7 + 0.18 mm (3).

Aedeagus (Fig. LF)

Parameres parallel-sided from basal piece, rounded
medially, parallel-sided, rounded to apex Penis sharp,
sides. obtusely angled away. Apophysis of basal piece
medium width, apically rounded.

Remarks

The distinct colour and pattern of this species
distinguish it from all other species, as does the
structure of the last visible abdominal segment in
females, in which the claws are unique. The specimens
examined were all caught by use of a colour lure in
an area. where no plants were flowering.

Erymology

The species name honours Master Paul Hasenpusch
its discoverer.

Castiarina hemizostera sp. nov.
(FIGS IN, 4B)

Holotype: of, Cardwell Ra., Qld, 22-xi.1995, J.
Hasenpusch, SAMA 1 2! 326.

Allorype: 9, Cardwell Ra., Qld,
Hasenpusch, SAMA T 21 327.

Paratypes: Qld: 1o, Cardwell Ra., 19.x1i,J995, 1.
Hasenpusch, JHQA; 20°C", same data av holotype.
JHQA: lor, 9. 22.xi1.1995, P. Hasenpuseh, FHOA;
3a oO, 30,xij, 1995, J. Hasenpusch, JHQA.

Colour

Head black with blue-green reflections, muzzle blue-
Aniemnae green. Pronotum with purple-green
reflections medially, blue-green laterally. Scutellum
preen. Elytra yellow: with black markings with bluc-
green and/or purple reflections coalesced leaving the
following yellow marks; pre-medial yellow fascia
reaching margin but not sutare, broad pre-apical yellow
fascia reaching. margin but nat suture. Ventral surface
black with bronze reflections. Legs blue. Hairs silver.

24. xi1.1993, J.

Shape and sculpture

Head closely punctured, median suleus broad.
muzzle short, Antennomeres 1-3 obconic, 4-11 toothed.
Pronotum heavily punctured, basal fovea extending
forwards to apical margin as impressed line: apical
margin straight, basal margin bisinuate; laterally
parallel-sided at base, rounded to widest pre-medially,
rounded and narrowed to apex, Scuieslum scutiform,
punctured, flat. Elytra punctate-stnate, intervals
convex, heavily punetured: laterally angled out from
base, rounded at humeral callus, concave, rounded
post-medially and narrowed to bispinose apex, sharp
marginal spine, minute sutural spine. margin rounded
and indented between spines, apices diverging Ventral
surface with shallow punctures, edges of abdominal
seginents glabrous, elsewhere moderately hairy, hairs
short, S,: males truncate; females rounded.

Size
Males, 70 + 0.2 x 2.5 + 0.09 mm (8). Females.
7.9 x 3,0 mm (2).

Aedeagus (Fig.. IN)

Parameres angled outwards from basal piece,
rounded and widened post-medially, rounded at apex
Penis sharp, sides acutely angled away. Apophysis of
basal piece wide, apically rounded.

Remarks

This species is allied to C.. bella (Saunders) and 1s
closest to C. aglaia (Barker). However, the post-medial
fascia in C. avlaia is red and the male genitalia differ,
(Figs IM, IN)

Etymalogy

The name is derived from Gk hemisys, hall, Gk
zoster, bell.

Acknowledgments

I am endebted to the following for assistance: Dr
T. FE. Houston, WAMA; Dr K. Walker, NMYVA, Mr
T. Weir, ANIC; Dr E, G, Matthews, SAMA, Mr D
J. Williams, Mr P. G. Kempster and Ms H
Vanderwoude, Department of Zoology, University of
Adelaide. Iam endebted to the following collectors tur
the loan and gift of specimens: Dr. F H- Uther Baker,
Cottesloe; Mr H. Demarz, Guilderton; Mr T. M. &.
Hanlon, Hunters Hill; MrJ. Hasenpuseh, Innisfail: Mr
R. P. McMillan, Kallaroo; Mr M Powell, Melville,
Mr S Watkins, Caparra.

References

Amneyy, R, Hy Jp, SAMUELSON OG. A. & Nisiwipa, G. M
(1993) "The Insect and Spider collections af the world”
2nd ed (Sandhill) Crane Press, Gainsville),

Banker, 8, (1Y87) Eighteen new species of Siipmodera

(Castiarina) (Coleuptera: Buprestidae), Trans, R. Sec, ¥
Aust. TE, 133-146,

(1993) Seventeen new species of Australian
Buprestidae (Insceta; Coleoptera) and a host plant ov
Castiarina uplani (Barker). [hid. (V7, (i-26.

SEVENTEEN NEW SPECIES OF CASTIARINA 59

____ (1995) Eight new species of Australian Buprestidae
(Insecta: Coleoptera). Ibid. 119, 149-156.

BLACKBURN, T. (1890) Further notes on Australian
Coleoptera, with descriptions of new genera and species,
Ibid. 13, 121-160

Devaquet, C. M. (1956) Notes on Australian Buprestidae,
with descriptions of three new species and two subspecies
of the genus Stigmodera, Subgenus Castiarina. Proc. Linn.
Soc. N.S.W. 81, 153-156.

Genin, J. J. B. (1855) Coléoptéres nouveaux ou peu Connus
Decade 1. Bull. Soc. Hist. nat. Metz. 7, 53-65.

KERREMANS, C. (1890) Espéces inédites du genre Stigmodera
Eschscholtz. Bull. Soc. Ent. Belg. 1890, 40-49.

Laporte, F. L. & Gory, H. (1838) “Histoire naturelle et
iconographie des inséctes coléoptéres, publiée par
monographies séparées.” Vol 2 Suite aux Buprestidae (P.
Dumenil, Paris).

SAUNDERS, E. (1868) A revision of the Australian Buprestidae
described by the Rev. F. W. Hope. Trans. Roy. entomol.
Soc. Lond. 6, 1-67.

Wart, J. C. (1979) Abbreviations for entomological
collections. N.Z. Zool. 6, 519-520.

A NEW GENUS AND THREE NEW SPECIES OF
CECIDOMYIIDAE
(DIPTERA) FROM OLEARIA SPP. (ASTERACEAE)
IN AUSTRALIA

By PETER KOLESIK*

Summary

Kolesik, P. (1996) A new genus and three new species of Cecidomyiidae (Diptera)
from Olearia spp. (Asteraceae) in Australia. Trans. R. Soc. S. Aust. 120(2), 61-67, 31
May, 1996.

A new gall midge genus, Trigonomyia, and three new species, T. ananas from Olearia
ramulosa (Labill.) Benth., T. cristata and T. tulipa both from O. axillaris (DC.) F.
Muell. Ex Benth., are described. Detailed descriptions of the adults, larvae, pupae and
galls are given. The species are distinguished from each other by both their
morphology and the appearance of their galls. The new genus is diagnosed and placed
in the tribe Oligotrophini within the supertribe Lasiopteridi of the subfamily
Cecidomylinae.

Key Words: Cecidomyiidae, Trigonomyia ananas sp. nov., Trigonomyia cristata sp.
nov., Trigonomyia tulipa sp. nov., Olearia ramulosa, Olearia axillaris, South
Australia.

Transactions of the Reval Society uf S, Aust, (1996), 120(2), 6167.

A NEW GENUS AND THREE NEW SPECIES OF CECIDOMYTIDAE (DIPTERA} FROM
OLEARIA SPP. (ASTERACEAE) IN AUSTRALIA

by PETER KOLESIK*

Summary

Kotesix, P. (1996) A new genus and three new species of Cecidomyiidae (Diptera) from (learid spp, (Asteraceae)
in Australia, Trany. R. Sec. 8. Aust. 120(2), 61-67, 31 May, 1996

A new gall midge genus, Trigonomyia, and three new species, 7. ananas trom Olearia ranudosa (Labill.) Benth..
T evistata and T. sulipa both from O. axillaris (DC.) F, Muell, ex Benth,, are described. Detailed descriptions
of the adults, larvae, pupae apd gal!s are given, The species are distinguished from cach other by both their
morphology and the appearance of their yalls. The new genus is diagnosed and placed in the tribe Oligotrophini
within the supertribe Lasiapteridi of the subfamily Cecidomyiinar.

Key Worps; Cecidomyiidae, Trigonomyia ananas sp. nov, Trigonomyia eristate sp, nav., Trigoremiyia tulipa
sp. nov., Qlearia ramulosa, Olearia axillaris, South Australia.

Introduction

Three new pall midge species are described here that
were found galling flowers of two species of ihe daisy-
bush, Olearia Moench (Asteraceae). Trigonemyia
anantas sp. nov. was found in Black Hill Conservation
Park, near Adelaide, infesting the twiggy daisy-hush,
Q. ramulosa (Labill.) Benth. Trigenori\ia cristara sp-
nov, and 7 fulipa sp. nov. were discovered at
Beachport, in the Lower South-East of South Australia,
both attacking the coastal daisy-bush, OQ: axilariy
(D.C.) FR Muell, ex Benth,

Olearia includes some 75 species in Australia and
25 in New Zealand and New Guinea (Cooke 1986).
Olearia ramulasa is an aromatic shrub, about 1,5
metres high, much-branched, witha woody stem anil
numerous, small, yellaw-white flawers which occurs
throughout Australia in mallee, woodland and coastal
scrub (Cooke 1986). It is common in Black Hill
Conservation Park where it often forms dense localised
populations on poor stony soils. Clearia axillaris is
a 2-3 metres high shrub, morphologically distinguished
from ©. ramulosa by larger leaves and minute ligules.
Olearia axillaris forms a dense scrub on coastal sand
dunes of moderate and temperate Australia (Cooke
1986) and is a dominant plant along the Beachport sea
shore.

A new genus is proposed for the three new gall
midye species. [t is placed in the subfamily Cecido-
myiinae and supertribe Lasiopteridi. It ts compared
to Rhopalomyia of the tribe Oligotrophini from which it

* Department ef Borticulure, Viticulture and Oenology
University of Adelaide PMB | Glen Oxmond 8. Aust. 5064,

is morphologically. distinguished by the male genitalia
and the larval neck segment. The three new species
differ from each other in morphology of the male
ponostyli, the ovipositors, the pupal prothoracie
spiracles, and the galls.

Material and Methods

Three distinct kinds of flower galls were sampled.
One was collected trom Q rarmu/osa in Black Hill
Conservation Park near Adelaide (17,ix.1994) and twu
from O. axillaris on coastal sand dunes at Beachport
(6,x.1994) The two types of galls collected. from 0.
axillaris were kept in separate bags and all galls were
processed in two ways according to the method
previously described (Kolesik 1995). Microscope
mounts of the type specimens were prepared by
maceration in 20% KOH, followed by processing
through distilled water, 70% and 99% ethanol, xylene
and were mounted jn Canada balsam for examination
by phase-contrast and bright-field microscopy. Larvae,
pupae and pupal skins were mounted dorso-ventrally,
Adults were dissected into four (females) or five
(males) pieces and their particular parts mounted
separately: wing, head frontally, thorax Jaterally, female
abdomen dorso-ventrally or laterally and male genitalia
and abdomen dorso-ventrally. Measurements were
made with an eyepiece graticule. Drawings were done
with the aid of a camera lucida. The type series ane
other materials retained in 70% ethanal are deposited
in the South Australian Museum, Adelaide [SAM], the
Australian National Insect Collection, Canberra
[ANIC] and the United States National Museum,
Washington DC [USNM]. Adult terminology follows
usage in Gagné (1981). Both larval and pupal
ierminglogy follows Gagné (1994),

ot P. KOLESIK

Gers Tngenemypia sen. mire,

Type species: Thigarremvnivia arnanar
Adults

Wings with Ry joining Caf wing apex, Re uhseng.
Ri joining C near wing mid-length, Ma absem, Cu
forked, Maxillary palpus with 3 segments, palpiger
well developed, Eye facets rounded, eye bridge 2-4
facets medially, Antenna with variable number of
flageiomeres, usually 16-18, first and second only
Weakly separated. Flagellomeres cylindrical with neck
longer in malethan in female, with long and stout setae
ay up ty three whorls; cirournfilar koaps short, forming
sparse network, similar in both sexes. Empodia longer
than claws, pulvilli stout, about half claw length. Claws
simple, broadly curved. Abdomen: lengite U sclerotized
in both sexes, with posterror setal row only, tergites
T- VIL in male and If - Vita female sclerotized,
with single posterior setal row inierrupted mesally, pair
of sparse setal fields laterally and one seta in both
anterior corners, female tergite VIL] not sclerotized,
with triangular field of scattered setae at posterior end;
stermtes IL - YIN in male and I - VIF in female
sclerotized, witl) dense, uninterrupied posterior band
of Selue, scattered setae anteriorly and isolated paw of
setae Of posterior end. Male genitalia: gonocoxites
cylindrical, unlobed, setose and setulose; gonastylis
situated caudally on gonocoxite, cylindrical, shehtly
tupenmng towards apex, with short apical tooth
comprising one elaw and several firm bristles, setose,
setulose throughout; cerci bilobed, with several stout
sclae on each lobe, setulose; hypoproct bilobed, with
one long sets on each lobe, setulose, parameres drveded
into LWe parts, basal lobe simple, asetose. setulose,
apical lobe asetulose, bearing 5-6 paralle) running
Janieliae, asetulose, bearing altogether 6-8 lange, serose
papillae, aedeagus robust, strongly sclerotized venarally
and apically, with apical end triangular. Oviposator;
protrusible; cerci fused into single, terminal lamella,
triangular in dorso-ventral view, with numerous strong
sclae, Setulose; hypoproct trapezoid in dorsa-ventral
yiew, short, bearing 2 setae posteriorly, setulose.
Pupra

Integument of abdominal segments cavered by
spiculue, Prothorax and abdominal segments |-VUIt
wilh spiracles. Antennal horns short, angular. Cephalic
pair of papillae with strong, long setae. Frons with one
pair of upper frontal weakly sclerotized depressions
and one of 2 lower facial papillae on each side with
short seta. Abdonunal segments I-VI with | pair of
ventral papillae, 2 pairs of pleural papillae and 3 puns
of dorsal papillae, Abdominal segments VIM and LX
with | pair of ventral, 2 pairs of plearal papillae and
| pair of dorsal papillae. All papillae setase.
Larva

Integument completely covered with dense spiculac.

Head: strongly sclerotized, posterolateral apodemes
shorter than one fourth of head capsule length,
untennge about three times longer than wide at bise.
<onical. Neck segment with | pair of setase pleural
papillae. Thoracic seginents without spatula but with
depression where spatula would normally appear, |
pair of Ventral papillae, 2 pairs of lateral papillae, 2
pairs of pleural papillae, 3 pairs of dorsal papillae.
Abdominal segments T-VIE with two. sternal
depressions, | pair-of ventral papillae, 2 pairs of pleural
papillae, 3 pairs of dorsal papillae, Abdominal segment
VIM with 2 puirs of ventral papillae, 2 pairs of pleural
papillac, { pair of dorsal papillae. Abdominal segment
LX with 4 pairs of terminal papillae, All papillae setose
excep! thoracic jateral ones. Setao long and clearly
apparent on all papillae with exception of ventral
papillae which are only slighty longer than integv-
mental spiculac.
Enrmolagy

Thganonwea combines “trgenge’, Gk lor trisnygle
Wluch refers to the shype ofthe apical end of aedeagus
and “myia”. Gk for fly. commonly used as suffix for
genera of Cecktomtyiidac.

Remarks

Thigonortyta belongs to the tribe Lasiopterid) because
it has male parameres and an irregular number of
antennal flagellomeres. The new genus belongs to the
Oligotrophini and within that tribe to 2 group of genera
that includes Rhepalomyia, and for which the tribal
name Oligotrophini (in the strict sense, not including
Dasineurini), is available. These genera share the
following derived characters: reduction of the palpus
to three or dewer segments, relatively short parameres
that do mot clasp the aedeagus along its full length,
\arvae each Iiving im @ separate cell in galls with
fupation occurring inside the larval cell. Other,
probably primitive characters shared by all members
of Rhopalomyia and telatives are the completely
setulose male gonostylus and the never divided female
cighth abdominal tergite, even when the ovipositer is
elongate (Gagné et al. in press). Trigonomyia differs
from Rhopalomyie in several ways, In Trigenomyia the
parameres are divided into two distinct lobes, the
uedeagus is sclerotized apically and the larva has setose
collar papillae. None of these characters has been noted
in Rhepalemyia or related genera, although presumably
separately derived, indistint to distinct divisions can
be found in the purameres in Dasineura (sl),
Lasioptera and Ledomyia (Gagné 1994).

Trigonomyia ananas sp. ner,
(PIGS 1, 2, 4-8, 1-17, 19, 21, 22)

Holotype: @. Black Hill Conservation Park, Sour

NEW CECIDOMYIIDAE FROM OLEARIA

6: ! in 4

Fig. 1. Sixth flagellomere of male Trigonomyia ananas sp. nov. Fig. 2. Head of male Trigonomyia ananas sp. nov. in frontal
view. Fig, 3, Gonostylus of male Trigonomyia tulipa sp. nov. in dorsal view. Fig. 4. Gonostylus of male Trigonomyia
cristata sp, nov. in dorsal view. Fig. 5. Wing of male Trigonomyia ananas sp. nov. Fig. 6. Genitalia of male Trigonomyia

ananas sp. nov. in dorsal view. Fig. 7. Last three flagellomeres of male Trigonomyia ananas sp. nov. Scale bars = 100
pm 1-4, 6, 7; 1 mm 5.

63

64

P. KOLESIK

rey

i pape

16:

eo

17:-——

Fig. 8. End of ovipositor of female Trigonomyia ananas sp. nov. in ventral view. Fig. 9. End of ovipositor of female Trigonomyia
cristata sp. noy. in ventral view. Fig. 10. End of ovipositor of female Trigonomyia tulipa sp. nov. in ventral view. Fig.
11. End of last tarsomere with claw, empodium and pulvillus of female Trigonomyia ananas sp. nov. in lateral view. Fig.
12, Sixth sternite of female Trigonomyia ananas sp. nov. Fig. 13. Sixth flagellomere of female Trigonomyia ananas sp.
nov. Fig. 14. Fifth tergite of female 7rigonomyia ananas sp. nov. Fig. 15, Last three flagellomeres of female Trigonomyia
ananas sp. nov. Fig. 16. Last two abdominal segments of larva of Trigonomyia ananas sp. nov. in dorsal view. Fig. 17.
Head and first two thoracic segments of larva of Trigonomyia ananas sp. nov. in ventral view. Scale bars = 100 xm 8-10,
13, 15-17; 50 pm 1; 500 wm 12, 14.

NEW CECIDOMYIIDAE FROM OLEARIA

ae) - } / 7 y,
we FA
fi \

22; —_- 23:24

Fig. 18, Trigonomyia tulipa sp. nov, - flower gall on Olearia axillaris. Fig. 19. Frons of pupa of Trigonomyia ananas sp.
nov. Fig. 20. Trigonomyia cristata sp. nov. - flower gall on Olearia axillaris. Fig. 21. Trigonomyia ananas sp. nov. - flower
gall on Olearia ramulosa, Fig. 22. Prothoracic spiracle of pupa of Trigonomyia ananas sp. noy. Fig. 23, Prothoracic spiracle
of pupa of Trigonomyia cristata sp. nov. Fig. 24, Prothoracic spiracle of pupa of Trigonomyia tulipa sp. nov. Scale bars

= 10 mm 18, 20, 21; SOO pm 19; 50 wm 22-24.

65

66 P. KOLESIK

Australia (34°54°S, 38°44 °E], 20.1%.1994, P. Kolesik,
reared from flower gall of Olearie ranulosa (Labill.)
Benth., sampled 17.1x,1994, 121294 [SAM].

Alloype> 9. same data, 21295 [SAM].

Puratypes (al) sampled with holotype): 1c [SAM),
Lo [ANIC], 2 9 9 [SAM]. 2 9 9 [ANIC], 2 pupal
skins [SAM], 2 pupal skins [ANIC], emerged
20, ix-8.x,1994; 2 larvae [SAM], | larva [ANIC].
Other material (all sampled with holotype):
39 Q[USNM], 4 pupae [SAM], 3 pupae [USNM],
emerged 5-8.x.1994: 4 larvae [SAM], 3. larvae
[USNM]}.

Description

Male (Figs 1-2, 5-7)

Colour: antennae grey. head black, ihorax brown,
abdomen with sclerotized parts black and non-
sclerouzed red (same jn other two species). Wing
length 3,1 mm (2,9-3.2), width I. mm (1.11.2).
Antenna total Jength 1.5 mm (1.5-1.6). Gonostylus 124
puny (121-127) long, 45 pan (43-49) wide, length of apical
claw of gonostylus I7 yan (16-18).
female (Figs 8, 11-15)

Colour as in male. Wing Jength 3.0 mim (2.9-3.1),
width 1.1 mm (10-11), Antenna total length J.4 mm
(1,3-1.5). Cereus 65 yim (60-68) long, 57 wn (57-58)
wide, setae 5-28 ym long,

Larva (Figs 16, 17)

Colour red (same in other two species), Total length
3.0 min (2.7-3.6). Head capsule: length 57 jan (50-62).
width 95 um (92-101), posterolateral apodemes 13 am
(11-16) long; antenna 17 pm long, 6 pm wide at base.
Length of setae; 2-3 jm in ventral papillae of thorax
and abdomen I-VII, 10-20 pm in remaining papillae.
Integumental spiculie 1-2 pm long.

Pupa (Figs 19, 22)

Colour: non-sclerolized parts of abdomen red.
remaining parts dark-brown (same in other two
species). Total length 3.3 mm (2.3-3.9). Length of setae
on cephalic papillae 361 ym (354-369). Prothoracic
spiracle 98 ym (93-103) long, trachea 70 wm (60-75)
long
Gall (Fig. 21)

Flower bud transformed into spherical, thin walled,
monothalamous rosette, 4-6 mm in diameter. When
fresh, gall wall green, malformed ligules violet, One
larva inside each gall. Pupation takes place within gall.
In the area surveyed, most shrubs were infested with
up to 200 galls per plant.

Etymology
The word “ananas’, a noun in apposition, is the

generic name of pineapple and refers to the
resemblance of the gall to a pineapple.

Trigenomyia cristata sp, nay,
(FIGS 4, 9, 20, 23)

Holorype; o , Beachport, South Australia [37°29'S,
140°00' BE], 85.1994, P. Kolesik, reared from flowee
gall of Olearia axillariy (DC,) F. Muell. ex Benth.,
sampled 6.x.1994, 121296 [SAM].

Allotype: 9, same data, 121297 [SAM].

Paratypes (all sampled with holotype): lor [SAM],
lo [ANIC], 29 9 [SAM].29 9 [ANIC], 3 pupal
skins [SAM], 2 pupal skins [ANIC], emerged
¥-19.x.1994; 1 larva [SAM].

Other material (all sampled with holotype). 29 9
[USNM]. 3 pupae [USNM], emerged 8-19.4.1994; |
larva [SAM], 3 larvae [USNM].

Descriprion

Mate (Fig. 4)

Wing length 3.4 mm (3.4-3.5), width 13 mm
(1.3-1.4). Antenna total length 1.7 mm (1,7-1.8),
Gonostylas 150 pm (145-157) long, 55 jam (50-58) wide,
length of apical claw of gonostylus 20 yam (19-22),

Female (Fig. 9)

Wing length 3.2 mm (3.1-3.4), width 1.2 mm
(1.11.2). Antenna total length 1.4 mom (1.3-1.6). Cercus
60 wm (55-63) Jong, 50 pm (47-52) wide, setae 17-45
pm long.

Larva

Total length 3,0 mm (2.4-3.4). Other measurements
within the range of 7 ananas.
Pupa (Fig. 23)

Total length 3.7 mrp (3.1-4.2). Length of setae on
cephalic papillae 386 ym (361-427). Prothoracic
spiracle 96 wm (86-107) long, trachea 57 jun (51-62)
long.

Gall (Fig. 20)

Flower bud transformed into monothalamous,.thick-
walled gall, 4-8 mm long. 3-6 mm wide, covered
entirely with numerous, densely-haired, malformed
ligules growing from proximal and. When fresh, botl
ligules and gal! wal! green in colour, One larva in each
gall. Pupation takes place within gall. At Beachport,
7 cristata was found with T, tulipa on the same shrubs
with up to 20 galls of each species per plant.
Etymology

The word “cristata” is L, for tufted, referring to the
general appearance of the gall.

Trigonomyia nilipa sp. nov.
(FIGS. 3, 10, 18, 24)

Holotype: &, Beachpert, South Australia [37°29'S,
140°O0' BE], 9.x.1994, P. Kolesik, reared fram flower
gall of Qlearia axillaris (DC.) F. Muell_ et Benth,
sampled 6.x.1994, 121298 [SAM].

NEW CECIDOMYTIDAE FROM OLEARIA Oo

Allotype: Q, same data, 121299 [SAM].

Paratypes (all sampled with holotype): 20" o" [SAM],
lot [ANIC], 29 9 [SAM], 29 9 [ANIC], 3 pupal
skins [SAM], 2 pupal skins [ANIC], emerged
9-17,x.1994; 2 pupae [SAM], 2 pupae | ANIC): 3 larvae
[SAM], 2 larvae [ANIC].

Other material (all sampled with holotype); 1?
{USNM], 4 pupal skins |SAMJ, 3 pupal skins
[USNM], i pupa [SAM], 3 pupae [USNM]. emerged
9-17,x,1994,

Description

Male (Fig. 3)

Wing length 3.3 mm (3.3-3.4), width L3 mm
(1,.2-1.3). Antenna total length 1.7 mm (16-18).
Gonostylus 121 jam (119-126) long, 50 jum (45-53) wide,
length of apical claw of gonostylus 13 am (12-14).

Female (Fig. 1)

Wing length 2.7 mm (2.4-3.2), width 10 mm
(0.7-1.1). Antenna total length 1.3 mm (1.1-1.5). Cercus
56 xm (52-61) long, 50 pin (43-55) wide, setae 620
pm long.

Larva

Total length of the only specimen 2.7 mm. Other
measurements within the range of 7 ananas.
Pupa (Fig, 24)

Total length 3.2 mm (2.8-3.4). Length of setae on
cephalic papillac 395 pm (364-455). Prothoracic
spiracle 51 um (48-53) long, trachea same length.
Gall (Fig. 18)

Flower bud transformed into smovoth,
monothalamous, thin-walled gall, 4-6 min in length,
3-4 mm in width, with tips of malformed ligules
sticking out al distal end. When fresh, colour purple.
One larva inside each gall. Pupation takes place within
gall.

Etymology

The word “tulips”, a noun in apposition, is the
peneric name of tulip and refers to the resemblance
of the gall to a tulip.

Key to species of Trigonomyia

1. Trachea reaching end of thoracic spiracle in pupa (Fig.
24). Apical claw of gonostylus diminutive, 1/4 of
gonostylus width (Fig. 3). From untufted galls of Olearta
axillaris (Fig. 18). cm an -T. wlipa
Traghes never reaching end of thoracic spiracle i in pupa
(Figs 22, 23). Apical claw of gonostylus large, more than
1/3 of gonostylus width (Figs 4, 6), From tufted ik wo
Olearia spp. (Figs 20, 21). ...-. 2
Longest setae on female cercus shorter than 2/3 of cereus
width (Fig. 8). From pineapple-shaped, glabrous, thin-

walled galls af Olearia ramulosa (Fig, 21)... ananus
Longest setae on female cercus longer than 2/3 of cereus
width (Fig. 9). From hairy, thick-walled galls of Olearia
axillaris (Fig. 20). - Hine Soot T. cristata

)

Acknowledgments

The Ministry of Environment and Planning South
Australia kindly gave permission to collect in the Black
Hill Conservation Park. Abid A. Munir State
Herbarium of South Australia Adelaide courteously
identified the host plant species, I am grateful to John
D, Gray Department of Horticulture, Viticulture and
Oenology University of Adelaide and Raymond J.
Gagné Systematic Entomology Laboratory USDA
Washington DC USA for their careful review of the
manuscript.

References

Cooke, D, A. (1986) Family Composilae (Asteraceae) pp.
1423-1486 Jn Jessop, J. P..and Toelken, H. R_ (Eds) “Flora
of South Australia, Part ILL (Polemaniaceae-Composituc )”
(South Australian Government Printing Division,
Adelaide).

GaGne, RJ. (1981) Fanuly Cecidomysicdae pp, 257-292 In
McAlpine, J. F.. Peterson, B. V., Shewell, G. E,, Teskey,
H. J, Vockeromh, J. R. & Wood, D. M. (Eds) “Manual
of Nearctic Diptera I" (Canadian. Government Publishing
Centre, Quebec),

______ (1994) “The Gall Midges of the Neotropical Region”
(Cornell University Press, Ithaca. New York).

, SOBHIAN, R, & Istporo. N, (Un press) A review al
the genus Psectrosema (Diptera: Cevidomyiidae), Ole
World pests of tamarix, and description of three new
species, /yrael J. Ent,

Koiesik, P, (1995) Asphondylia dudonaede, a new Species
of Cecidomytidae (Diptera) damaging leaves and branches
of hap-bush, Dodonaea viscosa (Sapindaceae) in Australia,
Trans. R. Soe. 8. Aust U9, 171-176.

FIRST FOSSIL RECORD OF THE HYLID FROG
LITORIA RANIFORMIS (KEFERSTEIN)

BRIEF COMMUNICATION

Summary

The Australian hylid frog Litoria raniformis (Keferstein) is a member of a group of
similar species known as the L. aurea complex’, and is one of the largest species in
Australia’ (snout to vent length up to 104 mm). The geographic range of the species
extends from South Australia through Victoria, the ACT and Tasmania to eastern
New South Wales’. The species also has been introduced into New Zealand and has
become established there’.

Transactions of the Royal Saciery of §. Aust. (1996), 120(2), 69.

BRIEF COMMUNICATION

FIRST FOSSIL RECORD OF THE HYLID FROG LITORIA RANIFORMIS (KEFERSTEIN)

The Australian hylid frog Literia raniformis (Ketferstein)
is a member of a group of similar species known as the L.
aurea complex!, and is one of the largest species in
Australia’ (snout to vent length up to 104 mm). The
geographic range of the species extends from South Australia
through Victoria, the ACT and ‘Tasmania (0 eastern New South
Wales’. The species also has been introduced into New
Zealand and has become established there?

it has been a source of surprise that such a large species
seemingly is absent from Holocene and Pleistocene sites in
south-eastern Australia where other extant sympatric species
have been found in abundance”.

Here We report the first specimens of L. ranifermis rom
the fossil record. The jlial descripuve terminology follows
Tyler®,

The largest and most complete specimen is a left ilium,
Jocuted in March 1995, trom muterial extracted at the ‘East
Low’ site at Henschke’s Cave, SA (Lat, 36°58'-06, Long.
140°45'-58). The specimen has been deposited in the
palacontological collection at the South Australian Museum
and registered as SAM P35305, The specimen has a length
of 28.0 mm which is less than the known maximum ilial length
of the species (35 mm). However, it is larger than other
Pleistocene frog ilia known {rom the area and ity identification
has been confirmed by comparison with extant specimens,
an example of which is shown in Fig. 1. SAM P35305 is
fragile and partly encrusted with matrix, hence the extant
specimen in more useful for identification purposes. A line
drawing of the sectional form of the ilium of this species has
been published elsewhere’,

"Tyler, M. J. & Davies, M. (1978) Aust. J. Zool. Suppl, 63,
1-47

“Tyler, M. J. (1978) “Amphibians of South Australia"
Handbooks Committee, Adelaide),

Tyler, M. J. (1994) “Australian frogs. A natural history”
(Reed, Melbourne).

Fig. | Pelvis of extant Litoria raniformis trom left, lateral
aspect, Length of (ium = 31 mm. (Photo: P. Kempster)

Salient features are (he poorly developed dorsal prominence
and dorsal protuberance, only slight elevation of the dorsal
acetabular expansion, a narrow and gently curved pre-
acetabular zone and a shallow longitudinal indentation upon
the literal surface of the ilial shafr-

We refer two other partial ilia from Henschke’s Cave (0 this
species: SAM P32249 and P3536.

The age of the deposit has been estimated to be from
35,0008 to 100,000" years. These pupers, provide informa-
tion on the depositional nalure and stratigraphic sequence of
the material.

We thank the Australian Research Council for funding
Michael Tyler's investigations of fossil frogs and Peter
Kempster for Figure |,

‘McCann, C. (1961) Tuatara, 8(3), 107-120.

*Tyler, M. J. (1977) Trans. R. Soc. S. Aust. 101(3), 85-89
oTyler, M. J, (1976) Ibid. W1(1) 3-14.

TTyler, M. J. (1986) Alcheringa 10. 401-402.

’Pledge, N.S. (1981) /bid. 105(1), 41-47.

*Barrie, D, J. (1990) Mem, Qld Mus. 28(1), 139-151.

MICHAEL J. TYLER, Department bf Zoology, University of Adelaide S. Aust. 5005, D, JOHN BARRIE, PO Box 227
Coonalpyn S. Aust. 5265 and RHYS W. WALKLEY, 48 Loan Street Black Rock Vic, 3193

THE TADPOLE OF LITORIA REVELATA INGRAM, CORBEN
AND HOSMER, 1982 (ANURA: HYLIDAE)

BRIEF COMMUNICATION

Summary

Litoria revelata Ingram, Corben & Hosmer, 1982 is a medium sized tree-frog that has
three disjunct populations; in northern Queensland (Atherton Tableland and the
Bellenden-Ker Range), mid-eastern Queensland (Eungella Plateau) and the extreme
corner of south eastern Queensland and northern NSW, Australia’. Herein we present
a description of the tadpole of L. revelata from the rainforest in the Eungella region in
mid-eastern Queensland. Habitat and life history notes are presented to assist
identification in the field but these are intended as a guide only and tadpoles could be
found in different habitats and months from those given.

Transactions of the Rayal Society of 8. Aust. (1996), 120(2), 71-73.

BRIEF COMMUNICATION

THE TADPOLE OF LITORIA REVELATA
(ANURA:

Litorta revelaia Ingram, Corben & Hosmer, 1982 is a
medium sized iree-frog that has three disjunct populations;
in northern Queensland (Atherton Tableland and the
Bellenden-Ker Range), mid-eastern Queensland (Eungella
Plateau) and the extreme comer of south eastern Queensland
and northern NSW, Australia!. Herein we present a descrip-
tion of the tadpole of L. revelafa from the rainforest m the
Eungella region in mid-eastern Queensland. Habitat and life
history noles are presented t6 assist identification in the field
but these ure intended as 4 guide only and tadpoles could be
found in different habitats and months from those given,

Tadpoles were collected in November and December Of
1993 at several stream sites near [he Eungella township,
approximately 70 km west of Mackay, central Queenslind,
Australia (Table |), A sample of larvae was preserved in 10%
formalin and others were reared to metamorphosis for
identification, Terminology fallows Altig’ and Hero’;
developmental stages follow Gosner*. Measurements were
taken using vernier callipers. Height of the caudal muscles
and fins was ineasured at mid-length of the tail, The drawings

Fig. 1. Tadpole of Literia revelata (QM J 59240; Gosner stage
35; TL 31.5 mm). Seale bar = 5 mm.

INGRAM, CORBEN AND HOSMER, 1982
HYLIDAK),

depict melanie patterns that persist in preserved specimens
(10% formahn). The colour descriptions should be treated
with caution as tadpole colour is often a function of water
clarity’. Drawings were made of two representative
specimens (Figs | and 2) placed in the Queensland Museum,
Brisbane (QM J 59239 and J 59240). The labial tooth-row
formula (LTRF) ix based on observations of all specimens
collected at Gosner* stages 25 through 45 (QM J 59241 and
4.59242; Tuble 1).

Description: Eyes lateral; eye diameter 14.5% of the body
length for stage 35 tadpoles and 14.7% for stage 40 tadpoles,
Nares dorsal, nearer to tip of snout than to anterior edge of
eye; narial margin without rim; spiracle paragyrinid (Fig. 2
C: located well below the horizontal longitudinal axis but not
on the midline so neither sinistral nor medioyentral is entirely
applicable®), unpigmented, opening directed posteriorly.

Vent tube dextral, attached to fin. Oral disc ventral. Single
row of large blunt, heavily pigmented marginal papillae with
wide anterior gap. Submarginal papillae present. Two rows
of labial teeth on aiterior labium with median gap in second
row: three rows of labial teeth on posterior labium with median
gap in first row: LTRF 2(2)/3 (1). Dorsal fin terminates at
tail-body junction. Both dorsal and ventral fins higher than
caudal musculature at midlength of tail. Tail+tip tapers uni-
formily to narrow point. These morphological features conform
lo the general characteristics for tadpoles.of the genus Litwria,

In life, body opaque, appearing “bluish” and heavily
Pigmented with lighter pigmentation around eyes; darkly
Pigmented supracranial patch (especially in larger tadpoles)
extending posteriorly over spinal cord (Fig. 28), distinct broad

Tante |. Dares, Localities and Museum Numbers for addinonal specimens examined in this stucly: Mt William {upper Catile
Cr.) Map 3655, MGR 666740), Mt David (upper Cattle Cr; Map 8655, MGR 678744).

Date Place Gosner Stage Body Length Total Length Qd Mus. Ne.
Collected (No) (mean) (Thean)
27.41.93 Cattle Cr, 25 4.6-4.1 248.3 J 59242
Mi William (7) (6.6) (15,3)
26 9.5 23.2
27 10.6 26.7
28 R.4 21.5
29 {1,3 27.4
a 12.0 29.8
33 WL.7 306
36 12.4 31.2
7 3.3 33,3
42 W.L-12.0 29.6-35.]
(2) (15) (32,3)
4s 10.5 .
44 W4-13.4 p
‘S) (12.3)
29.51.93 Cattle Cr. 25 88-106 2.1-25.2 J 5924)
Mt David (A) 9.7) (23.1)
27 14-11,9 26.1-27.6
(2) (11.6) (26.8)
3 12.0 33.0
37 14.6 43.1

‘i
PATTER Wy
ST
9-7

“a = yer ea
ce ieee
— wets areld

Fig. 2. Tadpole of Litoria revelata (QM J 59239; Gosner Stage 40; TL 40.8 mm), A, Oral disc. B. Dorsal view, C, Ventral
view. D. Dorso-lateral view. Seale bars = | mm (A), 5 mm (B.C,D).

vertical subdermal lines on dorsal side of each paris.
Horizontal band or patch from snout to eye (Fig, 21D),
Pigmentation often lighter during earlier stages (Fig. 1) than
at later stages (Fig. 2B-D). In ventral view intestinal maxs
visible, intestinal coils partially visible and obscured by heavy
pigmentation; branchial region semi-transparent. Tail
musculature un even shade of grey/brown with additional
melanophores concentrated dorsally (Fig. 2D). Dorsal and
ventral fins transparent, with even stippling of dark
melanophores, oflen outlining venation,

A tdpole at Stage 35 (Fig. 1) had the following
Measurenichts (hiv): total length 31,5, body length 17.7, body
width 6.5, body height 60, tail height 7,2, interorbital distance
5.6, internarial distance 2.1, eye-nary distance 2.0, A tadpole
at Stage 40 (Fig, 2) had the following measurements (mim):
total length 40,8, body length 14.3, body width #.2, body height
7A, tail height 8.3, interorbital distance 6,2, internarial distance
2.0, eye-naris distance. 2.6. Tadpoles vary in total length from
1.2 mm-al Stage 25 to 43,1 mm ut Stage 37 (Table |).

Diagnosis, Avihe sites studied, live tadpoles of L. revelara
can easily be confused with L. chloriy as both species oceur
in mid-water sections Of isolated streamside pools and they
have similar body shape and oral dise formula. Live tadpoles
of L. revelata have a bluish sheen covering the intestinal mass
and the intestinal coil is partially visible (Fig, 2C). In contrast,
1, chloris has a golden sheen covering the intestinal mass,
the intestinal coils are visible mid-ventrally and golden
chromatophores cover the heart.

In preservation, fidpoles of L, revelate have pigmentation
covering the intestinal mass making the intestinal coils only
partially visible. In contrast, 4. chloriy has a transparent
ventral surface and the intestinal coils are clearly Visible The
position of the spiracle, paragyrinid in L. revelata and sinistral
in LE. chloriy and (he dark pigmentation on the oral papillae
of L. revelata (with anly few scattered piyzments on the oral
papillae of, chloris) also distinguish these two species.
Interestingly, we onty know of one other Litoria sp. in
Australia with a paragyrinid spiracle (L. rubella, unpubl.).
Tadpoles of L. revelara were found in aympairy with tadpoles
of 1. chloris and Taudactylus ltemi. Adult frogs. of T
eungellensis and Mixaphyey fasciolatus were also observed
in adjacent streams,

‘Ingram, G. J., Corben C, J. & Hosmer, W. (1982) Mem,
Qd Mus. 20, 635-637,

*Altip, R. (1970) Herpetotogica 26, 180-207.

*Hero, J.-M. (1990) Amazoniana tl, 201-262

73

Habitat: Tadpoles of L. revelate were found in isolated
bedrock pools adjacent to fast-flowing rocky streams
surrounded by pristine rainforest, Each pool contained leaf
liter and algae and was between 1.5 and 2.5 m from the
stream, No fish were observed or captured by dip netting the
pools. Waler temperdures were noticeably higher in the pools
that in the adjacent stream (Table 2). Pool dimensions in
November were 100 em x 50 em x 10 cm deep for pool |

TABLE 2, Miler temperatures (°C) af pooly-and the adjacent
stream al [wer sttes,

Poo! | Pool 2 Adjacent Stream

Site Date

Cattle Cr iBin.94 14.9 16.0 14.2

MI Willian lia 94 lo% 1722 15.2
W094 19.7) OS 9
ii.xi§d 200 196 18.0)

Caltle Or IYix a 75 «160 13,5

Mi David 7x94 JL3 - 15.0
4.xi.94 200 7
2en94 22.5 22.8 17.9

and 200 em x 100 em. & 25 ci deep for pool 2. Tadpoles
were penerally observed in the midlwater and surface water
rather than the benthic layer of the water column and were
frequently Observed rising to the surface 10 gulp air

This research was partially funded hy the Australian Nature
Conservation Agency, the Queensland Department of
Environment and Heritage, the Wet Tropies Management
Authority and the Cooperative Research Centre for Tropical
Rainforest Ecology and Management, Research was carried
oul under a Qld Dept of Environment and Heritage “Permit
lo Take” no, TOOI77, We wish to thank Michael Cunningham
for his contribution to (his paper and Julie Martin who
prepared the illustrations and the voluftcers who assisted in
the LC.U, Eungella Frog, Search in Novernber/December,
1993. Ross Alford provided logistical support at J.C,U. and
Marion Anstis gave Valuable conyments on a dratt.

‘Gusner, K. L. (1960) Herpetologies 16, 183-190.
*Brage, A. N. (1957) Copeia 1957, 36-39,
‘Johnston, G. F, & Altig, R. (1986) Herp, Rey, 17, 36-37,

JEAN-MARC HERO Wet Tropics Management Authority CRC Tropical Rainforest Ecology and Management Dept of Zoology
James Cook University Townsville, Qld 48il, SHEREP. PICKLING & RICHARD RETALLICK Dept of Zoology James

Cook University Townsville, Qld 4811,

OBITUARY
NELLY HOOPER LUDBROOK, MBE, MA, PhD, DIC, FGS.
14.vi.1907 — 9.v.1995.

President of the Royal Society of South Australia Inc. 1961

Summary

An “obituary” is usually an account of a deceased person, but Nell Ludbrook deserves
more than just that. She really meant something to us so you must excuse me if I
depart from the kinds of ledger account statements that often follow the death of those
people who leave a significant mark on our community.

I first came across the name N. H. Ludbrook when I was a student at the University of
Adelaide in the late 1960s. While I was searching through the stacks in the Barr Smith
Library on some aspect of the evolution of interior deserts, her name appeared a
number of times in a paper dealing with geomorphology. The more | searched related
papers the more her name recurred. | must confess that, then, I didn’t know whether
N. H. Ludbrook was male or female, All I knew was that the name was referred to in
an array of papers dealing with stratigraphy, geological evolution, palaeontology,
palaeoclimate, ancient glaciations and the list went on. And it didn’t seem to matter
what part of the Phanerozoic either. I admit I thought that a person touching so many
aspects of geoscience had to be of great physical and scientific stature. It was not until
twenty odd years later when | actually met her that I found I was wrong on one count
but I was certainly not disappointed. What a marvellous person and an extra-ordinary
scientist | found her to be.

74

NELLY HOOPER LUDBROOK
MBE, MA, PhD, DIC, FGS.

At her office at the Department of Mines and Energy, Core Library, Glenside, 1985. MESA photo no. 34475

Transactions ef the Royal Society of 8. Aust. (1996), 120(2), 74-77.
OBITUARY

NELLY HOOPER LUDBROOK, MBE, MA, PhD, DIC, FGS

14.vy.1907 - 9y,1995,

President of the Royal Society of South Australia Inc. [961

An “obituary” is usually an account of a deceased
person, but Nell Ludbrook deserves more than just
that. She really hieant something to us so you Must
excuse me if | depart from the kinds of ledger aceount
statements that often follow the death of those people
who leave a significant mark on our community.

I first carne across the name N. H. Ludbrook when
| was 4 student at the University of Adelaide in the
late 1960s. While L was searching through the slacks
in the Barr Smith Library on some aspect of the
evolution of interior deserts, her name appeared a
number of limes in a paper dealing with
geomorphology. The more I searched related papers
the more her name recurred. | must confess that, then,
I didn't know whether N. H, Ludbrook was male or
female. All | knew was that the name was referred to
in an array of papers dealing with stratigraphy,
geological evolulton, palacontology, palaeoclimate,
ancient glaciations and the list went on, And it didn't
seem to matter what part of the Phanerozoic either.
T admit T thought that a person touching so many
aspects of geoscience had to be of great physical and
scientific stature. [lt was not until lwenty odd years later
when J actually met her that | found | was wrong on
one count but L was certainly not disappointed, What
a marvellous person and an extra-ordinary scientist |
found her to be,

Nell (never Nelly) was born Nelly Hooper Woods
at Yorketown, Yorke Peninsula on 14 June 1907, and
educated at Mount Barker High School in the Adelaide
Hills, During her undergraduate studies at The
University of Adelaide she became fascinated with Late
Tertiary Mollusca in the St Vintent Basin, a course
of study not easy at that time because palagontolopy
was not offered by the university. This faseination
broadened to the whole Cainozoic and continued
through her long career, Nell graduated as BA (1928)
and MA (1930), and was awarded the Tate Medal of
the University of Adelaide for a research paper on
molluses obtained from an Adelaide Plains borehole.
Even during her period of teaching at Mount Barker
High School, she still found time to extend her
knowledge of Mollusca.

Following her marriage in 1935, she and her
husband, Wallis Vereo Ludbrook,, moved to Canberra
where, undaunted, she continued her interest in Caino-

zoic¢ Mollusca. It was fortunate al this time that the
Commonwealth Palaeontologist function was moved
trom Melbourne to Cunberra, undoubtedly facilitating
her continuing interest in palacontology, While 11
Canberra, from 1942 to 1949, Nell worked as Assistant
Geologist in the Commonwealil Bureau of Mineral
Resources dealing with statistics of strategic minerals.

In 1950 she travelled to London. Here at Imperial
College und as a Visiting scientist at the British
Museum (Natural History) she continued to extend her
palacontological studies, Nell was awarded her PhD
in geology (1952) from the University of London and
the DIC in palaeontology for research on Pliovene
Mollusca from the St Vincent Basin, Out of this
research (leveloped an authorative chapter on fossil
scaphopuds in the first edition of the “Treatise of
Invertebrate Paleontology” (1960),

Sam ling Early Cretaceous rasa basin vation near
urreé, 1963, Photograph by B.G. Forbes. MESA photo
no. 20035,

76

Following the death of her husband and on returning
to Australia, Nell gained the position of Technical
Information Officer with the South Australian
Department of Mines in 1952. At this time
palaeontology was seen to have little economic value
- something more esoteric than having any practical
application. It was no mean feat, therefore, that Nell,
having been charged with the added responsibility of
demonstrating the application of micropulaeontology
in stratigraphy, succeeded way beyond expectations in
this role, She won the enormous respect of colleagues
around her and established biostratigraphy as an
important function of the Department. a role that
continues today.

During the heady days of early petroleum exploration
in the Cooper Basin, Nell was the key scientist in
determining the age and stratigraphy of samples from
deep wells drilled into unknown strata. Even the then
Premier of South Australia, Thomas Playford, waited
with great interest for Nell’s conclusions. Actually Nel!
admitted to me on one occasion that she did not really
know what the age of some rock samples was, so she
took a “stab”. As it turned out, later work, employing
far more sophisticated methods, showed her
determinations to be correct - such was the great range
of her knowledge,

In 1957 Nell was appointed Palaeontologist with the
Department of Mines, and later, Senior
Palacontologist, in which capacity she continued
biostraigraphic research until her “retirement” in 1967
During this time she developed an expertise in
foraminiferal biostratigraphy, essential to unravelling
the stratigraphy of largely buried strata in sedimentary
basins throughout the State and aiding in the search
for groundwater and petroleum. She travelled into
remote areas of the Eucla and Eromanga Basins with
tapping and drilling parties to undertake fossil
collecting and stratigraphic inyesugulions, offer
camping out in the open, Nell always insisted on seeing
the field relationships of the sediments and faunas she
worked on. It was through her field activities that
biostratigraphy became firmly recognised as an integral
part of geological mapping by the Department of
Mines. This work culminated in the publication of two
important monographs on the Murray Basin (1961) and
the Eromanga Basin (1966), still very much referred
to today as are the stratigraphic units she defined during
the course of her studics.

“Retirement” really meant the continuation of her
love of geology and especially for fossil Mollusca. She
worked as a consultant in palaeontology to the
Department of Mines and Energy until 1993, at which

N. H. Ludbrook und J. Spence examining Cainozoic sediments at North West Bend, along the River Murray. Photograph

by A. R. Crawford. MESA photo no. TOO2001.

time she had reached the age of 86, In addition to the
publication of a number of research papers during this
time she wrote the highly successful “Guide to the
Geology and Mineral Resources of South Australia”
(1980) and later the “Handbook of Quaternary Molluscs
of South Australia” (1985). As a demonstration of the
great respect and admiration that her colleagues from
all over the world had for her, a special honour volume
of papers dealing with stratigraphy and palaeontology
was published by the Department of Mines and Energy
in 1985. Until only a short time before her death in
1995 Nell was still researching a large volume on
Tertiary Mollusea.

Although the yast number of her publications (over
70 scientific papers and monographs) und Government
reports was known to-me, f only became aware of the
full extem of her extraordinary energies whilst ! was
researching material for the J995 Volume 2 of the
“Geology of South Australia”, During the course of
Tummaging through filing cabinets in the
Biostratigraphy Branch containing countless numbers
of her Report Books 1 came across a huge number of
unpublished Jetters and personal communications to
geologists in Companies, academia and povernment
carefully outlining the results of Work undertaken for
them, each almost of quality to be published notes in
their own right. We are now the custodians of Nell’s
journals, books and notebooks, donated by her in 1904
and now housed in the N. H. Ludbrook Memorial
Library at Mines and Energy South Australia,

Nell's interest in geology and the influence she had
on the science (and related. sciences for that matter)
extended far beyond the workplace. She was very active
as a member and office holder in the Geological
Society of Australia from its inception, She was the

77

founding Secretary of the South Australian. Division
(1953-56) and. Federal Secretary (1956-59), and a
Member of the Stratigraphic Nomenclature Committee,
in the early days of its operation, Nell was a driving
force in the preservation of key geological sites and
in the promotion of geological monuments. Nell was
elected Federal President of the Geological Society in
1968 and Honorary Member in 1976 - such was the
high respect that the geological community held for
her.

Her great energies extended into the affairs of the
Royal Society of South Australia. She was elected
President in 1961-62, awatded the Sir Joseph Verco
Medal in 1963, the highest honour fram the Society,
and was Editor of the Handbooks of the Flora and
Fauna of South Australia from 1967 to 1980. Nell
became an Honorary Assocrate of the South Australian
Museum in 1981. In recognition of her service to
science, in 1981 Nell Ludbrook was made Member of
the Most Excellent Order of the British Empire,

During her great devotion to research in
palaeontology and stratigraphy and her committment
to the affairs of scientific societies Nel! still found time
to guide and advise colleagues in many aspects. of
geoscience. She travelled widely throughout the world
pursuing her love of geology - into many places where
European women were rarely seen. Nell had the rare
gift of being able to devote herself to this pursuit and
yet still maintain an enormous interest in the cultural
and musical life of Adelaide and the world at large.
She loved entertaining at her home at Toorak Gardens
- many an overseas visitor was delighted with hes
hospitality.

It was an honour and a pleasure for all of us to have
known Nell Ludbrook,

NEVILLE F. ALLEY

OBITUARY
STANLEY JOE EDMONDS, BA, BSc, MSc, PhD, Dip Ed.
13.11.1909 — 16.vii.1995.

President of the Royal Society of South Australia Inc. 1965

Summary

Stan Edmonds died quietly in his sleep on 16 July 1995 aged 86. He is sadly missed
by his many friends from all walks of life who miss his sense of humour, joie de vivre
and scholarship.

His working life fell roughly into three periods each about twenty years’ duration. He
was a school teacher at Adelaide High School from 1931 to 1952, he then taught and
conducted research in the Zoology Department of the University of Adelaide from
1952 to 1974 and finally, as an Honorary Associate at the South Australian Museum,
he continued his research from 1972 to 1995.

mn

/
ee
a.

oe

_
8

STANLEY JOE EDMONDS
BA, BSc, MSc, PhD, Dip Ed.
Photograph courtesy of the SA Museum

Transactions uf the Royal Society of S. Aust. (996), 120(2). 78-82.
OBITUARY

STANLEY JOE EDMONDS, BA, BSc. MSc, PhD, Dip Ed.
13.11.1909 - 16.vii. 995.

President of the Royal Society of South Australia Ine: 1965

Stan Edinonds died quietly in his sleep on 16 July
1995 aged &6 He is sadly missed by his many friends
from all walks of life whe miss his sense of humour,
joie de vivre and scholarship.

His working life fell roughly into three periods cach
of about iwenty years’ duranon. He wax a schoo]
teacher at Adelaide High School from i931 fo 1952,
he then taught and conducted research in the Zoology
Department of the University of Adelaide from 1952
to 1974 and finally, as an Honorary Associate at the
South Australian Museum, he continued his research
from 1972 to 995,

Stanley Joe Edmonds was born in Adelaide (South
Australia) on 13) February. $909. He attended the
Thebarton Primary School fram 1915-1922 and the
Woodville District High School from 1923-1925,
obtaining his Interrnediate Certificate in 1924 and his
Leaving Certificate the following year In 1926 he
joined the Lands and Survey Department of the South
Australian Public Service with the imenton of
becoming a suryeyor. During this tume he studied
scieqve part-time as a private student at the University
of Adelaide doing Mathematics | during the day and
Chemistry Tand Physics 1 al night-

In 1927 he joined the Education Department and
entered the Adelaide Teachers College. He praduated
in 1929 with a BSc majoring in Inorganic and Organic
Chemistry. and began his teaching career in 1936,
teaching for six months at Woodville High School.

In 1931 he began his impressive twenty-year
association with Adelaide High School, ullimately
hecoming a Spécial Senior Master in Chemistry and
General Science and teaching Leaving Honours
Chemistry from 1945 to 1951. During ths time, he
obtained three further degrees - a BA in 1935 in which
he majored in Latinand English, a First Class Honours
in Zoology in 1945 (after completing Zoology 1, UW and
Il in 1941. 1945 and 1944 respectively) and an MSe
in 1947,

However, it was his broad interests and sense of

humour combined with his great teaching ability thar
endeared him to his students. He was interested in
sport, pafticularly tennis and hockey at which he
excelled, acting, singing and the arts.

During his last few years at Adelaide Aigh Schoo!
Stan became interested in Zoological research and
began a senes of collaborative studies with T_ Harvey
Johnston. the Foundation Professor of Zoology at
Adelaide University and a noted Parasitologist, These
studies on Australid4ng Acanthocephala (spiny ‘headed
worms. parasitic in the alimentary canals of various
fish, birds and mammals) were first published in 1947
and continued for several years after Professor
Johnston's death in 951. During this period Stan
widened his interests to include free-living marine
organisms and in 194% had a paper published on “The
common species of animals and ther distribution on
an intertidal platform at Pennington Bay, Kangaros
Island”. These counting interests. in Zoology led in
1952 to his resignation fram the Education Department
fo lake up an appointment ws a lecturer in the Zoology
Department, University of Adelaide under the newly-
appointed Protessor of Zoology, W. P. Rogers, an
authority on the physiolagy and biochemistry of
parasitic nematodes.

It was at this Lime that Chad the pleasure of meeting
Stan as [ had been appointed ta the Zoology
Department as a Demonstrator earlier chat year. One
always associated him with laughter or at least a smile
His miming of sewing his fingers together and then
threading the needle through various parts of his arm
so that the whole would be moved mechanically by
pulling on the thread, was always demanded of him
al departmental parties and was always accompanied
by gasps of horrar from the faint-hearted and much
amusement from the initiated.

Later, in his chapter on Zoology in “Ideas and
Endeavours. The Natural Sctences in South Australia”
Stan described What 4 busy time his early years in the
Zoology Department had been, as student numbers
were increasing rapidly and Rogers was building the
Zoology Department. Needless to say, in this chapter,
Stan gave himself scarcely any mention.

In addition to broadening his taxonomic interests,
describing new species of Australian marine
invertebrates, notably sipunculans and echiurans, Stan
began ta conduct a range of physiological and
biochernical experiments on them. These physiological
experimenis were extended to the parasitic acantho~

80

cephalans and to the species Moniliformis dubius in
particular. This species was maintained in Use
laboratory in cockroaches. the mtermediate host, and
rats, the primary best. The distinctive rasile and odour
of Stan’s experimental cockroaches, as one entered the
constant temperature room in which they were boused -
are sounds and smells not casily forgotten! The
nutrition and egg laying of these animals were studied
and teported upon and, together with B. R. Dixon, a
paper was published in Marure on he uptake of small
particles through the body wall of M. deine:

Around this time Stan collabunited with H. BS
Womersley in what was the first significant paper on
the intertidal ecology of South Australia. It was also
the first paper published on this topic in Australia that
dealt in equal detail with both flora and fauna.
Furthermore, it dealt with the relatively sheltered
coastline of South Australia which, with its gulfs amd
bays and Kangaroo Island, differed from the more
exposed coastline of the Eastern States. In die light of
their work Womersley amd Edmonds were able to
supply evidence, previously unavailable, for the
biogeographical nomenclature of the southern
Australian coastline.

In addition to bis researches on intertidal coology.
Stan also worked with Manan Specht on ecological
siudies of heathland in the Keith region of South
Australia, This work involved monthly visits over a
period of three years between 1952 and 1954 and
resalied in the wocumulation of a vast amount of
information that permitted judgements to be made on
the faunal rhythms of heathland in South Australia.

Tn 1958 Stan's researches on sipunculans resulted in
his being awarded a PhD.

Stan Edmonds’ work of the Public Examinations
Board, a member from (96) to 1974, Chief Examiner
in Biology for ten years and deputy Chairman from
1973-1974, was a measure of the regard In which his
waching expenence was held.

Stan was to continue to undertake research on the
sipunctilans and the somewhat similar echiurans for
many more years, In 1972 he co-authored a book with
his late friend Dr A. C. Stephen of the Edinburgh
Museum entitled “The Phyla Sipuncula and Echiuni™

At the time of Dr Stepher’s death, miuch remained to
be done and it was recognized that Stam was the only
person who had the scholarship and energy to complete
this task, His share of this important contribution to
marine studies was a large one, bringing information
up to date and checking descriptions, records and
translations with original specimens and data, He
arranged species into genera, provided keys for
identification and was alone responsible for the sixty
full page illustrations. Some 320 species of sipunculans
and 130 species of echiurans had been described at the
time this 527 page book was published by the Tnistees
af the British Museum (Natural History) London. Is
was. the first systematic monograph of the two phyla
to be published this century and is likely to remain
the standard reference Work for many years ta come,

Stan retired from the University of Adelaide in 1974,
having been made a Reader in Zoology in 1973, Re
became an Honorary Associate of the South Australian
Museum and over the next twenty years published a
further thirty papers including several chapters in
books. He was a strong supporter of the Royal Society
of South Australia Inc., occupying the positions of
Council member, Secretary. Vice President and
becoming President in 1965. In 1982 he was awarded
the Society’s Sir Joseph Vereo Medal for his
distinguished scientitic researches.

In conclusion T quote from C. M. Ward MA, a Latin
teacher and scholar of high repute and, wl the Lime,
Acting Principal of Adelaide High School who wrote
on. 17 September, I948 the following words. “Mr
Edmonds has a lively, genial personality, 2 good sense
of humour and a resourceful cultured mind. He is of
strong, independent character bul always fiemdly and
unassuming. His honesty and integrity are
unjuestioned™ A most fitting tabute to a much liked
and respected personality who maintained these traits
throughout his life,

Stan Edmonds is survived by his wife Barbara (née
Fy) and a daughter Elizabeth.

ALAN F. BIRD

Ribliography

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48 Australian Acai No. 7. Trans. R. Soe, §, Aust.
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1948 The commoner species of animals and their distribution
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South Australia. [bid. 167-177.
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1953 Acanthow a from Auckland and Campbell Islands.
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1956 Chlonnities of coastal waters in South Australia, Trans
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fresh, Res. 8, 55-63.

1957 The catabolism of nitrogen compounds in Beviclroseorruary
ivineducewe Edmonds (Sypunculoides). Ibid |31-135,
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1929-1931 Rep, B 6, 92-98.

1957 Austrtlian Acanthocephala No 10. Frans. R. Sac. 8.
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58 A Seueral account of the intertidal eculogy of South
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60 Some Australian echiurnids (Echiuroideay. Trey, #.
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[960 Sipunculids from New Zealand und Chathan tstand.

_N.Z. Depi Sci, & Indust, Res, Gull, 139, 154-167.

96) On Sipunewdics aeneus Baird (Sipunculaides), Ann, May,
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1962 Some notes on Ue abundance, environment anll nutrition
of Sipuncubes rudus L. at Morgat, Brittany. Cul, Brot. pir.
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(865° Wwe new echiunoids (Fehiurmidea) fron) Australia. Trans:
R, Soe, §. Aust, $7, 243-247.

1064 Australian Acunthocephaia No. 11. Abid, 88, 4).95,

1965 Sipunculoictes of the Russ Sea, .N. 2. Oupt. Sei. db deedieit
Res, Bull, W7, 27-34

4965 Some experiments om the nutrition of Munihifurmes
dulius Meyer (Acanthocephala), Parasitalapy $5, 337-344.

966 Siphonaome hawaiense 4 new sipunculoid tea Hawaii
(Sipunculuidea) Fre. Sci. 20, 386-388,

Wet Uptake of smal) particles ty Moanilifarnis eltibiies
(Acanitiocephala). Widture (London) 209, 99 (Will RR
Tiana.

19%) Hatching ot the eggs af Moanilyiirmis dabins Meyer.
Exptl. Purasizol, 1, 216-224.

1966 Sipuneuliades and Echivnvides, Port Phillyp Survey
1957.9. Mem. Na. Muy. Melb, 27, 175-178.

1987 Fardcunthorhynchws gilixiasue, a new genus aad species
of Acamhocephala trom fish. Australian Acanthocephala
Na, 12. Trams, &. Sae. S. Aust. 91, 41-43

968 Distribution of selected groups of murine invertebrates
in waters south of 3$°S lar. Folic li, Antarctic Map Folio
Senes, pp 23-24.

Sipuncula and Evhiura. Amer. Getigrah. Soe.

1969 Moastrements of the osmotic pressure in the habnat of
Folymorphus mitutes (Acanthocephala) in ihe intestine of
the domestic duck. J) Exp, Bjo. 50, 69-77. (Wilh BD. W.
T. Crompton)

197) Some sipunculans and echiurans chiefly from Guam
(Sipuncula and Echiura), Mieronesiva 7, 137-151.

1971 Australian Acanthocephala No. 13. Three tew species.
Trans, Rv Soe. & Aust. 97, 19-21.

1972 Marine invertebrates trom Adelie Land, collected by die
th and 15th French Aatarclic Expeditions. 5. Sipunculu.
Tethys, Supp. 4, 83-26.

1972 "The phyla Sipuncula and Echiura® (Trustees British
Museum (Nat. Hist.), London). (With A. C_ Stephen),

1973 Australian Acanthocephala Nu, 14, On iwo species of
OH el ee one new. Trans. R. Soc. §. Auyt. 97,

oy

KI

1973.4 new species and genus of earthworm (Megascolevidae,
Oligochaeta) from South Australia, Mbid.23-27. (With Tk
G. M, Jamieson).

1974 A new spocies of Sipuncula (4xpidosiphon exienus)
belonging to the imterstilial fauna of marine beaches,
collected by Mr L.. Bolosansenu during the second Caby-
Romanian Biospeleological Expedition to Cuba, 1973. bat
J. Speleel, 6, 187-192.

76 BraivaGnin in sipuinculans. Proc. [nuermational Symp.
Biol. Sipuncwla and Echiura, Kotar (1970) 2, 3-9,

1976 Thive sipanculan species (two new) from New Zealand.
N.Z, J Mar. Freshus Res. 10, 217-224

1979 Intertidal invertebrates pp. 155-167 Jn Tyler, M, 1.
TWidale, © R_ & Ling, J. K, (Fats) "Natural History of
Kangaroo tsland” (R, Soc, & Aust, Ino, Adelawe) (With
1, M. Thomas),

1979 Inwntidal ecolagy of marine orgailisms pp. 1f7-177. Janet
(With H., B.S. Womersiey)

1980 A revision of the sytemuatics of Australian sipunculans
Ree. S, Aust. Mas, WA 1-74-

1981 Dark Island heathland, ‘South Australia: faunsi rhythms
pp. (5-27 da Specht, R. L. (Ed_) “Reosysiems of the World,
9B. Heathlands and Related Shrublands, Analytical Studies”
(Elsevier, Amsterdam), (With M. M. Speech).

1992 Sipunculans pp, 299-31) de Shepherd, S. A_ & Thomas,
LM. (Eds) “Marine Invertebrates of Southern Austrtlia
Part 1. Handbook af Flora and Fauna of South Australia”
(Govt Printer, Adefaides,

1989 Echiurans pp 31231 thre.

1982 Ausiralian Acanthocephala No, 15. Four species. Trans,
R. Sac, S. Aust. 106, 71-76.

1982 Echiura pp, 65-66 Jn “Synopsis and Classiliention of
Living Organisms” (McGraw-Hill, New York)

1982 A sipunculan. répered no be “Tyterstitial” tear Lhe
Notherlands Antilles. Bijdrag, Dierkunde 52, 225 230,
1984 Phylum Sipuncula snd Phylum Gebiurs p- Ol-62 In
Mather, P, d& Bennett. 1. (Eds) “A Coral Rect Handbook”

(Great Barrier Roct Soviety. Brisbane)

8S A oew spcents of Pian eofusema (Sipyncula) [rom
Australia. Traks. & Stu SN 4net, TOS 43.dd

WSS A new ectiuran Sluierina daikourae (Belliurs:
Honellidae) from) New Zealknd und & nete on New Fenland
echiurans. WZ. Mar Fresh! Res 19) 6D L60d,

3986 Sipuncula p. 209 Jn Bolosuneanu, 1., (Eit,) “Stypofauns
Mund 96° (Brill & Buckhuys. Letden),

1986 A checklist of helminths from Australian birds. er:
S Aust. Mix. 19, 279-325. (With FM. Mawson & L, M,
Angel)

J9R6 A note ori pome sipHoculans fom rhe Norhem Territaacy.
Australia, The Beagle 3, 7-0

1986 Zoolopy pp. [62-212 Jn Twidale, ©, RB, Tyler, M,
& Davies, M. (Eds) “Ideas and Endeayours, The Natural
Sciences in South Australiv” (R. Soc S, Aust, Tnec.,
Adelaide),

1987 The sipunculun fauna (Sipuncula) ol Wextern Australia.
Ree, Wo Aust. Mus, 13, 215-224-

987 A note on the uccurrence of Balboxena capitan
(Linstow, [880) (Acanthocephala) fran) a false Killer whale
stranded on the coast of Western Australia, Ibid, 317-318,

1987 Obituary: 1. M, Thomas. Rec. 8, Aust Mus, 21, 61-63

1987 Echiurans [rom Australis. /bid. (19-130,

1987 Sipunculans and Echiurans pp. (85-212 Ja Devaney, D-
M. & Eldndge, LG, (Rds) “Reef and Shore Fauna of
Hawaii. Section 3" (Bishop Museum Press, Honolulu),

WBE dustraliformis semoni (Linstow, 189%) no. gen., 9. conib.
(Acanthocephala: Moniliformidae) froni marsupials ot
Australia and New Guitwi. J. Perasital. 75, 215-217. (With
G. D. Schmidt).

1989 A list of Australian Acanthocephala and their hosts. Rec,
3. Aust, Mus. 23, 127-133.

82

1991 Sipunculans and echiurans from the Phillipines and New
Caledonia (Estase 2, Musorstom 3 & 4). Mem. Mus. Nat.
Hist. Nat. (A) 151, 83-90.

1992 A new species of Acanthocephala from the green back
flounder, Rhombosolea tapirina Giinther. 1862. Trans. R.
Soc. S. Aust. 116, 35-40. (With L. R. Smales.).

1992 A note on Phascolosoma turnerae (Rice) (Sipuncula).
Ibid. 151.

1993 Sipuncula and Echiura pp. 94-96 In Mather, P. &
Bennett, I. (Eds) “A Coral Reef Handbook” (Australian
Coral Reef Society, Brisbane).

1993 Index to the Transactions of the Royal Society of South
Australia, Vols 102-113 (1978-1989). (With L. W. Parkin).

VOL. 120, PARTS 3 & 4
29 NOVEMBER, 1996

Transactions of the
Royal Society of South
Australia

Incorporated

Contents.

Anstis, M & Littlejohn, M. J. The breeding biology of Litoria subglandulosa

Dyson, I. A.

Dyson, I. A.

and L. citropa (Anura: Hylidae), and a re-evaluation of their
geographic distribution - - - - - - - - ~
Stratigraphy of the Neoproterozoic Aruhna and Depot Springs
subgroups, Adelaide geosyncline - - - - - - -
Stratigraphy of the Neoproterozoic Tent Hill Formation and
Simmens quartzite at South Tent Hill on the Stuart Shelf,
South Australia - - - - - - - - - -

Taylor, G. S., Austin, A. D. & Davies, K. A. Biology of the eucalypt

Bird, A. F.

Kolesik, P.

Nicholas, W. L.

Smales, L. R.

Brief Communications:
Baker, G. H.

O’Callaghan, M.

gall-forming fly, Fergusonina flavicornis Malloch (Diptera:
Fergusoninidae) and its associated hymenopterans in South
Australia, with a description of a new species of Bracon
(Hymenoptera: Braconidae) - - - - - - - -
Studies on the soil-inhabiting tardigrade, Macrobiotus cf.
pseudohufelandi, from South Australia - - - - - -
Rhopalomyia goodeniae, a new species of Cecidomyiidae
(Diptera) damaging Goodenia lunata (Goodeniaceae) in inland
Australia - - - - - - - - - - -
Robustnema fosteri sp. nov., gen. nov. (Xyalidae, Monhysterida,
Nematoda), a common nematode of mangrove mudflats in Australia
A redescription of Aspersentis zanchlorhynchi (Johnston & Best,
1937) comb. nov. (Heteracanthocephalidae: Acanthocephala) - -

Seasonal activity of the earthworm, Gemascolex lateralis
(Megascolecidae), in a Eucalyptus woodland in South Australia —-
G. & Beveridge, I. Gastro-intestinal parasites of feral cats in the
Northern Territory - - - - - - - - -

Terrace, T. E. & Baker, G. H. Predation of earthworms by the land planarian,

Australoplana sanguinea (Moseley) var. alba (Dendy) sensu Jones,
1981 (Tricladida: Geoplanidae) - - - - - -

Tyler, M. J. & Williams, C. R. Mass frog mortality at two jeeniriee’ in South Australia

PUBLISHED AND SOLD AT THE SOCIETY’S ROOMS
SOUTH AUSTRALIAN MUSEUM, NORTH TERRACE, ADELAIDE, S.A. 5000

83

101

117

177
179

TRANSACTIONS OF THE

ROYAL SOCIETY
OF SOUTH AUSTRALIA

INCORPORATED

VOL. 120, PART 3

THE BREEDING BIOLOGY OF LITORIA SUBGLANDULOSA AND
L. CITROPA (ANURA: HYLIDAE), AND A RE-EVALUATION OF
THEIR GEOGRAPHIC DISTRIBUTION

By MARION ANSTIS* & MuRRAY J. LITTLEJOHNT

Summary

Anstis, M. & Littlejohn, M. J. (1996) The breeding biology of Litoria subglandulosa
and L. citropa (Anura: Hylidae), and a re-evaluation of their geographic distribution.
Trans. R. Soc. S. Aust. 120(3), 83-99, 29 November, 1996.

The known range of Litoria subglandulosa is extended and that of L. citropa 1s
revised. Population trends observed at the type locality during the 1960s-70s and
1990s are compared.

The advertisement call, adult colouration in life, behaviour and embryological
development of L. subglandulosa are described and compared with those of L.
citropa. The single egg mass of L. subglandulosa shows adaptation to the lotic
environment, being compact and strongly adherent. The embryos and larval stages of
the two species are very similar in shape and colour in life from stage 17 onwards, but
are readily distinguishable by mouthparts. Comparative notes on larval behaviour are
given.

Key Words: Litoria subglandulosa, Litoria citropa, distribution, population trends,
advertisement calls, oviposition, embryology, larval behaviour.

Transactions af the Raval Soctely of S Musl (1996), T2004), 83-09,

THE BREEDING BIOLOGY OF LITORIA SUBGLANDULOSA AND L. CITROPA
(ANURA: HYLIDAE), AND A RE-EVALUATION OF THEIR GEOGRAPHIC
DISTRIBUTION

by Marion Ansiis® & MuUkRAY J. LIvTLeionn:

Summary

Amsitis, Mok Lirrirtonis, MEd, (1996) The breeding biology of Lirarig subylandilose and 2. cirrapa (Anuta:
Hylidae), wand a re-evaluation of their geographic distibunion. Trams. Ro See. 8) Aust 120041, 83-99, 29

Novertber 199%.

The keown range of Citerie suigtandalese is extended and that ot Lh. ciapa is revised, Population trends
observed ait the type lnedlicy during the 1960870s and 1890s. are Cormparcd.

The advertisement call, adull colournmion in lite. behaviour aim embryologieal development af 1,
subglandulose are described and compared with those of Lo crrmpa. The single ee mass al Lo subularuticlaser
shows adaptation to ure lotic environment being compiict und strongly udherent The embryos and larval stages
M the Wwe species sve very simile in shape and colour in life from sue 17 onwards, but are readily
distinguishable by mouthparts. Comparative dates on larval behaviotic are given.

Riv Warps: Zifern subctandulose, Lilorte ciropa, distribution, popubilion tends, advertisenient calls,

oviposition, embeyolopy, larval behaviour,

Introduction

Litoria Subglandulasi wis. deseribed as Lateria
glandulosa “Vyler & Austis, 1975 bul renamed
becuse Of primary homonymy (Tyler & Anstis,
1983), A member of the L. efmopa species group
(Tyler & Davies 1978), Lb. subglandilosa was
previously known only From the Quechshina/NS Ww
border south to the New England ranges of northern
NSW (Tyler & Anstis 975) The type description
included wdeseripuon of the kirvae. but no dita were
available on aviposition, embryological development,
larval behaviour or the advertisement call.

The species was found 1600 km south of as
previous known distribution inthe mid-rerih coastal
Rinwes wd Barrington Tops region by one of ts
(M.A,). in 1977) lis presence there and the absence
oF 2 eitrape, prompted a re-examination of the
distobution of both species. In addition, observalions
on oviposition, the morphology of embryos, larvae
and adults aid a comparison of the advertisement
calls of Lo cinwpe and L. subulanedilose were made
and are reported here.

Materials and Methods

Linta sibglandulosa

Adult specemens exantined: Australian) Museun
(AM) R1I7577, 35525, 42934-35. 50163, S1096-7.
Sti0d, 5173549. Point Lookout, R34458 - 14k
Fast of Ebor: R36724 - Oakey Creek neur Ebor,

~ Id Witeview Rul Beton Heights NSW 2082,
Deirnientel Zoalagy, Liniversdity at Melbourne
Parkville View 40529

36975 - Guy Fawkes River, Ebor, R7EIO9-T L114 -
Back Creek (Barwick River) pear Point Lookout
R37017 - Skin Sof Waleha >: R39056 - SOkim E of
Glen Innes (Gibraltar Range), R52031 - Sandys.
Creek, Dorvigo; RALI7S-80 — Styx River, Point
Lookout; R76S19 - Gloucester Tops: R31683
Upper Allyn River, Barcingion Tops: RLO4932
Ellenborough River, Bulua State Forest, NSW.

Litorea crtrape
Adult specimeny examined, Australian Museum
R7560_ Orbost: 7562. Aberfeldy, Vie 19237, 18234.
18236. 18245 Stunwell Tops: 79436, Stanwell Park;
24500-24505, 27590, Fauleanbridge; 45858,
Thitlnvere Lakes; 3)68S, 7112. 78927 Telensburgh;
45424, Tianjara Falls; 5188, Megialong Valley: 7110,
Hizelbrook: S008. Blackheath, 69034. Bell,
Kurrngjong Rd. 76625. 16 km N of Lithgow; 8459,
Pennant Hills; 14495, Cols Vale: 79100, 76623,
Culoul Range N of Cole Hts. 4261. Bundanoon:
TISYS, 2h ke N of Moss Vile; 15462, Gosford:
7T8204-26, 78698, Kuringai-Chase: 60425. Nadgee
Reserve; 79439, Cialston Gorge: 7563, Manly, NSW.
Three adults etted as 2. efirepa by ‘Tyler & Austis
(1975) from Barrington Tops localities: Dept Zool,
Univ. Melhourne (MUZD) 1792/64 - Upper Allyn
wid MUZD 1690-91/63 - Wombul Creek, were re-
exumined because oFupparent overlap in range with
the Barrington Tops localities for 2. swhehiedtlose.
These specimens have since been registered by the
Nuavonal Museum of Victoria (NMV) as 132666
(Upper Allyn Rivers and 193266d-65 (Wombat
Crock). Similarly, two specimens (NMV DO709-10),
vied by Copland (1957) as. avreape from neat
Crafton. noeth-castern NSW. were exuniied,

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BREEDING BIOLOGY OF L. SUBCLANDULOSA & L- CTTROEA BY

Ovipasiian and embryos

Observations on three cuplive breeding pairs ol
mivh species, collected by MLA, tre Summarised in
Tables 2and 3, la each case, a calline male was [first
collected at night, then a grayid female was found
during daylight the next day. in the same vicinity as
the male. The pairs were cach placed in a large
inflated plastic bag containing stream water, a flat
tock ind aguatic yegelution. The bay was covered
with opaque material for the duration of ampleaus,

Litera subeleandulase

Styges [-25 (Gosner, }960) were stuidica [ror
lhree sepirate ege misses. One from (he type locality
(luculity 10. Table 1), and the others from jae new
focalities 3 and 4h. Hereatter, numbered localities
will refer to Table 1 (unless otherwise stited),
Purther saiples frompece nusses found th (he stecary
al locality & were mauntuined until sige 25 to
confirm identity. by Dr A. While of the National
Parks & Wildly Service, NSW (NP& WS). Finbryas
and Jarvue were held ip dishes (40 em diam.)
containing streaiy Water, rocks, sedients aid
aquatie vexetation, und miintamed at 14-20C
Qoeulity 9), amd T5'-24° (localities 3 and 4h).

The eggs nuiss from tocality 4b tare on 7aal so
(hible 2). was submerged wathin a metal tea Strainer
in the cool, flowing water ol the stream for the titial
(wo days of development, but both the eg misses
from localities 4 and 4h were maintained at higher
lemperatiires of up to 24°C uway from the stream
from the third day afier deposition, Embryenie
developient Was observed under a Wild M5
MEPeOSCOPIE MILroscHpe,

Litorta cubkeype

Staves |» 25 were studicd from two egg masses
fron) Darkes Forest and one fron Ourimbsh
Coc#liies 15 & 16, Table 3), Samples of eges found
scullered over the substrate inthe stream were raised
lo stage 25 10 canfirm identity, Adults in breeding
condition were pliced in an inflated plistic bag
eovered with opaque mylecal during amplexus and
the resulting embryos maintained al 16°-27°C,

Larvae

‘Ladpales were measured (to 0.) mm) with vernier
cullipers and un ocular aicrameter attached to the
niicroscope. They were anwesthetised in Chlorbutol
solution before preservation in 3% formalin, The
Stuping system ots that of Gusner (1960),
Abbreviations for larval measurements shown in
Table 6, follow Anstis (1976); TL = total length, BL
= body length. BD = maximum body depth, TD =
maxinounm til depth, TM = inl musculature depth
(measuced in line with TD), 10 = interorhital span.
IN =internarial Spun. EN = the distance between eye

und maris and MW = maximum mouth width,
Hlustritions Were tmde using a drawing cube
ultdched to the microscope. Preserved and living
larvae OF L, sibelumedidose fro sites | - 9 were
examined for compartson witht those from (he type
locality and measurements are given in ‘Table 6.
Feeding and swinnnng hebavicur of several larvae
of both species was observed if captivity and i the’
nalwral dale eayiroament,

AdVvertisernrent calls

The cally of L. wubglaridulosa were recorded at a
tape speed of 4. 76 chi sec! Using a Sony TO-DSPRO
portable cassetie recorder with a Uher M516
microphone and a Grampian parabelie reflectar,
Calls of 1. crrrapa were recorded with a Nagra 4.2
open-rec! Jape recorder af a tape speed of | ern see |
anda Beyer M-X& cardio dynamic microphone.

Por Lo snbglandulesa, the lupe cusselie was
replayed on uw Nukamich) Dragon tape deck. and for
L. cite. the open-reel tape wis replayed on enther
a Revs Be 77 ora Sony PCS [0-2 rape recordey. The
calls Were dmilysed on a Kay tlemetnes Digital
Sonu-Graph. Model DSP-S500, Addylional analyses
ol waveforms were made hy way pf a Souned- Blaster
Lo card (Creative Technology) installed ih un IBM
PC-compatible desktop camputer, and osing the
Wave Studio (Creative Technology ) ail Speetra Plus
Protessional. Relewse 3,0 (Pioneer Hill) sotlware,
Both systems yielded consistent results for analysis
olf the sume Signals,

The Gomimndnt (= petk) frequencies were
calculiled us those of greitest ainplitude ina power
spectrum or an averaged spectral display, Numbers
af pulses were determined by jospection of
wuyelorins. Pulse rates were culealited fram the
interval between (he peak of (he first pulse and the
peak ol the last pulse tra pulse trai and the number
of pulses reduced by one (Le, n-1 pulses), Becuuse
of the difficulty in-determining the hegingings and
ends (LG. zero implitudes) of pulses: and) pulse
(ruins. the peak - peak interval was tiken as the
duralion. Where upproprrite, pulse trains are termed
‘notes’ TP two distinetly diflerent types oF temporal
unil uve present ina call. then the signal is described
ds diphasic (sens Littlejohn & Harrison 1985).

Results

Distriburion ane habitat
Liloria subslandulosa

The new localities (1-9) recorded in Table | extend
the known soulhern range of this species about [80
km All localities are permanent streams/rivers of
basalt or metamorphic rock courtry ussuciated with
nunfarest, montane or wet selerophyll forest (except
for [3a & (3b) and are at S10Q0m or higher ‘The

oH) M ANSTIS & M
14x” 150° E 152°
28°
a"
ee e
32° u Barrington Toes te bd
inigp
34"
30"

O = L. cilropa
@ =L. subglandulosa

ay

a
100 200

Nig. 1. A revision of the distribution of Dfterta eliropa and
Liania sbelandulosa provided by Tyler & Amstis
M1975). invluding wg nimher ol mew lovalines tir fl. whe
fanidlulosa,

km

sOuthernitiost locality at which the species has been
found is locality 9 Pal Brook, Mount Royal State
Forest. NSW. The National Parks & Wildlife North-
cust Forests Biodiversity Study (1991-1994) records
1. subglandiloya ata number of sites between the
Barrington Tops region und the northernmost forests
of NSW, including Doyles River State Porest, MI
Boss Stale Porest, Nowendoc, Wernikimbe National
Park, Gibraltar Range National Park, Styx State
Forest, Spirabn State Porest und Boonon State
Forest. This mdivates the species tas a fairly
continuous distribution along the range country, from
locality 9 in the south to near Stanthorpe, just north
af the QIU/NSW border (151 40"30"E, 28° 40" 20"
S) (big. 1),

On a daytime visit to localities TO and bl oon
19x01. 1994, no tadpoles of this or other species were
located, This was ula tine when numerous (adpoles
af L. swhelandulesa, Lo hewrantongensis ‘and
Mixepivey hathus would be expected to be present
(based on annual studies in the 1960s and '70s).

LLPPELISOUN

Observations by Joho de Baviy and Paul Webber
contirm that there has been little evidence of this frog
over recent yours at the type locality. stuggesuig that
the species may be undergoing a decline there. The
National Parks and Wildlite Biodiversity Study lies
records of five males of (his species culling al three
sites on 2.ix.1995 in the Styx River State Forest in
the region ol the type locality:
1) Rely Creek - lat/long. 30° 34/39" F, 152° 14?
43"S, (altitude L060 rn)
2) Rely Creek - 30° 35°26" BE, 152" 138° 18"'S,
(S90 nn)
3) Waule Mat Camping Area - 30" 35°28 "9, 152"
12°3K"S (870m),
Observations on 20.01.1994 at localities 12, 13h
and J4 (all northern localities), indicated the
presenee Of L. sebelanditlasc tadpoles

Litovia citrapa

Speehinens NMV D32606 (Upper Allyn River) and
D32604-65 (Wombal Creck) were examined und, on
the basis of the indistine! tympanum, prominent
suprutympune fold and head width. were found to be
L. subulandulosd. NMYV 196709-10 cited by Coplund
(1987) as Co cftropa from bear Gratton om the jeoeth-
east coast of NSW, form the basis of the stilement by
Heatwole eral (1995) that L. efrape “extends from
northeastern New South Wales to southeastern
Victoria’. Upon examination, these specimens were
found to have the beady proportions of
subglandulosa, but because both were collected in
1KOS and ina poor state of preservation, itis difficuh
fo come to a definite conelusion as to their Wentity.
The two species have not heen found in sympatry al
any site examined, and this fact im combinuion with
the examination of museum material, indicates (hil
the drvinage of the Hunter River appears. to be a
natural geographic burrier separating them (Pig...

Lurvae were observed by M.A. on 1.i.1076 und
251.1996 ut Boardinghouse Dam in the Watigan
Stiute Forest, south of the Hunter tiver NSW (33° 00°
17H. (S1° 24°7S"S) and hy R, Welly further north
in the Pokolbin State Forest, near Cessnock. in
January 1993) This is the northernmost known
locality Jor this. spevies,

Litoria subghkindilosa appears to replace 1. eiiraper
in the Barrington Tops region north of Neweustle
(Mig) £ cinepa oveupies § wider variety oF
habitats that £. subgdandilose, including permanent
streams in basall country associated with wet
sclerophyll or montane forest, (0 similar streams in
sandstone country. Although found at an altitude of
1066 m at Aberfeldy, Vie, and Blackheath, NSW. /,
citrapeuilse has been Jound in lower coastal areas to
50m (locality 16, Table 3).

BREEDING BIOLOGY Ob 1, SUBGLANDELOSA& 1. CM ROPA |

Adult colour in life
Litaria subslandulasa

Specimens front northern docalities were
predaminunily ween. whereas those fron mid-north
costal localities (f= 9) ranged from untlorm golden
brown with scattered darker mothne over the
dorsum, lo specimens with some small apcus ol
green ofien alone fhe canthus rostralis or under the
eye. ‘lwo mes fram Jocaity | each bad a broad
dorsal patch of green over the head or dorso-lateral
regions Two specimens, AMR763519, from
Gloucester Tops NSW. und another observed by TL
Hines (NP&WS) at Pal Brook (locality 9), were
uniform bright green. apart fre the characterise
golden dorsa-literal stripes.

Some polden-browi specimens developer large
poLht green patches over the dorsune at night (5,
Gow pers. comm). The tuner surtaces of the Hind
limb and groin area were wanstucent yellow. as
found tmadults fron the (ype Joculity.

Lior eitrapa

Lilorta eirepe bias aumitorn golden brown dorssl
colouration (with vreen along the canthus rostralis
and sides ob the bodw). similar to toust specimens of
1. snbelemedilosea Pron localiges 1-9. The principal
Wifference between the species i the colour of the
Inner surkaces OF the hind Limb and grom, whieh iL,
cirope is brick red.

Colling activity
Litorta subulandulesa

Culling begins in spring and) was observed on
20 1994 at localiry B. when water fenmperatures at
Hight were very low. e.g. 6'C. and the diy bulb air
femperatore ab locality Sb (1900 hy was W&'C (A,
White, S, Gow pers. comm,), Other observations by
NLA, at the type locality daring: annual three-week
periods (Dee /Jan. 1966-74). and al all ather
localities fisted in Tuble 1, tndicute that calling
perists drouvhout Decemberanuary ina variety oF
weather conditions, with increased activity during, or
wer. linht pin, Evening dry bulb air lemperatuces
taken ducing periods ob spring/summer actly at the
logalifies ia ‘Table To were 13°C-19.5°C (mean
s7C) At the lower temperatures (13°-14°C).
calling was Jess intense and hy aural comparison
only, Holes were aba slower repetition rate.

Sporadic ditvnal calling Was common daving the
breedine season bul males were most active al might.
Diurnal calling look place trom concealed positions
such as under rocks or from within vevetation. either
near the stream. oc at times up te about four mewes
away from the water, A single male or a small
number of individuals, called from as early as 0742
h (ep, Jocality 7b) Nocturnal calling was. initiated
by one frag, qonmally followed by others ia

distinetly polyphonic chorus, The calls of frogs atthe
southern localines could not be differentiated froin
those of males at the type locality.

Males observed calling ut might were often perched
on broad leaves of trees and stiribs approximately
05-15 mabove streams, on ferns at the edge of the
stream, or on yepelation further from the water's
edyve. They were frequently found calling in small
groups, (Wo or mare metres apart, On 22-1994 at
locality Sb. 40 males were Guilin at night Me groups
of up to six along o 50 um stretch of the stream (S-
Gow pers, com.) Abt locality 7b on 7X0 J994, four
males were calling 25 nm apart (KR, Thum pers,
CONT ).

Anainalysts bf the advertisement call rs provided
helow and Comparison made with that atl errr
‘Two vddilional cal) sequences, altabutable bo,
svhglimdnlose, are in the Broavoustic Library of ihe
Department of Zoology. University of Melbourne.
both recorded by M. J. Litlejott and bis asseeiutes.
The first. fron Coy Fawkes Creek Ehor NSW (3)
24° 20" EB 152° 20’ 46" Si was recorded on
28.8, 1964 aliewel bulb air lemperabiec of BSTC. and
the second. from Flat Roek Creek § hi Woof Port
Lookout NSW (close to the first site), on Lox. 1968
atu Wet bulb oir Temperature oF 13°C) They ute
similar in all pertinent respects to the call deseribedt
here.

Litaria ciirape

Males at Darkes Forest (locality 13. Table 4) were
observed during spritig and suiier callie frog low
branches neside the Seam, on racks near the edee of
the water or on exposed rack shelf ja midstream
close ta shullow. slowly flowing water Ns with 2,
Nuhelardulosie mules called while two ur more
metres wou and aelivity inepeased on overcast
evenmys during or after ram, Dey-hulb air
lemperalures on several nights when males were
calling in September — December 1972-1980. were
14°-22"C" No dina callin was observed.

Advertisement calls
Literta subalandulosa

The advertisement call of 1. subelandalosa wits
recorded by J. Courtney al Diehard Creek, Glen
lines (locality 13a), on 204.93. The dry-bulb an
leniperature was 13°C. The lollowing data were
obtained from the fourth call in the sequence (Pig.
3A). The call has a duration of 9475 5 and consist
of [3 pairs (doublets) of pulse trains (notes), with
each of those in the first five pairs all being of
relanvely low amplitude (fig, 4b). In the subsequent
seven pairs of nutes. the second nate is of much
prediler anyphiude than the first. Thus, all but one of
the first notes Owhich is of equal amplitude) are
softer, with the amplitude of second notes being

a

M. ANSTIS & M. J. LITTLEJOHN

Fig. 2. Live egg mass of Litoria subglandulosa attached to a leaf from submerged overhanging foliage in Tuckers Creek,
Barrington Tops ( locality 8), Seale bar = 10mm.

BREEDING BIOLOGY OF L. SUBGLANDULOSA & L. CITROPA 93

0 2 4 5 8 10
seconds
B Fig. 3. Waveforms of advertisement calls of Litoria sub-

glandulosa and L. citropa.
A. The complete advertisement call of Litoria sub-
glandulosa from which the values given in the text were
derived. This call was recorded at Diehard Creek, Glen
Innes, (locality 13a), at a dry-bulb air temperature of
13.0°C.
B. An expanded waveform of the eighth doublet in the
call depicted in A.
C. A waveform of the complete advertisement call of
Litoria citropa from which the values given in the text
were derived, This call was recorded at the Rocky River
Road crossing on the Brodribb River, 17.5 km NNE of
Orbost, Vic. at a wet-bulb air temperature of 17.5°C.
<i al a Oe
0 100 200 300 400 500

milliseconds

seconds

oy MOANSTIS YM OE LEPTRRIOHIS

greater by upto Ban pais 1-7, cud by 12 120 dB
i pairs 8-13, Durations of doublets range from 29]
te 372 ins (mean = 3344): intervals between doublers
runee from §4 to 159 ms tneun = 11a). The
repetition rte of the doublets is 135 ss! The
doi iant od peak frequencies we within the binge ol
1360-1480 Ha, with means of 1405 Hz for the first
notes and [454 He for the second notes, There are
L005 pulses (incu = leh dain the first notes of cach
of the first seven pairs, and 4-6 pulses (med = 48)
i the remainder, with J4-23 (mein = boob) tn the
secpnd nole of the first seven pairs amd YET (mean
= HO.3) in the rentoinder. Ranges ef durations at test
rds dire 72-80 mays (neu = 85.9) for the first seven
pilirs, 43-69 Ars (mein =59.8) forthe others and (26
IVY ae (mewn (47.0) and 134-159 ms tine 144.2
respectively forthe sceond notes, Pulse rates of first
Holes rHige fuin 125-167 ps! (mean 182.5) in the
first Seven pairs, LO 46-h5 po! (rmeun 64 9) ae thee last
iy. For (lie second notes, the faliges bay plifse fakes
wre 42-182 prs | (mean 145.0) for fhe [est seven pars
and 56-75 ps! (mean h49) forthe last sp

Litopia clirapa

Lhe advertisement call of (his species Wis
deserihed by Litejolin et af (1972) from the
audiospectrographic and oscillographic analysis ol
two calls Of one individual recorded (Nagrne HIB
mcorder, Electra-Voice EV 644 nierophone) at
Tonighs Creck 9 kin Woof Cyn River Vic (149° 08!
BL aT? 34'S) on 244.1960, The mile was calling on
the bank wi a wel-bulb air temperature ef TQaoC.
Owing to background noise levels mm the cecording.
only uiracing of a waveform was provided.

This relatively long call (3.2.3.6 8) was described
as of Complex lernparal stractire (he, strongly
diphasic), with a long introductory pote (910-920
Me Lot high and ropular pulse rie (46 ps), followed
by a sequence al inegularly praduved pulses in
groups of 5-7. The groups have duratigns between AO
vind (20ms and pulse rates OF 34.57 pos! near the
Stall, and ure lonwer (245-500 mis) ahd of lower pulse
rite (1-21 8!) near the end. The (deminant
frequencies ringe trom 135010 1800 He within a
hroud bind of frequeneies between 1250 te 3600 He,

To confirm this deseription, and io provide un
Hididalion of possible effects of emperature, the list
clear wall im the recorded sequence of anothes
individual oF 4. ci/eope was anilysed. The recording
was made al the Rocky River Road crossing an (he
Brodribb River 17.5 km NNE of Orbost Vie. (L487
43°R, 37° 40'S) by Mod, Littlejohn oo 25.81.1981.
This frog was calling from vegetation ala heh of
about 50 em, udjacent to the river, ata wet-bulb air
teniperature of 175°C, A wave fort of this call as
presented in Fie. AC.

The call whieh hus an overall duration of 2.866 s,

consists of a distmet first note whigh is ua regubur
pulse train willy a duration of 81-1 rms, a pulse rite of
{87 ps! and a dominant frequeney of 1040 He. 4
sinwle pulse (duration approximately 7.0) as) wath
dominunt frequenty of 1600) Hy follows. The
remainder of (he call consists of eleven groups ol a
5 pulses bul four pulses and one paral pulses cannot
reaistvally be grouped to allow caleulation of a
pulse rules otherwise. pulse rates range frome 3] ty eb
pal The dominant frequencies of these piilses ranpe
from |200 to 1300 Hea and the durations-of the pulses
ranve from 4.6 to 129 ms, The variable pulsatile
second part of this call has ad maxim dumplituee
wboul SUB higher than that of the miroduciory nate.

Cyiposdton
Literia subelandulase

The advertisement call ah the imale wiry heard on the
bag before umplexus oeeurred. Detuils ol ge musses
lad we presented In Table 2. Qyiposition was jer
observed, bul for cuch of the three capuive: pars
studied, a single eve ness was found adhering to the
Side ol the bag. just below water level. Phe bys were
laid in a small compact clump of two to diree layers
of extremely steky, coabering capsules, Ege
complements for two females were 292 und 425,

Another eight cee masses of this species were
fold wt loeality & Gh 4x7,1994 by AL White and
$, Gow Each mass was attached lo an overhanging
leah Wir ob a rogk Mee i) verrical oc near vertical
orentalion, just below the ywaler surface jn a stuwdy
lowing section of u pool (some in mid-sirean), The
pool Was heavily shaded by an almost complete
canopy cover, Steady tam bod fullem three cdiys
earner ond the surhies water Lempeniture it
1400 Twas YAPC) One OF these passes, removed
from the stream oon a leat and photographer) is
shower in Pig, 2

Litetie citrypa

On 25.40.1973, oviposition ovcurred after the male
and female had been collected at Ourimball Creek,
NSW (loculity 16, Table 4), ut 2200 bh, The Props
were pliced iu phisie bay Che pile saan began to
cull and the pair Was th dimplests three hours aber
capture. AC OLS6 ho on 26.89.1973, the imitial twe
Sequicnices Of OV [POsHTOn acHvity occurred! ACOLST Ir,
i further four ovipesition sequences. foflowed. with
only about (hree seconds belween each, Ovipositian
was complete by OTA9 h,

I) nw typical sequence. the temate dorsilexed her
body with outstretched hind limbs and prodived a
batch of eggs. The inale fertilised ther) while
cupping his feet ina lanning motion around the ewes
The fermale Hen scattered the eges with Uiree sudden
kicking moyements of ber hind limbs, The eggs sank
und spread in a single layer across the bollon. She

BREEDING BIOLOGY OF L, SUBGLANDULOSA & TL CIROPA

Tabet 4. Comperiven of embryes ef Litora subglandwosa and Litoria citeopa

5

Developmental stages are those of Gosner (1960)

Mean embryo
diameter/length
(mm)

Mean capsule
diameter (moi)

Stuge Sample Deseripiion
1. subylanidiilese

Lasiibgland. Locittape

4 tt L. subyland. L. citropa Le subglaned. £. citrepa
2 4 4 1.50 \,73 A457 6) Animal pales black!
vegere! poles dirk grey
7 9 \5 217 3,12 - Body: dark grey,
yolk sae light grey
Head (lateral view):
uculely ungled
Optic vesicles small,
distinet bulge
21-22 j 9 653 6.76 Gilly: anterior 3-4 branches
posterior 4-5 branches
24-25 8 7.34 W65 Lateral lines: pigmented

1, ertrapa

dark browe/
creamy white

dark brown/
creamy while
actitely angled

larger,
indistiner bulge
1-2 brinches
2-4 branches
(shorter)
non-pigmented

Motth-paris; no toath rows or tooth rows,

Keruinised jaw sheath keralinised
jaw sheath
Hatching times 6-10 days 4-6 days

then swim ta another site and the process was
repeated, During the final sequence, the ‘emale
remained in the dorsiflexed position about three
seconds longer, but produced bo eges, The male then
released the female at the point when she began
kicking ber hind limbs.

The exe complement wis 655, Embryos hatched m
four - five days at water temperatures of |7°-23°C.
Two other egy masses laid in captivity continned 890
eves und 928 eggs and took four - six days to
complete hatehing at 16°-23°C (see Tables 3 & 4).

In the field, the eggs were found scattered over the
substrate in shallow pools or slowly Mowing sections
of the stream. which is similar to the mode of
deposition of eggs observed in cuptive pairs.

Tasie 5. Dimensions oe} preserved embryos of Litoria sub:
glandulosa
(mica in mn, range 1 parenthesis, stage -
Gosner, 1960)

Stwe Saniple Embryo diam. Capsule diam,
a 4 1.59 AST
(1.36-1.04) (RI24511
7-4 5S 156 356
(].40-1.50) (3.36-3,85)
16 y 2.05 3,39
(2415-2.05) (3, 283,53)
7 ) 27
(2, 13-2.34)
20) 6 3.80
(5 40-6.24)
24<22 3 HAS
(6.48-6,04)
23 1 O80, 7,34
a4 | 759

Embryonte development
Litoria subglandulesa

Mortality rates of embryos mamtained in captivity
were high. The survival rate (after removal from the
slreum), was greatest amongst embryos in the top
layer of each mass, Those below this layer mostly
ceased developing beyond about stages 8-12,
Embryos from the egg mass held at locality 10 ip
water temperatures of J4™-29°C, survived the
longest; hatching vecurred from ditys 8 -— LO and only
(7 reached stages 20 - 25. The mass from locality 3
did not develop beyond stage 1%, Initially, the
embryos from Jocality 4b continued to develop
during the two days of immersion in the siream
before higher temperatures away ron) the stream
were experienced, Hatching occurred al stages 20 -
21 from days 6-8. with only exght embryos surviving.
Embryos from the southern localities mulch the
following description of those from the type locality
10,

Embryos laid early on 1a.1974 Cocality 10) were
al stage 2 when a sample was preserved at (945 h.
The animal pole is black and the vegetal pole dark
erey. There ure (Wo layers of jelly surrounding the
perivitelline inembrune. Meusuremenis of embryos
are given in Table 5, The embryos were at stages 7-8
after 8 h, and &-9, alter 12 h. Six embryos at stages
7-8. measured after preservation, have a smaller
mean diameter than the same embryos measured live
(1.7 mm live, 1.5 mm preserved: capsule diameter
3.6 mm live, 3.3 mm preserved). After 23h, embryos
were at stages 10-11, and after 38 heat stages 12-13,

Stage 17 was reached after 62 h. A specimen drawn
ulstige [7 (Fig. 4A). 1s desertbed:- prominent optic
vesicle, pronephric swelling. slightanal bulge, large

Oy M ANSTIS & M, J. LITTLEJOHN

——
tomes penemenesssey mB ATTN 4
eee

Hip 4d Brobryos of dilovio subelandalova ad L eltpa
ANF subelanedulose cemoved from its capsule cu ste 17.
Bo Lo ortrape ronperved fram jis cipstle, al ste 17,

OC. Losthvlandidosa just hitched, a approximately stage 21
YF tthe just hatched, avapproximately shige 2).
Seale bur > boa, Shes wre rom Gusner (19604

BREUDING HIQLOGY Ob L, WWRGLANDELOSA & 2 CTEROPS "7

vil-plate swelling. with beginnings of inuseulier
ridges along dorsal Surface just below neural tube. U-
shaped adhesive orvan, slight stomodacal graeve
beguiting te form. Head truncate. acutely angled in
literal vrew. Tiul bud short, rounded, with strong
Jepressing on each side below neunil lube. York suc
grey, resLol body very dark grey After some years in
preservation, body uppears dark and yolk sac lighter
brown.

Embryos examined at 71h were in stages 17-152
growing tail bud pointing acutely ta the left side of
the body within firm jelly capsule: two visceral
arches forming, nurial pits beginning to develop,

After 95 hy stage |&:-optie vesiele more detined
wilh groove forming between this and gill plate;
peutal tube, dorsal muscular ridges, ourial pits. and
divided adhesive organs all more developed.

After 13) b, stages 19-20; spall external gills. gill
cireulavion not apparent, head small, more ronnidec
over eranmal région, adhesive organs diminishing,
ope vesicle depressed shehtly am centres five
vinbryos dark grey dorsally, lighler grey over volh
sim, moving actively within capsule.

Hatching begun eight days. afer ovipostuon, all
surviving embryos had hatched ulter ten days.
Erobryos hatching first on day & were at stage 20 (in
relation to uplic development. but no all
ereeuhihond:- Opbe vesicles indistinet, yolk suc
large deepened stomodaeal pit with adhesive organs
close legether ab anterior chd, a divided ruge ar
posterior end; gills. developing. noticeably sno
adyateed On sitisiral sider vent lube ool well
differentiated: lail fins dusky grey, slightly arched
dorsally; body dark grey brown i preservative. head
region slightly durker,

Stave 2] wis reddhed on day 10) i relation to etl
developinenh and tack OF dail fin cirewlaven only,
(iy, 4C)~ two puirs of well-developed Tunetional
extemal wills. comprising 2-4 branches on interior
pal TS on posterior pair, udheseve organs srall
Iranslieents optic vesicle undefined: fins wanslicent,
degpemmg further, circulation nol apparent: tal
nisculalure poorly developed,

Hive final hatchlings al 1900 hon day 10 were at
slave 22 in relation to bul eireulation. but other
development was assockned wath stages 2)-2 1
corney Sot hol lransparent, prominent but only
purtially plemented optic vesicle. bul fins deepening;
fills al maximum development, fully tunctional,
longer in some specimens than ofhers: adhesive
organs merging to form suall ndge- mouth
thigulir tine of pigment from up af snout through
cach nial pil to eye.

Sue 23:- cornea transparent, eyes well developed,
heavily pigmented; unterior hall at body becoming,
trauspurent around nares; gills diminishing. operculern
developing,

Stage 24;- vent tube more discernible, oral dise
developiag. with sroall triangular funnel above large
oval depression lo become lower labrum.

By day 13. most remaining embryos were at early
stage 25+ golden iridophores scattered iO spots over
dorsum, eyes black wilh seatlerod golden imdophores,
patches of melanin over dorsal surface of til
rousculuture: til fins. body wall mosdy clear. with
some dusky pigment present. Internarial reason
polideably delineated with pigment, lateral line
ors becoming visible,

By day 17, the development of the mouth was
almost complete with the exeeption of the fine black
filanvents, which were either hot yet present, or only
short Unpigmented roots. Dorsal surluce further
piemented with more golden iridophores over areas
pigmented with melanin. ineluding itis: tail
musculatuce pigmented dorsally, im well-spaced
broad bands: fleeks of pigment found aver fins in
Gilder daryae. syel aot Obvieus; ventral surface clean
excep for broad pennmeter of iidophares.

Liturta cinepa

Limbryonie development wis described by Tier
& Anstis (1975). A cojnparalve sunny of embryos
Of L. cittepa and 1, svhglanditesa during stages 2.
17, 2) and 25 is given in Table 4. Pigures 4B,
D shaw stages 17 and 21 In general Lo ectrepet is
larger than 1. sufelattidova theiugheut embryonic
development. with adhesive oygans mere prominent
wud wills sitaller ane less numerous al stuee 21, At
slave 25 and heyond. the lateral tine ormins remain
unpigipented snd mouthparts possess Louth rows ane
aw kerainesed jaw sheuh (Pig. 4, ‘Tyler & Ansiis
1975S). Otherwise, the two species have distinctly
Irujeinte. anuke heads im stage 17 anu similar
hady/ta) Shape throughout embryenie aad larval
development.

Larval behevisntr
Linerter subylanedutese

Tadpoles of this species. observed at all lovalitics
in Table 1. were mostly found on the subswate
shallow, slowly-Howing seehons of the stream on
sind. anmengst tucks or leat liner They were
frequently (ound at the sides of Ure stream:
swimming fast lo deeper mid-stregin or aimonyst
rocks Hf disturbed. They were well camouflaged
whilst on sind or graging amongst rocks and
appeared to feed oo Moveulent sult and algae.
Tadpoles defaeeated rapidly aller capture und the
ubdominal region, while similar a width ti the
branchial region (or shehtly less) in live specimens
HO the streanh, wits Commonly darrower in preserved
specimens,

Tudpeles observed udhering fo the substi
rapidly pulled the body forward 4 distanee of 2-3 mim

98 M. ANSTIS & M. J. LITTLEJOHN

by the use of the oral disc alone, in a rasping action,
This process was repeated continually, resulting in a
distinctive form of locomotion during feeding, which
has not been described in other Australian suctorial
species. Particles of a fine silt suspension were found
amongst the dense, incurved papillae, buccal cavity
und gut of recently-captured specimens.

The fine black filaments of the mouth were broken
or missing In some specimens, or each was present
only as a shorter white filament or core, without the
black outer surface (or pigmentation).

Litoria citropa

The tadpoles were found in small rock pools
(either associated with the main stream or segregated
when river levels were lower), and in larger pools or
slowly flowing sections of the stream. They were
also found on the substrate, but unlike L.
subglandulosa, were not observed moving forward
by the use of the mouth alone; the tail and body were
also involved. They appeared to feed on flocculent
silt and most individuals examined live in the
streams, had well-filled intestines (the abdominal
region being as wide as, or wider than the branchial
region). When disturbed they took cover under rocks
or leaf litter. They were well camouflaged on the
sandy floor and the dorsal colour varied from light to
darker golden brown, depending on the colour of the
substrate and light intensity.

Discussion

Population trends

Comparative field observations of the 1960s-70s
and 1990s showed a marked decline in the
population status of L. subglandulosa at the type
locality, indicating a need for comprehensive studies
on population trends of this species across its entire
distribution.

Advertisement calls

The calls of Litoria subglandulosa and L. citropa
differ markedly in structure (Fig. 3A, C) and cannot
be of any assistance in the confirmation of
relationships based on other criteria. As noted by
Watson et al. (1991), the audiospectrograms of the
advertisement calls of L. citropa and L. spenceri are
of similar diphasic structure; they differ, however, in
that the following notes in the call of L. spenceri are
more regularly pulsed and of higher pulse rate.

Oviposition and embryos

From observations of oviposition sites of Litoria
verreauxti, L. dentata, L. phyllochroa, L, caerulea, L.
chloris, L. freveineti, Limnodynastes peronii, Lim.
tasmaniensis, Lim. ornatus and other species of
Australian frogs, it has been noted that each deposits

TABLE 6. Comparison of body proportions of larvae of
Litoria subglandulosa.
Type Locality 10 compared with new localities
2, 6a & 7a (Table 1).
(Measurements in mm; mean with range in brackets).
Stages 35 & 36 (Gosner 1960).

Morphometric = Type Locality 10 Localities 2, 6a, 7a
Character n=8 n=8
TL 29.84 31,50
(26,40-35.00) (28.50-33.75)
BL. 12.19 11.88
(11.64-12.63) (10,82-13.13)

BW 7.42 7.64
(6,15-8.04) (7.05-8.45)

BD 6.17 6.10
(5.74-6.64) (5,58-6.72)

TD 5.86 5.87
(5.17-6,48) (5.42-6.40)

TM 2.01 2.35
(1.64-2.29) (1.89-2.71)

10 2.49 2.75
(2.13-2.87) (2.46-3.29)

IN 1.88 1.94
(1.80-1,97) (1.80-2.05)

EN 1.46 1.37
(1.15-1.64) (1.15-1.64)

MW 4.55 4.48
(3.77-5,25) (4.10-5,00)

eggs in a similar manner whether in the field or in
captivity (Anstis 1976, Anstis, unpub.). Similarly, L.
citropa scatter eggs over the substrate in both captive
and field situations, and L. subglandulosa attach the
entire egg mass to a surface just below water level.
The egg mass of L. subglandulosa is adapted to the
lotic environment, being compact in form and highly
adherent.

Embryos of L. subglandulosa that survived beyond
stages 8-12 were mainly from the outside layer of
capsules. Mortality may be attributed to reduced
oxygen levels associated with higher sull — water
temperatures in the laboratory of up to 24°C,
compared with 9.4°-15°C in flowing streams. The
embryos from the egg mass at locality 4b continued
development during the initial two days of
immersion in the stream but, after removal and
placement in the laboratory, development gradually
ceased over the next four days in the majority of
cases.

The periods of 6-8 and 8-10 days taken by two egg
masses to hatch (while maintained in containers) are
slower than those of other known stream-dwelling
hylids of lower altitudes, including L. citropa (Tyler
& Anstis 1975; Anstis unpub.). Further comparisons
can be made when data are available for
developmental rates of egg masses within the stream.

The egg capsules of L. citropa are not as adherent
as those of L. subglandulosa. As they are scattered
over the bottom of still pools or very slowly flowing
sections of the stream, stronger adhesive properties

BREEDING BIOLOGY Ob 1, SUBGLANDULOSA & LCITROLA oo

would nor be advantageous. The embryos developed
faster and hud a much lower rate of mortality than
those of L, subylaidulose, possibly attributable to
the individual capsules being scattered over a broad
area, lacilituting oxygenation.

harvie

Whilst slight differciees ih body proportions were
noted between some of the northern and: southern
tadpoles of L. sdbelandilosa (Table 6). only a small
sample from cuch urea was examined.

A sample of 2. sdhelandilosa tadpoles was also
very difficult to muimnliin in captivity al higher
temperatures and 4 second sample maintained in
dented water With Hinution fared io better, Lacking
keratinised. juw sheaths, they could not eat foods
such as boiled lettuce and commercial fish food,
Introduction of siltand dewital sediments Giken frou
(he Streams in their natural environment resulied in
some fecding. although the tilpoles did not grow as
well us those in (he streais,

The distinctive locomotive behaviour of the
fadpoles thyvolving forward proptilsion with the use
of the oral dise alone. distinguishes them [rom the
simiku sympatric species 2. plvilochrea and La.
fesnenei, both of whieh employ some tail movement
during locomotion associated with feeding. Cracdwell
(1975) sides that the M3e muscle ib.
Mbelandulose tadpoles ts inserled in both the upper
wid lower labia. resulting in both Jabia being “pulled

cuudad simultaneously”. whereas “most other
suctorial tadpoles move thetr upper and lower jaws
toward each other during their seraping detion”. This
could explain the Mechanisin behind the distinctive
movement observed in live ladpoles in the stream,
Gradwell also notes that this species has, for its size.
“the longest and densest papillae of the buceul
Nucose’, and these “may act as a steve to exclude
suspended particles above a certain size”,

Examination of gut contents and firther
observations of feeding mechanisms are required to
determine the functional morphology of the unique
mouthparts ol this species.

Acknowledgmeuts

We are grateful 10 the Australian Museum and the
Muscuin oof Vietona for access to and low of
specimens. The NSW National Parks & Wildlile
Service is acknowledged for permission to refer to
records Ol Literia subglandilesa. John Courtney
provided the tipe recording of L. subelandulasa
unalysed. Arthur White provided Figure 4, and with
Kuren Thumm. Jacquie Reese and the late Shane
Gow, added valuable observations on L,
suhvlandiasa Vo this study. David. Duleie and Ron
Anstis, John de Bavay und Roy Seott assisted with
field work, Stephen Richards, Michauel Mahony, Joli
de Bavay, Fred Parker and Michael ‘Tyler
constructively reviewed the manuseript,

References

ANSTIS, Me (1976) Breeding biolowy abd larval
(ewcopnient ol Lite vernuer (Adis bly Hele |
Trams. Re See. S. Aust VO 193-202,

Coprakb. Sh (MYS7) Australian tree frogs of the gents
Aylin Pra Line Soe NS WEB2. (1) Oe TOK.

Gosnen, KL. 1960) A simplified table for staging anes
embryos and larvae WHT Roles oi identheition
Herpetiloeive Vas (43-190.

CRADWELL, NLU T99S) Experiments on orl suction and will
hreahing in five speeies OF Australian Gidpole
(Anuricbhyhidoe anu Leproduetyldtae), Zeal, London
W727, K-98

Hiwiwobb. A. ab Bavay, 2. Wong, Bo & Wiis, (1995)
Paninal Survey oF New Pnghund. (Yo Vhe Prous, Alen, (le
Muy, AR(1), 224244,

Lirrigonig. M0. & Piareisors, POA. (H98S) Phe funcional
sinifigunee of the diphasie advertisement call at
Carini Victarhtnn CAMURE Leplodaety lide). Bedi
Peel ol Soctubiel, VW, 464-373,

) HORUS THEEAS Bb, Matin, ALA A WATSON,
6. (1972) Amphibian Tiana aw? Vietrria, Contirnation
al records of Litoria (=/ yk) etirope (Tschudi) in
Cippstarad. Ver, Med, 8& StS,

Tynen. Mod, & Assis, Mo 11975) Taxondny and biology
ottrogs of the Lirerte cimepe eomples (Anueas bly tear),
Rew NS. Aint. Muy. U7 (5), A1-50.

& (1Y83) BReplugement nun lor
Lively eludulpse Tyler & Anstis, F875
(Arr E ylides), Pray A See NA TOT 2), 1297 30

a. & Davis. VM. C1Y7S) Species groups within the
Australopapoan hylid fre genus fiterie Uschndi Avy
d, Zool, Suppl. Sertes 03, |--47.

WATSON, GF, LIPTL BION, My, HERO, T-M &
RoBherson. B99 t) Conservation statis, ecology anid
monuvement of the Sported Tree Prag ore spencer,
Tech, Kop, Series TG, Arthi Rytote bist Laine Bey
Pepe Cros. Envpam.. Weidethers. View 4000 ve pp

STRATIGRAPHY OF THE NEOPROTEROZOIC ARUHNA
AND DEPOT SPRINGS SUBGROUPS, ADELAIDE
GEOSYNCLINE

By IAN A. Dyson*

Summary

Dyson, I. A. (1996) Stratigraphy of the Neoproterozoic Aruhna and Depot Springs
subgroups, Adelaide Geosyncline. Trans. R. Soc. S. Aust. (1996), 120(3), 101-115, 29
November, 1996.

The Sandison Subgroup of the Lower Wilpena Group is unconformably overlain by
the Wilcolo Sandstone and, together with the Bunyeroo Formation, comprises the
Aruhna Subgroup. The Bunyeroo Formation is in turn unconformably overlain by the
Wearing Dolomite which, together with the overlying Wonoka Formation, is assigned
to the Depot Springs Subgroup. A number of subgroups in the Umberatana and
Wilpena groups is also capped by dolostones that display similar characteristics to the
Wearing Dolomite of the Depot Springs Subgroup. The dolostones are interpreted as
having been deposited on major, sediment-starved hiatal surfaces under cold water
conditions, each of which is adjacent to either a major incised valley or submarine
canyon fill. The differentiation of these unconformity-bounded subgroups is based on
their recognition as genetic units in terms of sequence stratigraphy.

Key Words: Sequence stratigraphy, Neoproterozoic, Aruhna Subgroup, Depot Springs
Subgroup, Bunyeroo Formation, Wearing Dolomite, Burr Well Member, Artipena
Dolomite Member, Wilcolo Sandstone, Wonoka Formation, incised valleys,
submarine canyons, dolostones, Adelaide Geosyncline.

Tremacnons of the Roval Soerery aS Aa O06), T2004), LOL 11S

STRATIGRAPHY OF THE NEOPROTEROZOIC ARUHNA AND DEPOT SPRINGS
SUBGROUPS, ADELAIDE GROSYNCLINE

by TAN A. Dyson*

Summary

Dyson. LA. 11996) Stratigraphy pr ihe Neoproterozoic Aruhna and Depot Springs subgroups, Adelitide
Crosyneline Trans, RK See SMe (1996). 12K, TOL-1IS, 29 November. 1946,

The Sandison Subgroup of the Lower Wilpena Group is unconformably overlain by the Wilcolo Sandstone
HW together With the Bunyeroe Forniation, comprises the Afuhna Subgroup. Phe Bunyeroe Pornmmarion ts i turn
weonformably averiun by the Wearme Dolomite which, lowether with he overlying Woooku Pormaton, 1s
issigned to (he Depul Springs Subgroup, A number of subgroups inthe Uniberatina and Wilpeniw eroups is abso
capped by dolostones that display similar charwleristies to the Wearing Dolomite of the Depot Springs
Subproup, The dolostones are interpreted is having been deposited on mor sedimentstervedt Hattal suriiees
Inder gold water conditions, auch oF which ts tdjacent 16 eithera major menmed valley or submarine envon all.
Nhe (iWerentianon of ese Unconhornity bounded subaraups. is based oh thelr recrgnihon as geaebe units
Tenis on sequence strilienaphy

Kiy Wokbs) Sequence siiatigraphy, Neoproterpzoig, Aruhoa Subgroup. Depot Springs Subgroup. Bunyeroo
Formation. Wearing Dolomite, Burr Well Member, Aptipent Dolomue Meniber, Wileolo Sandstone. Wonoku

Formation. ingised valleys, submarine canyons dalostones. Adelaide Geosynctine,

Introduction

The sthitigraphic nomenchuwure of the Adelaide
Geosyncline emphasises the distinclion between

chronostratigraphie and lithostrangeaphie units

(Preiss LY87a). The positious of the chrono:
Straugruphic units da nok always correspond to
Hithostratiyraphie boundaries. Some lithostaugraphic
boundaries are Unconformities and therelare assume
chronostratignaphic significance, While others are
mappable ttholosteal changes of regional sigmfcance
(Preiss }987u), These differences helween the lwo
stratigraphies con best be accommodated by adopting
io osequence stratigraphic scheme. tt depends on
the recognition of mappable rock units within ot
chrovostratgraphie Framework of repetitive.
genelically-related strata hoonded by uncentormities
or thar corrclauve contormites. “Thus. a revised
stratigraphic nomencluture of Neopraterazoie
suiceessions i the Adelaide Geosyneline could: be
based on differentiation of subgroups within a
sequence stratigraphic framework (Dyson 1992a, b,
1996), Forbes & Preiss (1987) suggested there
was merit in uniting related depositional units ina
siigle subgroup.

* National Centre for Petrotcuin Geology and Geaphysics,
University of Adelude Adelwide & Aust S005

‘Dyson 1A, (1995) Sedimentology and stratigraphy of
the Neoproterozawe Sandison Subgroups a stormedoon-
nated shallow nmarine sequence jn the Adelaide
Geosyncline, Soul) Austra PHD thesis, Planers
University af South Acstralae (iimpub,

Sequence analysis of the Urnberakin Group
(Dyson 19924, 1995! 19960, by) und Wilpenat Group
(yon der Boreh eral JOSS: Dyson 1992b) bas led to
the recognition oF sever! tnconlormity-bounded
depositional sequences. Ina stidy of the Sandison
Subgroup (Dyson 1995!) stratigraphic units
immediately overlying this sequence were examined
order to understand better the spatial and temporal
relationships af the Lower Wilpena Group. The
Sandison Subgroup is wneonformahly overlain by the
Wileolo Sandstone and together with the Bunyeroo
Formation is herein assigned to the Aruhna
Subyroup. Similarly, the Bunyeroo Pormation is
unvonlormably overlain by the Wearing Dolomite
und tovether wilh the Wonuku Pormatian 1s assigned
lo he Depot Springs Subgroup. The Sandison,
Aruhna and Depot Springs subgroups (Piz. 1) are
Uefined uy genelic unity Chat are considered mayor
unconformity-bounded, depositional sequences in
the sense of Mitchum (1977). OF particular
significance is the dature of the Weariny Dolomrre and
iis relationship to other Neoproterozoie dolostones or
unils (hat contin appreciable dolomite in the Adeline
Geosyneline, Le. Nucealeena Formation, Tindelpina
Shale. Warcowie Dolomite and the Artipena Dolomite
Member (new name) of the Enorama Shale. The
names “Wilcola Sandstone’, “Aruhna Subaroup’,
“Depot Springs Subgroup and “Aripena Dolonijte
Member” have been reserved by the Central Register
of Australian Stratigraphic Names,

Aruhna Subgroup

li the southern and central Flinders Ranges, the
ABC Ragge Quartzite is overlaah wilh local

Wh?

MORALANA
SUPERGROUP

WILPENA
HEYSEN GROUP
SUPERGROUP

UMBERATANA
GROUP

oO
(2)
N
So
oF
Li
fo
jo)
oc
oa
oO
we
=

WARRINA

SUPERGROUP CALLANNA

GROUP

ARCHAEAN & PALAEOPROTEROZOIC COMPLEXES

SAN

1. A. DYSON

HAWKER
GROUP

EARLY CAMBRIAN

POUND SUBGROUP

Wonoka
Formation

Wearing
Dolomite

Bunyeroo
Formation

Wilcolo
Sandstone

ISON SUBGROUP

MARINOAN

"EDIACARIAN’

ADELAIDEAN

STURTIAN
TORRENSIAN

WILLOURAN

Pre-ADELAIDEAN

CURDIMURKA SUBGROUP
ARKAROOLA SUBGROUP

Vig |, Stratigraphy of the Aruna Subgroup and Depot Spriigs Subgroup with respeet to selected lithostratigraphie units of
the Adelide Geosyneline. Note the stratigraphic position of dolostones within the Umberatana and Wilpena Groups.

discontormity by a thin (2-5 m), massive, purple,

coarse-vrained lw pebbly cross-hedded sandstone of

fluvial origin (Plummer 1978), In places. it 1s
interbedded with conglomerate and purple shale. It
is, in farm. overlain by geevish red shale and. thin,
imerbedded lenticulir sandstone of te Bunyeroo
Formation with a sharp, conformable contiet, Dyson
(1992b, 1995!) vecogmsed the regional significance
of this unconformity and the nature of the channel
fill facies overlying the unconterniuty. The channel-
il facies is referred to herein as the Wilcolo
Sandstone and is conformably overlain by shale of
the Bunyeroo Fortnation,

The Wileolo Sandstone and Bunyeroo Formation
together constitute (he Aruhna Subgroup (Fig. 2). Tt
ina third-order eyele that is overall transgressive ancl
was deposited ducing one custitie hill and rise ot
relative sea level, A reference section is designated in
Bunyeroo Valley between Aroona Ruins and Wileole
Creck on PARACHILNA, ‘The Aruhna Subgroup
was studied al Bunyerou Gorge, Mount Terrible,
Partacoona. Pettana Gorge, Trebileock Gap and the
Mount Goddard and Angepena Synelines (Fig, 3), A
type section for the Wilcoly Sandstone (hig. 4) 1s

soul of
33!

designated in Wilealo Creek, 2) kin
Bunyeroo Gorge Cat, 31° 25° 10S, long, 138
12" BE).

Lower sequence boundary

The Wileolo Sandstone represents iin incised
valley lil near the lop of the ABC Runge Quartzile,
A shallow palaeovalley can be traced fram the
Aroona Valley (30 in thick) to south of Bunyeroo
Gorge where it attains a thickness of 3 m (Fig. 4).
The base of the incised valley fill is interpreted to be
wsequence boundary that was cut damig a lowstiond
of relative sea level, At Purtucoona (Fiz. 3), the base
of the incised valley is interpreted as a combined
sequence boundary/transgressive surtiee, A possible
sequence boundary exists near the top of the ABC
Range Quartzile at Hidden Gorge (Fig. 3). Here, the
sequence boundary is overlain by g thick (> 10 i),
very course-priuned sandstone or conglomerate that
is Lypically bimodal and very well-sorted. Internally,
diagenetic chert occurs us replucements and
overgrowths. The same texture is observed in the
Wileolo Sandstone near Buryeroo Gorge.

ARUHNA AND DEPOT SPRINGS SUBGROUPS

EARLY CAMBRIAN

vv vv Vv V

POUND SUBGROUP

re
Zo .
cs Wonoka Formation
ae

ral ; :
3 2| Wearing Dolomite
tay Bure Well Nbr,

o
<5
=O] Bunyeroo Formation | MARINOAN
rare
<I Wilcolo Sandstone

ABC Range Quartzite

=o 7 ;
26 Brachina Formation
Fan) oS T ‘Bayley | Ronge Sillst. Mor.
2m Moorilloh Sillst. Mbr-
5 a Moolooloo Siltst. Mbr.

a

Nuccaleena Formation
L Reynello Siltst. Mbr.

Elatina Formation

Trezona Formation

Enorama Shale
Fe ag as ee a

MORALANA | HAWKER
SUPERGROUP | GROUP
WILPENA
GROUP
OQ] HEYSEN
© | supercROUP
MN
O
aa
Lu
ke
O
aa
al UMBERATANA
O GROUP
LJ
Zz

|e ee ee ce ae a aa

BURRA GROUP

WARRINA

SUPERGROUP | CALLANNA

GROUP

ADELAIDEAN

Sunderland Mbr.
Mt Coernorvon Greywacke Mbr.

Tindelpina Shale

STURTIAN

Wilyerpa Formation

Warcowie Dolomite

Holowilena l|ronstone

Pualco Tillite

TORRENSIAN

CURDIMURKA SUBGROUP

UN—NAMED|UN-—NAMED|UN—NAMED]UN—NAMED] UN—NAMED]UN-NAMED
SUBGROUP} SUBGROUP | SUBGROUP | SUBGROUP] SUBGROUP| SUBGROUP

WILLOURAN

ARKAROOLA SUBGROUP

Lo OE ee es

ARCHAEAN to MESOPROTEROQZOIC COMPLEXES |Pre—ADELAIDEAN

Fig. 2, Unconformity-bounded subgroups conforming to depositional sequences in the Umberatana Group and lower

Wilpena Group (alter Dyson 1995!).

V4

104 I. A. DYSON

- © TMUNDAWERTINA
WELL

DEPOT SPRINGS ‘7+ t+
. ian - ANGEPENA,

~ | g SHEPHERDS BOR
“f° @ MOUNT
* , GODDARD
“@BELTANAHILL

STUART

«@ BRACHINA GORGE eae
+ @ BUNYEROO GORGE - ,
“fs D+ 9 @ ARTIPENA HUT
LAKE *:: “os. geE
GAIRDNER. rin Ra
a Karat ae erariccou NG PROVINCE
a ADELAIDE
|. GEOSYNCLINE :.

PORT AUGUSTAPS | ~

NEW SOUTH WALES

{ \. HIDDEN GORGE |,

<
c
‘@)
ke
oO
S

100
KILOMETRES

Fig. 3. Tectono-sediumentary provinces of eastern South Australia, showing localities of stratigraphic sections in the Adelaide
Geosyncline and their relation to other localities on the Stuart Shelf and in the Torrens Hinge Zone (after Dyson 1995!).

ARUHNA AND DEPOT SPRINGS SUBGROUPS 10s

_- Brachina Gorge =
7 10 km _—»"-

Wonoka Formation

Wearing Dolomite

Bunyeroo Formation

Wilcolo Sandstone Strike and dip

ABC Range Quarizite Geological boundary
Brachina Formation inferred

Nuccaleena Formation Type section

Elatina Formation

Trezona Formation

Fig. 4. Geological map of the lower Wilpena Group at Bunyeroo Valley showing the type section forthe Wileola Sandstone
(aller Dyson 1995").

106 1, A, DYSON

Fig. 5, Pebbly cross-bedded sandstone (2 m thick) of fluvial
origin, assigned to the Wilcolo Sandstone, overlying
shallow marine sediments of the ABC Range Quartvite
about 2 km south of Bunyeroo Gorge. The channelised
base of the sandstone is immediately lett of the native
pine in the centre foreground.

BUNYEROO

FORMATION outer shelf

shoreface

WILCOLO
SANDSTONE

ARUHNA SUBGROUP

fluvial

tidal flat

6 dF & &3 € |

inner shelf

ABC RANGE

QUARTZITE SCS
HS

current ripples

frough cross—bedding
herringbone cross—bedding
wave ripples

lenticular bedding
horizontal planar lam.
clasts

sequence boundary
transgressive surface

SANDISON SUBGROUP

Fig. 6. Stratigraphic log of the Wilcolo Sandstane at the proposed type section, south of Bunyeroo Gorge.

ARLUNA AND DEPOT SPRINGS SUBGROLPS W)7

si
mi

nA

Fig. 7. Fluvial channel at the base of the Wileolo Sandstone
is overlain by a 20 emethick mature sandstone thal ism
turn wractienally avertaind by shale of the Bunyernw
Formation, The head of the hammer marks the sharp con
lact between thetwo sandstones, mlerpreted as the tris
gressive surlace

Wileola Sandsroune

The fluvial channel at the base of the Wilculo
Sandstone (Pig. 5) near Bunyeroo Gorge js overlain
by athin (c. <! m), mature sandstone (Pigs 6.7) that
Oficn disphiys swaley eross-stratification 1SCS).
hummocky eross-stratiticution (CS) and sym-
metrical ripples (Dyson 1992h). The hase of the
swaley cross-strutilied sundstune is uilerpretcd as a
Wanseressive surface. At Purtacoona. an un-
conformity al the top of the ABC Range Quartzite is
overlain by 25 m of mature, off-white quartzite (at
displays ough cross-bedding of tidal origin: and
large symmetrical wave nipples with abundant well
rounded clasts of gravel to pebble size. The quartzite
wis deposited in a possible incised valley of sunilar
dimensions to that observed in the Aroona and
Bunyerou valleys. A contact wil overlying shale of
the Bunyeroo Formation was not obseryed. Near
lrebilcock Gup west of Bellana, the Wileolo
Sandstone varies in thickness from 20-50 m where it

consists of interbedded metre-thick, pebble lo cobble
conglomerate, medium to coarse-grained sandstone
and shale, The conglomerate and sandstone display
planur-tubular cross-bedding and SCS respectively,
and are interpreted as having been deposited in a
shorefuce environment within an incised valley (ll,
About 1S km cither side of Trelitcock Gap, ‘the
incised valley fH contains large (S00 x 100 m) nalts
of diapiric breccia, fhought to have slumped int the
incised valley during the early stages of depositor.
On the south limb of the Mount Goddard Syneline. a
greyish red, line-grained sandstone erosively overlies
the Ulupa Siltstone. The sandstone, sbout 1m thick,
contains ungular to sub-rounded clasts of diapiric
material suggesting exposure of a nearby diapir und
is interpreted us being of fluvial origin (Pig. 8).

At Pettana Gorge (Fig. 3), the Wilcalo Sandstone ts
absent bul fora thin remnant of gritty and gossanous
sandstone fl 1s erosively overliin by a boulder
conglomerdte at the base of a submurine canyon in
the Wonoku Formation (Dyson 19951) nat mapped
previously on ORROROO (Binks 1968). The base of
the Wilcolo Sandstone is bot exposed at Hallett Cove
(Fig. 3). but al Mount Termble it is overlam by
interbedded greyish red silistone und mature
sandstone. At this locality, the base of the Wileole
Sandstone is. interpreted as a combined sequence
boundury/trinseressive surface, A. similar situauon
exists at Finke Springs on the north limb ot the
Angepena Sytcline where a thick-bedded, medium-
grained, swaley cross-stratitied Sandstone of
shoreface origin overlies the ABC Range Quarizite
On the south limb of the Angepena Synecline near
Shepherds Bore (Fig, 3), uv decimetre-thick. lididly

Fig. & Conglomerate [rom the Wilcola Sandstone on the
soulher Limb of the Mount Goddard Syncline th con-
tins cirhonite clasts of possible diapirie erigin Coin 4
28 iO tO diameter,

10 | A DYSON

erpss-hedded sundsione erosively overties the Ulu pa
Siltsione. Tt ty overkun by a 30 m-thick section oF
imerbedded greyish red Shile and. sandstaye (hat
grades upward into reddish shale of (he Bunyeroo
Formation,

Buiwverae Parinanon

The Bunyepoo Formation (Dalgarny & Johnsen
196+) is TOO ma thick in ils type seetion at Brachina
Gorge (Mig. 3) where it consists of Liminuicd ia
massive. durk reddish brown shile The overall

Up Ward-Tiiing SUCCESSTON Ts pUNneligred BY a senes oF

subie, Upward-coursenng cycles Hit i places ringe
from 5-10) ms thick. Sedimentary straerurces
dsseciated with very fine to fine-gritined sandstone al
the lop of sume cycles include snillescale cpoys-
bedding and ners TICs.

The Bunyeroo Formation was tor te most pari
deposited below stormy wave base ia middle to
miter shelf setting. Dysow (1992h) placed the
Bunyerou Formation ii at irinseressive systems tract
that wis capped by the forme: Weariig Dolomite
Member of the Wonokia Formation,

Upper sequence haundary

The sequence boindary at the top of the Bunyeroo
Formation as eomerndent will the former Wearing
Dolomite Member of Thomson (1963). leis clevated
herein to formation status to reflect ity regional
signilicunee, Deposition af the Wearing Dolomite ts
Interpreted as having been contemporaneous wilh the
canyon uncontormity ul the base of the Wonoka
Formition (Dyson LO9S' 199Ga, bh).

Depasitional environment

The Wileolo Sandstone wits deposited in a Muvial
und estuarine lo shallow marine envirounent. The
Bunyeroo Formation was deposited in progressively
deeper water ina middle to outer shell’ setting und
Comsiilutes a transgressive systems trict. Thus,
sedimentiuon of the Aruhna Subgroup was unable to
keep up with subsidence, pesultitig ia a depositional
(riliseresston in the sense of Curray (1964), The
Bunyeroo Formation thickens eastward of the
Torrens Hinge Zone (Vig. 3). Adjacent to diapirs,
onlapping sediments of the Aruhna Subgroup tire
thin. However: localised thick development of {he

Hoes. PW. (1987) Carbonate shell and basin
sciliinenbition, Iate Praterozoie Wenoka Korat
South Australia, PhD thesis, Universiny ob Adeiide
iunpuls).

' Dr Bowa, BAL (1989) Guolowic History — SUYUICHIC®
Stefigniphy ot the late Proterazoie Wonoka Mormiution,
nomhor Flinders Ranges Soule Australia, PAD (hess,
Elinders University of South Australian (uupuly),

Bunyeroo Formation oecurs adjacent lo some
diapirs. A thick. black stiecession of sulphide-rieh
shaleadjacent to the Mucatoona Diapir (Coats 1973)
sugeests tinowie, deep water deposition of the
Bunyerao Formation, perhaps associated with the
formation ofa erestil gtaben over the diupir due (1
salt depletion. The diaipirs of (he Flinders Ranges
offen contain volcanic xenoclasts (Preiss 1987h). aml
Coats (1973) suggested that nany diapirs on
COPLEY were uetive and exposed Curing deposition
oF the basal Bunyeroo Pormation, An inferreel
volcanic Component of the redheds (Mawson (039)
Plummer (978u) may be related to deposinunal
onlap Of Bunyerao sediments adjavent to exposed
dhupirs,

Dalrymple (1992) Sugpested Uhal estuarine
sandstones were transgressive in origin because
estuaries owed (heir existenee to marine (ovding ut
incised vatlleys. On the other hand, Exxon researchers
(eg. Vuk Wagoner er a/. (987) aieied thar Muyial
sediments at the buse of incised valleys should he
assigned lo the lowskind systems trace deposited
during an initial fall and subsequent curly rise of
relative seat level, Alternatively, such fluvial (arts
may be the updip equivalent of Lrapseressive murine
sandstones. The kick of beuch deposits between the
Mivial and estuarine sandstones of che Wileule
Sandstone at Burryeroo Gorge sugwests thal the hase
of the estuarine sandstone is the transgressive
surface “The ensuing transgression eroded und
reworked (he former beach sediments.

Depot Springs Subgroup

The Woroka Formation (Dalgarna & Johnsen
1964) and Wearing Dolomite together represent a
transgressive-regressive (ER) cycle that is referred
tous the Depot Springs Subgroup (Fig. 1). The Depot
Springs Subgroup constitutes an uncontormity-
hounded depositional sequence and was studied at
a number of localities on PARACHILNA, COPLEY
and MARREb including Petltuna Gorge, Wyucer
Blut, Brachina Gorge, Bunyeroo Gorge. Beltuia
Wilk, Mount Goddard. Shepherds Bore ond
Mundawertinu Well (Fig. 3). A reference section for
the subgroup is designated ih and adjacent to
the Patsy Springs canyon of the Angepena Syncline
fear (he Depot Springs HLS., 40 km east of
Copley (Fig. 3),

The base of (he Wonoka Formation was mapped on
PARACHILNA (Dalgarno & Johnson 1966) where ii
colour change oecurred above greyish red sandstones
at the top of the underlying Bunyeroo Forination,
This boundary corresponded ta a rather abrupt
increase in lime coment, Gostin & Jenkins (1983)
defined a deeimetre-thick dolostone overlying
reddish shales of the Bunyeroo Formation. referred

ARUIINA AND DEPOT SPRINGS SUBGROUPS uy

Fix Y. Sharp contact bewween the Bunyeroo Formation and
Wearing Dolomite fh the Angepena Syneline, The
Wearing Dolomite is about 30 cm thick. Hs hase is
marked hy the head of the hammer with sedimentary fic
ing to the right.

Fiz. 10. Wearing Dolomite displaying micro-HCs,

Angepena Syncline.

to informully as the Wearing Dolomite Member
(Thomson 1965), as the base of the Wonoka
Formation, Jenkins (1993) detined the base of the
Wonoka Formation in Bunyeroo Gorge at the base of
an intraformational conglomerate within the Wearing
Dolomite Member, Haines (1987?) divided the
Wonoka into 11 litholacies uints. These units were
subsequently adopted by other workers (e.g. Di
Bona 1989; Christie-Blick e7 af L990) with the
prefix W.

Lower sequence boundary

The base of the Wearing Dolomite is defined as a
sequence boundary. [can be either sharp or diffuse
iw matte, Jenkins (1993) interpreted a sequence
boundary at the base of an intraformational
conglomerate within the Wearing Dolomite,
However, the intraformational conglomerate displays
edgewise clasts thal hive in the pust been iiterpreted
us storm rosettes (je. Dyson & you der Boreh
1986). Dyson (1992b) suggested thal the Wearing
Dolomite fepresented deposition within a condensed
section thal included a possible maximum flooding
surface. Furthermore, the Wearing Dolomite was
deposited on a sediment-starved hiatal surface below
storm wave base (see below),

Wearing Dolomite

The former Wearing Dolomite Member of
Thomson (1965) is a thin cream. dolostone or
dolomitic siltstone that has been mapped over
extensive areas of the Flinders Ranges (Forbes &
Preiss 1987), IL corresponds to unit | of Haines
(19872). The Wearing Dolomite often sharply
overlies the Bunyeroo Formation with apparent

_—

Unil W3

Unit W? ff

Unit W3t
Unit W2 _ Unit WI ee

A a A acl

Unit WS

thin bedded lempestite facies

Unit W2

‘wall plaster’ dolostome focles

QPL & HCS tempestite focies

Burr Well Member

stromalolitic Imestane fovies

SCS sandstone facies

Cop Dolmslone Member
lominaled dolostono foctes

| Cenyen Fill Member
‘wall plaster’ doloslane facies

WONOKA
FORMATION

WEARING
OOLOMITE
unlit WI

slump facies

3 cross~bedded pabbly sandstone facies

Fig 11 Schematic cross-section through the Patsy Springs Cunyon in the Angepena Syneiine Gifter Dyson 1995!), Note
the relationship between dolostones of the Wearing Dolomite,

Wo La DYSON

Mig. 12. Breeciola limestone af the Wearing Dolomite
uneuntormably overlying the Bunyeroo Pormation east
of Beltane Tih,

conformity (Pig. 9). (bis characterised by wavy to
parallel kumination ind, less commenly, by micro
HCS (Fig. LO). A similar sittation exists on the south
limb of the Angepena Syncline and north limb of the
Mount Goddard Syncline. Near Mundawertina Well
and on the northern limb of the Angepena Syneline,
the Weuring Dolomite splits into two thin dolostones
that ure separated by Up ta 25 m of lanmimated grey
given shale, AL Petlana Gorge (Fig. 2). a 2 m-thick
boulder conglomerate cuts downward into the ABC
Range Quartzite. I marks the base of a shallow
cunyon with an estimated relief of 70m. About S km
south of Pettana Gorge near Wyaceu Blull the
Bubyeroo Formation altins a thickness of about 30
in Where iL is sharply overlain by a deeimetre-thick
dolostone of the Wearing Dolonite with sharp,
planar contact, It is interpreted as the correlative
conformity adjadent to the submarine cunyon,

Other dolostones are also genetically related to the
Weuring Dolumite at this stratigraphic level (Pix.
(1), About 2 kin cast of Beltana Hill (Fig 3), the
Weuring Dolomite passes laterally tito a several
Mietre-thick. creium to orange laminated dolostone
that is offen brecciated’ and present as detached
slump blocks (Leeson & Nixou 1966), Here, iL
overlies reddish shales of the Bunyeroo Formation
wilh an angulay unconformity (Pig. 12). This
dolostone ty thought to represent the “wall plaster
veneer” Of Rickoll (O88). Ao similar relitionstip
can also be observed near Mundawertina Well, and

ise on the south limb of the Angepena Synetine

where the Wearing Dolomite cun be traced into
slumped wall plaster sil the edge of the Pitsy Springs

CHICKHOW, KAA PU 88) Geologie Hteey and nig ak Me bie
Protenozae birtress Alt Canyon Chrples. Adelaide
fwesyneling, SON. PAD Theses. Tiers Crisereity at Seneh
AUSTELL (pub),

oanyon (Dyson 1995'). On the south limb of the
Maunt Goddard Syncline, the Burr Well Menther of
Di Bons (1989 consists of mature, medium
grained, swuley cross-siratified, crarbunite-cemented
sandstone or intraformutional conglomerate with
edvewise clasts of dolostone, and passes: literally
into stromutolitie dolostone (see Fig. 11), 4 basal Tne
comprises diapiric detritus. Where the Wearing
Dolomite displays such uncontormiry, it is defined as
the Burr Well Member un the sense of Di Bons
(1989').

The base of the Wearing Dolomite ts Tlerpreted by
he wdeep Water sequence boundary, A masimun
flooding surface al the top of the Buniyeroe
Formation may caineide with this sequence
boutidary. The diffuse base of the Wearing Doloinpite
sugeests (hat carly post-depasitional dolomitizaton
took place on a sedimentstaryed hiatal surface.
Wavy to parallel lamination and mero BCS lurther
suggests depositiun below storm wave hase (ee.
Dyson 1995). The Wearing Dolomite ean be traced
into the Burr Well Merber on the south limb of the
Mount Goddard Synetine. The Burr Well Member
was deposited in a storm-domimated shorefuce to
lide-dominwted. higoonal environment ts sharp,
erosional base is a cOmbined sequence boundary!
transgressive surhice, The “wall plaster veneer”
Was deposited on the canyon shoulders, to ts
interpreted fo be coeval with deposition of the
Wearing Dolomite and unit W2 of the Wonoka
Formation (Fig. 11).

The Wearing Dolomite, together with the Wonaka
Formation, is present on the Stuart Shelf where it cunt
be observed in deillcore (e.g., Bupeechee 2 at 367.2
in) A possible Wearing equivalent, only a few
centimetres thick, crops out south ef Bills Lookout
near the north-western side of Luke Torrens within
Whil was previously interpreted as Yarloa Shale
(Jobns era, 1966). This stigvests that the Wearing
Doloniite transgressed the Stuart Shelf prior to
deposition of the Wonoka Formation.

Wonake horniation

The Weuring Dolomite is overlain by unit W2 of
the Wonoka Formation, consisting of greyish red,
fine-grained sandstone and interbedded culearcous
shale, Tt grades upward jote whe domiiantly
cul¢urcous shale of unit W3. The sueecession
represented by units W2 und W3 is averall
transgressive. A ¢olour change to areenish
limestones in the middle seetion of anit Wa murks
the base of regressive sedimentation in (he Wonobki
Formation (Pig. 11),

The lithofacies units W3-7 inclusive of the Wonoka
Formation display an overall upward sunding trend.
li culminates ing thick, storm-dominated, mixed

ARUHNA ASD DEPOT SPRINGS SUBGROLTS Ut

curbonate/siliciclastic succession, commonly displaying
HICS (Haines [Y8X) that was deposited in un inner to
middle shelf setting, Units W3-7 of the Wonoka
Formation. as interpreted by Maines and other
Workers, represent regressive sedimentation from
inti) deposition below storm wave base to
deposition abowe fiirweather wave -byse.

A number of metrethick, greyish red, meditin
wrained sandstone beds in unil WT that ure
vharactemsed hy SCS may represent toreed
regressive deposits associted with a lilling stage
systems tet (ca. Dyson 1996c). In the Angepena
Syneline. a disconformily ut (his Stratigraphic level ys
overlain by a several metre-thiek medium-grained
sundklone that in many places disphiys horizontal
phn Janination and SCS. Il is interpreted to
represent deposition on the lower shorelace in a
Invine environment Di Bonk & vor der Boreh
(1993) interpreted a lowstand delta al this level in the
Uinberating Syvneline Unit W7 is overliin by a
suecessivim Of shallow nmiarine sandstone and
carbonate of tidal and lagoonal origin (Huines 1990)
that corresponds to units WR EL. Occasionally, these
unis cu be observed to grade upward ito the tec
shile and. simdstone of the Bonney Sandstone, cat,
hear Mount Goddard,

The TER cycle of lhe Depot Springs Subyrouyp is ae
unconformiry-bounded, third-order depositional
sequence, The cunvon Oil represemed by units WI,
W2 ikl to a lesser extent W232 eensttute tie
Ininsuressive evele of the Depot Springs Subgroup,
Deposition of unit Wl in the canyon Fil wats
qiintemponuimous with deposition of (he Wearing
Dolomite On a sediment-sturved Tati ssulaee A
possible condensed seetion is represented by uni
Wa, The mille of this unit corresponds to air qhrupt
cylour change and increase in lime content, A
THM Moodie surfioe may be contimed: within
Wit Wa and oy dierelore equated with a cownlap
surface His overliin by the meressive Hrhafieres ol
(he Wordka Foriition,

SUA OTHE: CON TES

The Weating Dolomite wis developed adjacent Lo
ormajor submiwine unconformity an a sediment
starved bigdal surface (hat corresponds to u combined
sequence boundary and migor flooding surtaec
(Dyson M954, Th cun he observed ty piss fierally
ints slump breeviokis on the shoulders of submarine
cunyons. eo. near Beltane Hill south tinh of
Anyepena Synchine, On the north Jitnh of the
Abdpepena Syneline, the Wearing Dolomite consists
of hwo dolostones that are separated by some 20 mot
shale. The upper dolostone overlies the (runeated
edge of a lower dolostone, sugegsting canyon
grosion occurred before deposition of the upper

dolostone. Furthermore. unit W2 overlies the Til of
the Patsy Springs canyou in the Angepeno Syneline.
Retrogressive slumping on the outer shell was the
precursor (o canyon ticision, and proeeeded up to the
level of unit W3 in the Wonoka Pornation which
marks the turn around from transgressive lo
revressive sedimentation im the Wonokie Bormution
(Hig, 11). This interpretation questions the timing ob
curligst Canyon incision expounded by orhey workers
ihal coincided with deposition of units W3. We oi
W5 (e... Haines P87: Di Bona 1989S) Christie
Bhick ete. L990, L995),

Upper sequen bomndary

The Bonney Sandstone alieo displays ao sharp
thoveh apparently conformable contiet swith the
underlying Woroki Porvaion, Flowever the
relationship becomes discontormable im the weinity
Of diapits. Adjacent te sume divpirs. eg, Pineda
Diuproon COPLEY und Prome Diapie on
PARACHILINA, Whe Booney Sandstone displays an
unconformible relalronship wath the Wonka
Formation (Dyson unpub. ‘Thus. (he base of the
Bonney Sandstone of the Pound Subgroup ts
iiterpreted us a Sequence houndary.. Dysan (1Yos!.
199g, UNpUb) suagested (hat (he development 0!
several prominvnl cneonformities or seyuehee
boundaries within the Umberatana and) Wilpena
Groups was associated with pertods of aetive: and
passive daprasn. which oF Lup Was related to major
catcnnioiil events during break-up of the
Neoprotenzore superconunent,

Discussion

The wwe of suberoup asa iicenferintiy: beaded Wit

The previous use of sdbyroup in oa lithe
stratigraphic sense has caused problems in regional
mapping programnies. Ao good example is the
differentiation of interglacial deposits in the
Uinberauink Group, They embrace the Paring
Suberoup (Coats ef af, L986) Thompson (969) and
ihe Willochra Subgroup ¢thamsei (969). Phe
infention of (he term Farina Sdberoup was to aeltee:
all relitively deeper walter sediments, in
conthadistinetion frein the dominantly shallow water
redheds of the Willuchra Sahyroup (Coats & Preiss
1987), There has been inconsistency Heapplication of
the terms. espeenilly af sotie Wansitfonal regions
where facies infertongue ornare inlerealwed (Coats &
Preiss 1Y87). The southern portion of PARACHILNA
is such a region Where a basis for this distinetion 7s
warranted. ‘Therefore, a revised stratigraphic
fomenckiutire of the inferghticial deposits could: be
bascd ou dplicren gilien of Subgroups tis i wenede
HO fers OF Sequence sini graphy

2 LA. DYSON

A seguenee stratigraphic lramework for. (he
Urpberatana and Wilpena groups is shown in Fig. 2
and is bused on dilferentialion of subgroups asa
genetic unit, An uncontormity-bounded sequence
incorporutes the "Tindelpina Shale and Tapley Ell
Formation, Similarly, ai uncoutornmity-bounded
sequenee is defined by the lop of the Tapley Hill
Formation to near the top of the Etina Formation, and
wroverlyiig sequence 1s capped by an unconformity
aM the base of the Elatina Formation, The Willochra
Subgroup previously meluded the Murinoun glacials
of the Elatina Formation (Thomson 1969). The
Klatina Poemation is capped by dolostune ol the
Nucealeena Formation, the basal unit of the
Sandison Subgroup. The Sandison Suberoup is
unconformably overlain by the Aruhna Subgroup
(hal comprises the Wileolo Sandstone and Bunyeroo
Formation, The Aruhna Subgroup is in tien
Unconformably overlain by the Wearing, Dolonite al
the base of the Depot Springs Subgroup (Fig, 2). A
similar scheme was proposed for the Kanmantoo
Group of Cambrian age (Dyson 1995"), More work
is required to differentiate genetic units within the
Pound Subgroup which contains possibly two
Uncorformity-bounded sequences, wilh sequence
boundaries al the buse of the Bonney Sandstone and
Eciacurd Member ol the Rawnsley Quurtzite (¢.2.,
Dyson, 1995?).

Lithostrutigraphic, biostratigraphie and) chrono-
stratigraphic units will continue lo be used as the
basis lor most stratigraphic studies. However, the use
of iinconformity-bounded units ts invaluable in
busins where the development of stratigraphic units
was controlled by (eetanie episodes id custatie sea
level eyeles, Th such basins. eg. the Adelaide
QGeosyneline, unconformities pass laterally into
correlative conformitics where traditional strari-
praphy is tatikely to differentiate the lateral and
vertical extent of genetic units aboye and below the
sgquicnee bouldary, Thus, the use of uncontormity-

bounded Units cah Conteibute to (he understanding of

the suadgraphy and geologic history of a basin, it
can provide the Iramework for regional stratigraphic
framework and it can enable the mapping and
expression of stratigraphic concepts for whieh other
stratigraphic units are inadequate (Salvador (994),
A hierarchy of Uneontormity-bounded units can be
formulated by determining sequence order ina basin.
Mitchum & Van Wagoner (1991) proposed a
sequence boundary Mnerarchy of five orders on the
frequency of boundary occurrence. Alternatively,
Embry (1993) suggested five orders of sequence
boundaries based on the nature of the boundary. The
latter Method is possibly less subjective, However, il
every pair oF unconformilties is used to recognise and
name an uncontormity-bounded unit such as in the
cise of the Cardium Formation i the Cretaceous

Western Interior Seaway of Canada (Walker 1990),
the stratieruphic units would grow unmunageably
(Sulvador 1994). The tise of the subgroup as a
depositional sequence of third-order cyelicily
thereby usefully lirnits the establishment al
meaningtul and usefol straagraphie unity Gnu
regional wna inter-regional busis.

The sientficaice of dolostones capping Adelaidean
NEGUCHCES

The Wearing Dolomite of the Depot Springs
Subgroup was developed adjacent (0 4 major
submarine unconformity on a sediment-stirved hiatal
surluce that corresponds lo a combined sequenve
boundary and major flooding surface. There are other
dolostones or units containing significant amounts of
dolostone that fre associated with sequcnce
boundanes in the Umberatuna and Wilpena Groups
(Fig 2), They are the Warcowie Dolomite. the

3

aT

4
A

Warcowie
Dalamite

3 Tindelpina

z Shale
sé al
om SAMPLE LOCALITIES
a. 1: MUNDAWERTINA WELL

-5 2: SHEPHERDS BORE 4
> 3: UPALINNA DIAPIR
2 4: WILLIPRA ANTICLINE
© St ARTIPENA HUT 5,
A
Artipena

Dolomite Mbr.

—10

A Nuccaleena 1
Wearing Dolomite Roptriatfen
Ulupa 1
1 Siltstone &
-15 ;
=10 —5 0

6'C PDB %,

Fig. 13, Lsotupic data far Adelaidean slolostones (based on
Dyson 1905!)

4RUHNA AND DEPOT SPRINGS SUBGROUPS 113

Tindelping Shale and the Nuccaleena Nornmtion, all
of which cap sequences of glicigenic origin. They
are commonly referred lo as cup dolostones (c.a.,
Williams 1979), Another dolostone occurs within the
lower Knorauma Shale and is prominent on south-east
PARACHILNA. c.a., between the Willippu Anticlne
aud Martins. Well Dome. This formally referred to
here is the Artipena Dolomiue Member of the
Enoruna Shale and a type section ts desizmuted 2 km
south of Ariipend Hut (Pig. 3), Flere. i consists of
two thin CLOS0 em) dolostones that are separated by
Am ol ereyish red shale, and marks the transithan
from lanmsgressive to overlying regressive
sedimenticion of the Enorana Shale.

These dolostanes of the Umberatand and Wilpena
sroups display similar charaeteristics and. are
interpreted Lo have been depositedt yn inajor Nooding
surfaces under cold wiler conditions. Their isotopic
Jana. based an Dyson (1995!) in Big. 13. display two
upparent Gends. ‘The curbon isetupe values show a
shift to more pevative values with possible incredse
im owaler depth, accompanied by ai inerease 1
diagenesis. This sugeests thal the Wearing Dolomite
was deposited in greater water depths than the other
Jolostones. and was most susceptible to secondary.
post-depositional alteration. The oxyven isotope
vilues showeu shifl lo nore negative 6MO aboye the
Siraligraphic level of (he Warcowre Dolomite. dO
is senmsinive ty lemperituce changes. then i) might
Show a similar trend to relative water depth is
suggested by BUC (ee, Buu eral, 994). However
the shift to more nesarive 60 above the Wareowie
Dolomite is interpreted Lo reflect the oyerwl| increase
in palucotomperature following the Startian
vlncnition, This iterpretivion must be viewed with
caution, particularly with respect Lo the Wearnig
Dolomite, becuse of overpriils associated wilh
secondary, pos-deposiionul alteration. Depositiin
at qhe Wareowie Dolomite iv a glacivenic
environment is suggested by the presence of
dropstones (Dyson 1995!, 1996b). Pulaeo-
temiperatines for the Neoproterozoic dolostones ire
thought ty range from S' © for the Nucewleensa
Formation and Wearme Dolomite. to 5° C for he
Warcowie Dolemile (CLP Rao pers. comin, 1995),

dach of the dolostones was depasited on a
miximiim flooding surface assoenited — wath
lergenous starvalion, The Milendella Limestone, a
carbonate oF Cambrian age, occupies a simifur
stratieniphic positing in the Kunbintwo Croup.
Jnemed valleys are associated with the Milendella
Limestone and Seaelil Sindstone (Dyson & von der

Borel 1994: Dyson 19964). Prograding slope
compleses ure assocned with the Tindelpina Shale
Such units are considered ty be the downslope
equivalents of incised valleys (Mitehuntet af, (99S),
A pertinent question is why dolostones of the
Wearing Dolomite and Wareawie Dolomite do not
appeur to be ussocited with imeisced valleys Uhat
show dominantly shallow water features. Mayor
extensional evenls comeiding with deposition of
these units, together with high rates of substdence,
restilled in no fallin relative sea level and precluded
development of sandy highstand facies, Instead.
dolostones cup rehlively deep water sediments of the
Bunyeroo Formation and Holowilena tronstone
respectively. Buel dolostone ts developed on an
Iiatul Surface that represented u period ol terrigenous
Starvation. In cuch case, this sticfiwe cad be followed
into the Submarine uncoulormity where a major
eanyon was possibly cul on the ouler shell In such a
setting. a flexural response of the continental margin
muy have oecurred as an is@stuhe readjustment to
eadnyort erosion (McGinnis en al, 1893). Fora wie
continental shell, (lesural uplift oF (he outer shell
would nol have influenced the position of (he
shoreline. resaliing a an erosional uncanformity
developed only aeruss the distal shell (MeCainnis ef
al 1993) The fits generated would be greatest
across [he distal shell and deerease ina landward
direction. Incised valley fills of fhe Seach!
Sandstone dnd Milendella Limestone are overall
(ranisgressive but display rekitively shallow water
features at their base. Sequence boundaries at the
base OF these meised valleys were formed dhiring
lesser exlensional events when the degree ob
subsidence was rehitively small or more likely, when
the widih of the pulacushell was relatively narrow.
Thus, a full in relative sea level on the outer shell
resulted in subaerial ieescd valleys comprising basal
uyial to estuarine deposits,

Acknowledgements

This paper rs bused on field studies uiidertiken
during postgraduate research al Flinders University
under the Sapervision ol Professor Chris von der
Beveh. Several useful comments were suggested by
reviewer's Clirts Nedin and Wolfgang Preiss. Aspects
of the current study were discussed with Peter Reid
and Wolfgang Preiss. Gail Jackson and Cihazi
Keaishnan kindly drafted the ffgures.

i | A DYSON

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STRATIGRAPHY OF THE NEOPROTEROZOIC TENT HILL
FORMATION AND SIMMENS QUARTZITE AT SOUTH TENT
HILL ON THE STUART SHELF, SOUTH AUSTRALIA

By IAN A. Dyson*

Summary

Dyson, I. A. (1996) Stratigraphy of the Neoproterozoic Tent Hill Formation and
Simmens Quartzite at South Tent Hill on the Stuart Shelf, South Australia. Trans. R.
Soc. S. Aust. 120(3), 117-129, 29 November, 1996.

The Tent Hill Formation and Simmens Quartzite represent regressive, shallow marine
sedimentation of the Sandison Subgroup on the Stuart Shelf. At South Tent Hill, the
Tent Hill Formation comprises the Tregolana Shale, Lincoln Gap Siltstone and
Corraberra Sandstone members and represents deposition in a storm-dominated shelf
environment. Sharp-based, swaley cross-stratified sandstone beds of the Corraberra
Sandstone Member are interpreted as forced regressive deposits formed above
fairweather wave base. The overlying Simmens Quartzite was deposited on a broad,
open shelf that was conducive to tidal amplification. These units are correlated with
their stratigraphic equivalents in the Adelaide Geosyncline.

Key Words: Stratigraphy, Neoproterozoic, Tent Hill Formation, Simmens Quartzite,
Lincoln Gap Siltstone Member, Sandison Subgroup, Stuart Shelf, Adelaide
Geosyncline.

Transactions of the Reval Seeier ap S Anyi (1996), 1204), LET 128

STRATIGRAPHY OF THE NEOPROTEROZOIC TENT HILL FORMATION AND
SIMMENS QUARTZITE AT SOUTH TENT HILL ON THE STUART SHELF,
SOUTH AUSTRALIA

by LAN A. Dyson®

Summary

Dysos LA. (1996) Stratigraphy of the Neaproterozore Tent Hill Porriation and Simmens Quartzite ar South
‘Vent Wilbon the Stari Shell, South Australia. Yams. Re See. S. Ave. 124K 3), 117-129. 29 November, 196,

The Tent Will Formation and Simmens Quartzite represent regressive, Shallow marine sedimentatoaa of ic
Sandison Subgroup on the Stuart Shell At South Tent Hill, the Tent Hill Formation comprises the Tregutinet
Shile, Lancoli Cap: Sitstone and Corruberra Sandstone members and represents depasittan tia stor:
dominated shelf environment. Sharp bused, swuley yross-straticd sandstone beds of the Corriberry, Sandstone
Member are interpreted as forced regressive deposilys formed uboye Turweather waye base, The oyerlyiny
Somens Quartvite was deposited ut a hrowd. oper shell thar was conducive to nidal amplification, Vhese units
ure Correlated with their siitigraphie equivalents in the Adehide Geokyacline

RY Words: Stratigraphy, Neoproterocoic, Tent Hall Pormaion. Sanimens Quartvile, Lincoli Gap Siltstone
Member, Sand)son Subgroup, Staart Shelf) Adelide Geosynctine.

Introduction

On the Stuart Shelf, the flat lying sediments of the
Toot Hill Pormiation crop aut west of the Torrens
Hinge Zone and Adelaide Geosyncline in South
Australia, The Tent Hill Formation (Browe (885)
wits named ufler the Flattopped hills 25 km
northwest of Port Augusta (Pi. 1), As part ofa major
study (Dyson 1995"), the sedimentology un
stratigraphy of the Tent Hill Formation and Simmens
Quarizite Were mVesugaled (1 the type section at
South ‘Tent Hill (Pig, 2) and represent the first
detailed syothesis of the sedimentology ane
stratigraphy of these formations. This paper revises
the Stratigraphic nomenclature for the Tent Hill
Formation on the Stuart Shell.

At South Tent (ill the Tent Mill Formation was

formally detined: as consisting af three members,

munely the Tregolina Shale Member, the Corraberru
Sandstone Member and the Simmens Quartvite
Member (Dalarna e7 al 1968), This study has
elevaled the Sinmmens Quarizite Member lo
formation status and redefined the Corraberra
Sandstone Member thereby incorporating the
Lincoln Gup Siltstone Member into the Tent fill
Formation (Fig, 3). The Tent Hill Formation ts
conelalve with the Brachina Pormation i the
Adelaide Geosyncline, Similarly. the overlying
Simmens Quartvite may be correlated with the ABC
Runge Quartante, The name “Lincoln Gap Silestone

National Conte for Petpaleii Cleahoey dark Gadpiysies.
Livers of Adelaide Adelaide S. Aust SOUS.

Dyson. 1A (OOS) Sedimeitvtopy amd staligraphy al Ue
Seoprolerovo Saindiseu Subproups uv stormeduniiiaited shallow
minine sequvace in the Adelaide Cleasynctine Soul) Austeatio
Phlo Cheses. Flinders. Liniversigy oF South Austrabia Ganpubs >

Mernber" has been reserved by the Central Register
of Australian Stratigraphic Names,

Previous work

The seerjion at South Tent Hill (Pig, 4), originally
Jeseribed by Thomson (1965), wis proposed by
Dalgarno eral. (1968) as the type section for the Tent
Hill Formaten., The underlying formidion was
referred to as the ~Tregolina shales by Miles
(1955), The term “Lincoln Gap Fligstones” was
proposed by Miles (1955) for theesandy suceesston
ol (he Tent Hill Formation. Crawford (1964) referred
to the lower part of tus unit ay the Corrabecra
Sandstone. ‘Thomson (1965) detined the Corriberra
Sandstone Mejaber and the Simmens Quartztte
Member as constituents oF ihe Tent Hill Formation.
Coats (1965) correlated the Tregolana Shale Member
wid Corruberra. Sandstone Member with the
Braching Formation, and the Siminens Quurtvite
Member with the ABC Range Quartaile, Johis
(1968) proposed the lerms “Woomera State
Member” and “Arcoona Quartaile Member” on the
northern Stuart Shelf where they were considered
luteral equivalents of the ‘Tregolina Shale and
Sunmens Quartzie Members, respectively (Coats
TYH4).

The sttubgraphy of the Tent Hill Formation was
reviewed by Coats (1965), Thomson (1965), Johns
(1968) and Forbes & Preiss (1987). TL was considered
tobe of Mannoan age by Chomsou (1965). However,
the scdimentology of the Tent Pil Pormucion has
never been studied in detail und the stratiraphie
column of the South Tent Hill scetion by Thomson
(1965) 1s the only previous stltempl to mentily and
document the sedimentary structures. Joyo (1968)
considered ihat fhe suite oF sedimentary structures i

118 L.A, DYSON

: _ FINKE SPRINGS
. PUTTAPA GaP “3

STUART

‘@ BRACHINAGORGE © .
— * BUNYEROO GORGE ' .
. BL ACK GAP

oe -

NEW SOUTH WALES

0 LAKE a Sse
: GARDNER eet ‘ NI - | WARRAKIMBO GORGE “Ea

- ‘ 3 ° =U WAGGA BLUEF@, @ PARTACOONA =. 82 +, PROVINCE
aN UFR © MIDDLE GORGE i:

aN |. ADELAIDE
SOUTH TENTHILL © be GEOSYINGLINE:

‘ : PORT AUGUSTA \ “ont

NG . HIDDEN GORGE

:@ QODLA WIRRA |

KULPARA

\

h \
VADELAIDE
HALLETT COVEY .

OCHRE COVE WA ae
* Sé_waunt

4 O* TERRIBLE

VICTORIA

100
KILOMETRES

Fig, |. Tectono-sedimentary provinces in South Australia, showing localities of stratigraphic sections in the Adelaide
Geosyneline and their relation to other localities on the Stuart Shelf and in the Torrens Hinge Zone (afler Dyson 1995!).

TENT HILL FORMATION AND SIMMENS QUARTZITE WW

Fig. 2. View of the type section for the Tent Hill Formation
(Dalgarno et a/. 1968) on the southern face of South Teat
Hill.

Fig. 3, Stratigraphic log of the Simmens Quartzite und Tent
Hill Formation at South Tent Hill (after Dyson 1995!),

SOUTH TENT HILL

f mc ve
mud sand gravel

Simmens

Quarizite tidal shelf Regressive
Systems

Tract
(FRST)

Set’ Member a de

Lincoln Gap
Siltstone

Member Highstand
Systems
Tregolana Tract

Siltstone
Member

outer shelf

SANDISON SUBGROUP

=
oO
==
oO
&
oO
Lu
Pa
_—_—
Lom
eb)
—_

= —< parallel lamination ~€ hummocky cross-stratification
—<s\__ ripple cross~bedding “~<& _ swaley cross-stratification

\A/ trough cross—bedding ewe clasts

i 1. A. DYSON

METRES

SIMMENS QUARTZITE oo Horizontal bedding

TENT HILL FORMATION
Corraberra Sandstone Member
Lincoln Gap Siltstone Member Limit of outcrop

Tregolana Shale Member Measured Section

Geological boundary

bin. 4 Geological map of South Tear Hill

+
yao?
a

(LNT TILL FORMATION AND SIMMENS QUARTZITE

the Tene FOIL Kormation was jadicutive af sinong
Currentuction and shallow wiiter scdimentluten,

Siratiuraphy

The former Simmens Quurtzite Member of the
Tent Thi Bormation is raised hereiy to formation
status to refleet its regional significunee. ‘Together.
the Tent Hill Portration abd Simmens Quartzite (Pig.
3) represent regressive sedimentation of the Sandison
Subgroup on the Stuart Shelf The Sundison
Subyruuip is un uncontormity-bounded depositional
sequence in the sense of Mitchum (1977). Dolostone
of the underlying Nuccaleena Formation dees rot
crop oulin tbe Tent Thily buts present in deilcore
from the Stuart Shelf This commonly about 2-6 rn
Wick and displays a shirp to gradational base, The
Nuccaleena Formation was deposited below storm
wave base on a combined sequence bound-
ary/transeyressive suchace Uhal represents am hiatal
suilave of terigenous starvation, Phe lower Tent Hill
Pormuhon comprises the Tregolina Shae and
Lincoln Gip Siltstone members that represent the
Hiwhstund systenty tract of the Sandison Subgroup,
An uterpreted falling stage or forced) regressive
systems tricl (Dyson l99Ga). comprising the
overlying Corraberru Sundstune Member of the Tent
Hill Formation and the Simmens Quartzite, is placed

REGION
TYPE AREA

STUART
SHELF

South
Tent Hill

FLEURIEU
ARC

TORRENS
HINGE ZONE

DEPOT
SPRINGS
SUBGROUP

hiatus higlus hiatus

ARUHNA
SUBGROUP

Hallett Cove Kulpara =| Hidden Gorge

2)

between the highstind systems tract and the
sequence boundary at the top of the Sandison
Subgroup (Pig. 3).

The Sandison Subgroup on the Stuart Shelf and in
the Adelaide Goosyneline (Fig. 5) is unconformably
overlain by (he Wileolo Sandstone and. together wih
the Yarlow Shule, is herein assigmed to the Aruhia
Subgroup (Dyson 1996b), The Yarloo Shale is m turn
uncontormithly overkun by the Wearing Dolomite
und. logether with the Wonoka Formation, is
ussigied to we Depot Springs Subgroup (Dyson
1906h),

Tent Hill Formation

The Tent Hill bormation, abour 200 nm thek at
South Tent Hill, isan upwurd-sanding unit consisting
of the Pregolaia Shale Member the Lincol Gay
Siltstone Member and the Corraberra Sindstone
Member (Pig. 3). Tt ts gradationally overlain by the
Simmens Quartzite.

Degolana Shale Meniber

The Tregoluna Shale Member consists ol
laminated to thin-hedded. very fine to Hoegraied,
dark greyish brown sandstone iiterbedded with
greyish rad shale (Fig. 6). It is about 60 m thick.

NORTH
FLINDERS
RANGES

Finke Springs

SOUTH WEST
FLINDERS.
RANGES

CENTRAL
FLINDERS
RANGES

Bunyeroo
Gorge

NACKARA
ARC

Oodla Wirra

Wonoka
Formation

Wonaka
Formation

Wonaka
Formation

Wearing
Dolomite

Wearing
Dolomite

Wearing
Dolomite

hiolus

Bunyeroo

Bunyeroo
(i Formation

Formation

Bunyeroo
Formation

Wilcolo. Sst.| Wilcola Sst, Wilcolo Sst.

VV oF
ABC’ Range ABC Range
Quartzite Quartzite

Simmens
Quartzite

Vw

Corraberra Corraberra
Sandstone Sandsione
Member Member
Moorillah Moorillah

Sillstone Sillsione
ember Member

Corraberra

Sandstore
_Member
Lincoln Gap

Sandstona
ember

Brachina Fm,
Brachina Fm,

voy ‘ABC
Range
Quarizite

ABC Range
Quortaile

Bayley
Range
Sillst. br.
Moorilloh |

Sillstone
Member

We Range
ABC Range Quarizite

Quarizite

Moorillat
Sillstone
Member

Ulupa
Siltstone

Ulupa
Slitstone

Moglooloo
Sillstone
Member

Tant Hill Formation

Tregolana i
Shale
Member

Siltstone
ember

Neaicaloo
M

Meaioalue Moolon|no
Sillstone iitstene
Member ember

Brachina Fm,

Brachina Fm.

SANDISON SUBGROUP

Seoclilf

h- pullers Sandslone

Nuccaleena
Formation

cL
a)
oO
oe
©
<
Zz
Lil
Oo
=
=

UMBERATANA
GROUP

Nuccaleena
Formation

Seacliff Sst.

Seoclif{ Sst_

Nuccaleena
Farmation
yo

Nuccaleena
Formation

Nuccaleena
Formation

Fig, 5. Nomenelitture aud correlation of the Samdisoo Subseaup (alter Dysan 1995 ')

)32 LA

hig. 6. Shale and very thin te thin-bedded fite-grained
sandstone, Trepoluia Shale Member of the Tent Hill
Formation,

Individual Sandstone beds are flat-based with
occasional grooves and scratches. Internally, the
sundstones display parallel lamination and current
ripple cross-lamination analogous to the Bouma
sequence for turbidites and are interpreted as having
been deposited at or below storm wave base from
waning. unidirectional currents of storm. origin.
Siltstone beds are characterised by parallel
lamination, The thickness and frequency of the
sundstone beds increases up-section as the Tregolana
Shale Member grades into the Lincoln Gap Siltstone
Member. The Tregolana Shale Member was
deposited below storm wave base and is a@ lateral
equivalentof the Moolooloo Sillstone Member of the
Brachina Formation,

Lincoln Gay Silttstone Member (new nqme)

The name for this new member of the Tent Hill
Formation is derived from “Lincoln Gap". 24 km
south of South Tent Hill. It resurrects. in putt, the
fomer “Lincoln Gap Flagstones” of Miles (1955)
That was previously used to include the siandy
succession above the Tregolana Shale. Here, the
Lincoln Gap Siltstone Member is used to deseribe
the lower half of the Corraberra Sandstone Member
that was originally defined by Thomson (1965), Lt is
about 40 in thick and consists of interbedded greyish
red shile and thin to medium-bedded, fine-grained
greyish brown sandstone. Flute casts and scratch
marks are common at the base of sandstone beds
suggesting current transport to the east (Pig. 7).
Internally, the sandstones commonly display
horizontal planar lamination. They are. in places,
capped by interference ripples or asymmetrical
tipples with sinuous. crests. Sundstones displaying
planar lamination or hummocky cross-stratification
(HCS) are capped by near-symmetrical ripples
Crests of the nearsymmetrical ripples show a

DYSON

FLUTE CASTS

WAVE RIPPLE CRESTS

Vig. 7. Palacocurrent data for the Lincoln Gap Siltgtone
Member

hexugonal pattern or are straight to wavy with
tuning-fork bifureations.

Planir-laminated sandstone beds cupped by current
ripples are interpreted as Bouma BC sequences and
suggest deposition at or below storm wave base in it
current-dominated environment, The [Mute cists
suggest the influence of unidirectional currents that
were directed off-shore (Fig. 7). Ripple marks on top
of these sandstone beds indicate the influence of
unidirectional and oscillatory currents, resulting
combined-flow ripples (Fig. 8). The presence of

TENT HILL FORMATION AND SIMMENS QUARTZITE 123

Fig, 8, Combined-flow ripples on top of fine-grained sand-
stone, Lincoln Gap Siltstone Member. The ripple crests

show imperfect bifurcation and note the presence of

wonkle marks.

HCS and planar lamination indicates that storms
were responsible for the generation of both
unidirectional and = oscillatory currents, Near-
symmetrical ripples are interpreted as wave-formed
in origin. The orientation of wave ripples suggests a
north-south palaeoshoreline. These structures
suggest deposition above storm wave base but below

Fig. 9. Chocolate-brown sandstone of the Corraberra
Sandstone Member in the centre-foregound, gradation-
ally overlain by quartzarenites of the Simmens Quartzite.

fairweather wave base. The Lincoln Gap Siltstone
Member represents deposition in an environment
where oscillatory currents were dominant over
unidirectional currents. [It is a lateral equivalent of
the Moorillah Siltstone Member of the Brachina
Formation and is sharply overlain by the Corraberra
Sandstone Member,

Corraberra Sandstone Member

The Corraberra Sandstone Member is about 25 m
thick and consists of greyish red, iron-stained, fine to
medium-grained sandstone (Fig. 9) interbedded with
greyish brown shale. The sandstone beds are
commonly micaceous and display heavy mineral
lamination, swaley cross-stratification (Fig, 10),
quasi-planar lamination (Fig. 11) and medium-scale
cross-bedding. They are, in places, capped by
symmetrical and asymmetrical ripples, interpreted as
wave and current ripples, respectively. Glauconite,
intraformational mud clasts, mud drapes, foreset
bundles, climbing ripple cross-lamination and
herringbone cross-lamination are also present.
Several upward-sanding cycles, commonly about 5
m thick, are present in the Corraberra Sandstone
Member where swaley cross-stratified sandstone
beds are commonly erosive into underlying cycles
that comprise tidal sand sheets. The Corraberra
Sandstone Member grades upward into the Simmens
Quartzite (Figs 3,9).

The lamination and cross-stratification styles
within the Corraberra Sandstone Member suggest
initial deposition at or above fairweather wave base
in a shoreface environment where oscillatory-
dominant storm currents were operative. Cross-
bedding near the top of the unit suggests the
increasing influence of tidal currents. A tdally-
influenced marine environment is also indicated by
the presence of glauconite, bipolar cross-lamination
and foreset bundles. The disconformity at the base of
the Corraberra Sandstone Member marks the onset

Fig. 10. Medium-grained sandstone of the Corraberra
Sandstone Member displaying SCS. Hammer for scale.

124 1. A. DYSON

Pig, 11. Quasi-planar lanination i medium-prained sand
stone, Corraberrn Sandstone Member, Note preserve of
loweangle cross-bedding showing palieotlow to the
right, Lens cup is 52 min in diameter

of forced regression in the Sandison Subgroup and
metre-(hick, swaley cross-stratified sandstone beds
ie interpreted as forced regressive deposits (Dyson
1995119960). Upward-shillowlig cycles ure
Interpreted ws parasequenees in (he terminology of
Van Wagoner (P985), On the Stuart Shelf the
Corruberra Sundstane Meniber represents partial
lateral equivatents of the lower ABC Range Quartzite
and upper Brachina Pormation troy the Adelaide
Geosyiicling, The ironerich facies are Similar to
cyuivulents thal crop aul east of the Torrens Hinge
Zone at Kulpara, Ochre Cove, Hallett Cove and
Puthipa Gap (Big. 1). 1h is thought (hat the iren was
derived from erosion of the Mesuproterozoic
Pandora Formation on the Gawler Craton.

Simmens Quartzite

The Simmens Quartzite is about 100 m thick and
consists of prey lo greyishewhite. fine to medium
eruined. thin to very thick-bedded quartzarenite.
These beds contain various clasts of voleanic and
grinilic composition, varying in size up to 20 mm,
Compound cross-bedded sets comprisiny herringbone
cross-stratification, sigmoidal cross-bedding displaying
foresel bundles, planar-tabulur cross-bedding, shale
clasts, horizontal-planar lamination and tinor swaley
cross-stratificadion (SCS) are abundant (ries 12, 13).
Large-scale, dough cross-bedding is common (Fig. 14),

Compound cross-bedded sandstone suggests
deposition of sand waves ti a tide-dominitted
environment Where asyiimetry of the dominant and
subordinate currents was pronounced, but bipolar
currents were significant, The lack of 1ICS and SCS.
except near (he base of the Simmens Quartzite
suggesis shoreface deposition above luirweather
wave base where tidal currents were dominant

Fig. 12. Quartaavenite of the Simmens Quariaite display ing
overtinned cross-hedding of tidal origin and SCS, Note
sinalleseale herringbone eross-bedding at base al bed
Brunton compuss tor scale

Fig. 13, Compound enoss-hedding i quartzurenite al! the
Simmens Quartile

Nig. (4. Rough eross-hedding in the Simmens Qaarvate.
Lens cap in 52 mm in diameter.

PENT HILL, FORMATION AND SIMMENS QUARTZITE 25

Palaeocurrent dala show a strong trend towards the

east and are interpreted as reflecting progradition of

ebb-flow tidal sutid sheets (Pig, 15). Following Couts
(1965). (he Simmens Quartaile is correlated with the
ABC Range Quartzite in the Adelaide Geosyncline.

Sequence boundary

The upper boundary of the Simmens Quuartzite
does not crop outat Sourk Tent Hill However, south
of Bill's Lookout near the north-western side of Lake
Torrens, an erosively-hased sandstone at the top of
the Simmens Quartzite crops out poorly where i is
contormably overlain by moderate brown lo greyisli
green shale of the Yarloo Shale. The metre-thick,
mediun-eraincd off-white sandstone contains basal
clasts of shale, lithic sandstone und occasional
volcanics, Sedimentary structures include trough
vross-bedding, SCS and symmetrical ripples.

The erosively-based sundstane at the top of the
Simmens Quarizite is interpreted as huving been
deposited ian estuarine environment. The style ar
ceross-bedding in the lower purl of the oil suggests
deposition on a fuvially-dominated shorevuce, A
stormy influence is indicated by the presenee of SCS

CROSS—BEDDING

n = 120

CURRENT LINEATION

Fig, 15. Palaeocurrent data for the Simimens Quartaite,

and the symmetnical ripples are interpreted as having
been formed by wave action, This unit is 4 possible
equivwtent of the Wilcolo Sandstone that overlies the
ABC Range Quartzite in the Adelaide Geosyneline
where it murks the development of broad (c. 1)-20
km) ine sed valley fills that in) places atin a
thiekness of some 25-50 m ip outcrop, They consist
of a basal, trough cross-hedded facies of thuvial
origin, overlain by SCS shorelace sands, The SCS
shoreface sands pass rapidly upward inte. basinal
shale of the Bunyveroo Formation. The Wilculo
Sandstone and Bunyeroo Formation together
constitule the Aruhna Subgroup (Dyson 1996b), Its
upper boundary is represented by the maximum
flooding surface ul the base of the Wearing Dolonite
Which is coincident with the development ot
kilometre-deep canyons previously assigned to the
overlying Wonoka Formation (Dyson 1995°, 1996b),

Stratigraphic equivalents of the Tent Hill
Vormation and Simmens Quartzite

The redefined Tent Hill Formation and Simmens
Quiurtzite uny be considered partial lateral
equivalents of the Brachina Formation. ABC Range
Quirtvite and Ulupa Silistone in the Adelaide
Geosyneline (Fig, 5), The Brichina Forinution was
delined hy Dalgarno & Johnson (1964) as the thick
sucvession of siltstone conformibly overlying the
Nuccaleena Formation and passing upwards mto the
ABC Range Quartzite (Mawson 1939). Together
wilh the Seaclit? Sandstone and Nuccaleena
Fornration. the Brachina Formation und ABC Ringe
Quativale were incorporated into the Sandison
Subgroup (Dyson 1992). The Brachina Formation
and ABC Range Quartzite crop out al several
localities. wilhin the Mount Lolly and Flinders
niuges, of which the latter occurrences uppeur lo
display lateral continuity. A study by Dyson (1992,
1995!) focused on well-exposed sections at Hallet
Cove. Ochre Cove, Mount Terrible, Kulpara, Hidden
Gorge, Wyaccu Bluff, Purtacoona ant Bunyeroo
Gorge (Fig. 1). The Ulupa Siltstone is best developed
to the east und northeast parts of the Adelaide
Geosyneline on the BURRA. COPLEY, OLARY,
ORROROO und MARREE 1:250 000 geological
sheets, The Brachina Formation and overlying ABC
Range Quartzite constitute an overall upward-
sanding succession, They cepresent regressive
sedimemation of the Sandison Subgroup aed are
therefore defined us all the strata overlying the
maximum flooding surface/downlap surface.
represented by the Noecaleena Formation. to the
sequence boundary at the base of the Arubna
Subgroup.

126 1 DYSON

Prachi Corneaiion

The Bruching Pormuion comprises four members,
cach ot whieh can he defined by [ts (jthafaciwes (Fig.
53) TWwe of these, tamely the Bayley Runge Siltstone
Mewiber aid the Corniberry Sandstone Member. ane
consmlered literal equivalents (Dyson LYYs4. The
Bayley Runge Silistone Member does not crap out at
Hallew Cove ov Kulpara (Fiz. 1), The Coreaberra
Sandstone Merpber of the Tent Gill Formation that
crops out onthe Stuart Shell hasan equivalent that is
Included in the Bravhion Formation at Halle Cove
(Dyson 19054). ‘Phe lithofieres al the Brachina
Formation ut Hallett Cove displiy a sui af
sedimentary structures Suggestive ol depasttion
uoder the influence of greal storms. These sirucnires.
Invading weonompanying pulaeweurment dak, ane
ilfustrated and deseribed tore Tully by Dyson
(1995! 1995).

ABC Range Guarizile

The ABC Range Quartvate is composed of pale
piikish wrey to greyish white, thick-bedded to
massive, fine to mediin-gralned, slightly feldspathic
quatzale sind ino interbedded shale and) fine
granmicd sundstone, It is characterised ty the
abundance of planartabular, herrighone and laree-
scale Trough eross-bedding. usymimetricy) cripples
with sinuous crests, Maser and lenuculur bedding,
muderacks, mud drapes and mud intrachaists.
Straigiil-crested symmetrical tipples also cup seme
sandstone beds, Compound cross-hedded sets consist
ul small-scale cross-bedding separated by ipelined,
master set boundaries, As the inclination of the
niaster bedding plines decreases, the bipolariry of
Ihe cross-hedding becomes nore prevalent
Megaripple cross-hedded sets, commonly uni-
direchonal display o thin-thick alternation of sanely
foréset bundles, bounded by ceachwation surtiaces, mid
couplets and/or mud drupes. Sandstone ehannels are
characterised hy genrly jnelined lateral aeeretion
Surfaces and low-angle truncation surlaees. SCS and
herizontal-planue laminae are eecusionally
Observed neur the base oF some minor quartyite
units FIOS is restricted to iitervals ob interbedded
shale and thin lo medium-bedded line sandstone and
quartzite, Thus, the suite of sedimentary structures in
Ihe ABC Range Quarizite deseribed here suggest.
(hal ii was deposited ina shallow mue cavironnment
where tidal currents were dominant over storm and
Wave aenion,

Ueapred Si 8tonte:

The Ulipa Siltstane is a regionally significant unit
that can be mupped over w large urea of the Adeluice
Geosyncline, The Lype section of the Ulupa Siltstone
was defined hy Miruns (1964) near Mount Bryan on
BURRA where it was described as a succession of

green, grey dnd locilly purple shales, Phimmer
(lY7S) pecounised the three members of the Brachinu
Fornativn in he Ulupa Siltstone at Qudla Wit
(Pigs |. 5) and) recommended the term ~Uhipu
Silane’ he relinquished. Towever, Porthos d& Preis
(1987) proposed retention ob the stratigraphic nume
Becsuse they aeued (hal reainnal muppability of the
constiuent members hid por been established
Cilhofieies of the Utupa Siltstone are disuil
equivalents of the constituent members of the
Brivhiog Vormation and are therefore tire
Ininsgressive, The Ulupa Salisfane represens
jeeressive shallow purine sedimentation on the
middle to outer shell, Palie@ocurrent data show 4
Wide range al current directions (Dyson (995),
sUgesins in environment where unidirectional aru
dscillory currents were interactive.

Depositional model tor the Tent Hill Vormation
and Simmmens Quartzite

The Simmens Quarizite und constituent mernbers
of the Tent Hill Formation may be defined hy: heir
lifhotages. Phe Tent HE Pormation is an upwarnd-
sanding succession of interbedded shale and tine-
grained sandstone, Sandstone beds display several
scdimentary struetires associated with stort
deposition such as BC Bouma sequences, HCS,
Micro-HCS and quasi-planar lamination, ‘The
Trevolana Shale Member way deposited below stun
wave buse. With progressive shallowing ahove ston)
wive hase, interbedded shale and sandstone of the
Lincohy Gap Siltstone Member were deposited, A
the top of this succession, the Corraberry Sandstone
Member consists of several sharp-bused shoreface
siidstunes about 1-2 in thick that display SCS,
These SCS sandstones are ceferced to as attached
shoreface deposits, Some SCS sandytunes ane
completely enclosed withinshale and we reterred 1
ux deiachec shoreface deposits. The huse of wae
SCS sandstone is a high-lrequeney sequence
houndiry. The ¢rosive shomfure depasus are
interpreted is forced repressive deposits The
Conuberns Sandstone Meriber was deposited above
falrweather wave buse in-an environment that was
stormedominated bul where tidal activiry was alsu
signifieunml, Th is gradationally overlain by pura
sequences of tide-dommnuted quartvurenites anid
lithic sandstones of the Simmens Quuartvite. The
Siimmens Quurtzite represents continued shallow ie
af the sea in which the Tent Hill and Brachina
Formations were deposited, Combined with a high
sediment supply, prowridation of (he lower shoreface
resulied in a wide, shallow shell whieh was
conducive ly tidal amplification. No submurine tan

deposits Have been revognised ait this Sadvraphir

level elsewhere in the busin.

TENT HILL, PORMATION AND SIMMENS QUARTZLTE a

The ‘Tent Hill Formation and Sinmens Quartzile
represent progradation ofa tide-dominated shoreliace
Wok starin-dominated, shallow shel? environment.

This regressive Succession is composed OF number

of upward>sanding eyeles fat ane conained within a
hierumehy of transgressive-regressive eycles, Tivse
cycles are inithuly aggradational mm character and
become Mmeresingly proghadational apsecnon. A
very thick afflup wedge developed fron) this
prowradkation) aid eravitational instability resultedt in
exfensionil faulting and enhaneed subsidence,
Palacocurrent dit suggest a north-sonth, “dally
Influenced shoreline, Sediment was derived fron the
GawlerCraton 1 the west ind tainsparted eastwards.
‘The depositional environment shalloweed to hove
lairwealher wave base across a telatively narrow
shell and sediment prograded geross the Torrens
Hinge Zone into the Adelaide Geosyncline. Here,
thickiess oF he Sajidison Subgroup Was affeeted hy:
syn-deposilinnal teclonies, Coats (1965) sugeestedt
thar the overlying Wonoka Formation amd Pownd
Suberoup were nat deposited op the Stuart Suet! bur
were restricted to the Adeliide Cieosyneline hecuuse
of syredepositonal faultine actos the Tortens Hinge
Zone, Vhe ureugle end represented by the “Correos
Hinge Zone marked the possible elze of the tormer
Shelf during deposifian of the lower Wonuka
Normation, However, identilteation of the Wanaka
Formation uf dnilcore from the caster Stuart Shelt
(eg. Bopeechee 2) sugeests thal it mayor trans-
oression ovcurred weross rhe Torrens Hinge Zone at
{his tine, Depositin of thick Bunyeroo sediments in
the Adelaide Geosviclineg Was contemporaneous
with delve subsidence. Rewronal instability
contributed (0 the incision of Wonukiu canyons on the
weater edge of the Geosyncline.

Disewssion

Torovd regressive deposits

I the chissig Exxon Sequenee stritigniphic model,
the ‘Type 1 depositoual sequence consests ol
lowstund, traiseressive aod Highstaod systems. tracts
whieh are schemartcally fled to specific Trerements
Of the custatie curve, However, ali docreasiige Vollime
OF lifcraiure sageests (halt deposition during a relative
fall insea level may be placed into a fourth systems
tract between the highstand systems tract (HST) and
the sequence boundary. This systems tract has been
previously yelerred (6 us the falling Stage or lorced
regressive systens tel (ca. Hunt & Tucker 1992).
Study Of progradiationsl tidal sand sheets and: sharps
hased shorelace deposits of the Tent Hill Formation
and Simmens Quartvite at South Tent Hill. and
similar deposits of he Brachina Pormauon und ABC
Runge Quartvite at Halletr Cove. Kulpara, Bunyeroo
Gorge and Trebileoek Gap (Fig. 1) sugacsts that they

may be wssipned to the falling sluge systetis fact
(Dysvo [996a). These Units represent pegresswe
sedimentation of the Sumdson Subernip on the
Stuart Shell aod a the Adchade Guasynctine The
lower dnd upper boundaries of the the Lalling stage
systems Wact (PSST) qin fixed on the neliive: seu
level curve, However the facrements of the other
svarema tracts dre nol Axed ane will vary due ie
subsidenve rae and sediinenc supply. Sharebee
sandstone displaying SCA ait the buse of the
Corraberra Sandstone Member corresponds uy the
base of the PSST. Ts apper boulidary is (hie sequence
huundary which is defined heneas the lowest parma
relative sea level. Vhe correlivive cunloriity may he
UN OgOHs tO the Gown lip sure ar discon Fornyiyy st
the lop oF stibmarine fans in earher Exxon models, 1
pusses updipr inte the subuerial uncentornily
ussdenited with the sequenve houndury,

Self dyiindes cel path eeaehrrones

Shoreface stev'ii depostis of the Cnrtaherny
Sindstone Member were possibly deposited ina
mesotidal environment with tidal ninge of seme 2
Som. Such enviriiinents abe Stora; ar wave-
dominuted (Dalrymple 1992) The Crrraterra
Sanuletoue Member is direethy Sueceeded by tial
sind sheal deposits of lhe ABC Range Quartaite of
the southern purt af the Adelaide Geasyneline This
sugwests that dial overpruting af sturm and wave
effects extended well Out veross the upper stioreface.
The roliative influence of storms deereased as the
fidal current speeds toereused. se thal distal parts ef
the sand sheet coutitined storm-penenmted structures.
The stornm-donnnated shorehuce system was replaced
by progriding Gde-dominuwed dels and open coast
Lidal fats. Mural wars deposited beyond the aleptly and
ringe ol tidal reworking. “Tidal channels within the
Sioimens Quurtzite were possibly jucised to shallow
subudal depths on the shoreluce. bused on the varity
ay SCS. The depth of incision Suggests oo high
mesonidal to possible daerofidal tange along the
palaeoshoreline. During deposition of the FSST, (idal
cunge miy have been limited dire to the celative fall
in sea level.

Wave ripple orfcntations suggest a regional aorth-
souub shoreline. Clastie muterial was sourced trom
the West. Asymmetry of the tidal regime is supported
by the dominant dminiodal trend for tipple etoss-
bedding, The wide spread ol these dita suggests Ui
longshoare currents were operulive, Shoaling
lauirweather aod stort waves initiated longshore
currents that transported sumd on the shore
rouvhly parallel to the shorebne. Pulaeocurrent ditt
feom storm-influenced litholacies in the Brachina
Formation at Hallett Cove are described by Dysoy
(1995! 1985),

128 I.

Litholacies relationships and correlations

The Corraberra Sandstone Member of the Tent Hill
Formmilion at South Tent Hill and oo the Stuart Shell
comprises the lower part of the sandy succession
overlying the Tregolana Shale Member (Crayford
L964), Litholucies resembling these of the
Corraberra Sandstone Member also crop out at
Qchre Cove (Dyson 1995!) Hallett Cove and
Kulpara (Dyson 1992) and at Puttapa Gap near
Beltana (Coats 1965), Granular, medium to course-
grained sundstone and dark red to reddish brown
shale at Ochre Cove (Pig. J), previously identified as
Bunyeroo Formation (Dyson 1993) was
reinterpreted us lithofacies of the Corraberru
Sundstone Member of the Brachina Formation
(Dyson 1995!) At Kulpara, favies of the Corraberra
Sandstone Member ure interbedded with the lower
ABC Range Quartvite, The Bayley Range Siltstone
Member crops oul north of Picht Richi Pass at
Middle Gorge, Partacoona, Warrakimbo Gorge.
Black Gap. Bunyeroo Gorge. Brachina Gorge and
Finke Springs (Mig. 1). South of Partacoona and
adjucent to the Torrens Hinge Zone, the storie
domimted fees of the Corriberra Sundstone
Member ure predominant, ‘This suggesis that the
Corraberra Sandstone and Bayley Range Siltstone
members, both deposited above fairweather wave
base, are lateral equivalents. Furthermore, they are
partial hteral equivalents of the lower ABC Range
Quartaile,

4. DYSON

The terms “Simmens Quartvite’ and “ABC Range
Quucizite” are used herein to deseribe silica-
cemented quartvite or orthoquartzite in which the
dominant mineralogy is over 909 quartz, Petlijohn
er al (1972) prefer the use of the tern
“quartzarenite” over “orthoquartzite™ for those
sediments m which the detrital traction is 95% or
more quartz, The ABC Range Quartzite has been
mapped at the first appearance of thick, lalerally
extensive white quartzite (e.g., Webb & von der
Borch 1962; Dalgarno & Johnsen 1966). Apparent
imertonguing of the ‘Tent Hill Formation and
Simmens Quartzite on the Stuart Shell, and between
the Brachina Formation and ABC Range Quartzite in
the Adelaide Geosyneline cun be generated by the
stacking of these lithofucies.on a purasequence scale,
Similarly, intertonguing between the constituent
inembers of the Brachina Formation and Tent Hill
Formation may be explained in this manner

Acknowledgments

The study of the Tent Hill Formation and Simmens
Quartzite comprised part of a PhD dissertation by the
author at Flinders University. Wolfgang Preiss and in
unonymous reviewer read (he manuscript critically
and contributed many helpful) suggestions. Gail

Jackson and Ghazi Keaishnan drafted the figures

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Stratigraphy - Applications to Hydrocarbon
Exploration.” Amer. Assoc. Petrol. Geol. Mem. 26, 205-
212.

PETTUOHN, F. J., Porrer, P. E. & Stever, R. (1972) “Sand
and Sandstone” (Springer-Verlag, New York).

PLumMMRR, P. S. (1978) Stratigraphy of the lower Wilpena
Group (late Precambrian), Flinders Ranges, South
Australia. Trans. R. Soc. S. Aust. 102, 25-38.

Tuomson, B. P. (1965) Erosional features of the Tent Hill
Formation. Quart. Geol. Notes, Geol. Sury. S. Aust. 13, 4-
3

VAN WaGconeR, J. C. (1985) Reservoir facies distribution as
controlled by sea-level change. Abstract and poster
session, SEPM mid-year meeting, Golden, Colorado, 91-
92.

Wess, B, P. & VON DER Borcu, C. C. (1962) WILLOCHRA
map sheet, Geological Atlas of South Australia, 1:63 360
series (Geol. Surv. S. Aust., Adelaide).

TRANSACTIONS OF THE

ROYAL SOCIETY
OF SOUTH AUSTRALIA

INCORPORATED

VOL, 120, PART 4

BIOLOGY OF THE EUCALYPT GALL-FORMING FLY,
FERGUSONINA FLAVICORNIS MALLOCH (DIPTERA:
FERGUSONINIDAE) AND ITS ASSOCIATED
HYMENOPTERANS IN SOUTH AUSTRALIA, WITH A
DESCRIPTION OF A NEW SPECIES OF BRACON
(HYMENOPTERA: BRACONIDAE)

By G. S. Taytor, A. D. AusTIN & K. A. Davies’

Summary

Taylor, G. S., Austin, A. D. & Davies, K. A. (1996) Biology of the eucalypt gall-
forming fly Fergusonina flavicornis Malloch (Diptera: Fergusoninidae) and its
associated hymenopterans in South Australia, with a description of a new species of
Bracon (Hymenoptera: Braconidae). Trans. R. Soc. S. Aust. 120(4), 131-146, 29
November, 1996.

Galls initiated by the association of Fergusonina flavicornis Malloch (Insecta:
Diptera: Fergusoninidae) and Fergusobia sp. (Nematoda: Sphaerulariidae) on
Eucalyptus camaldulensis Dehnh. at Goolwa, South Australia were monitored during
a single, late summer generation for emergence of flies and associated Hymenoptera.
The morphology of the galls is described for the first time. A total of 12
hymenopteran species was reared from galls, twice the number previously recorded
from Fergusonina/Fergusobia galls. There was a strong positive correlation between
emergence of flies, combined totals of flies and wasps and gall size. Bracon
fergusoninus sp. nov., a probable primary parasitoid of F. flavicornis larvae, is
described. Notes on the biology and taxonomy of each hymenopteran species are
presented along with an illustrated key to their identification.

Key Words: Eucalyptus, gall-forming, Fergusonina, Diptera, Fergusoninidae,
parasitic Hymenoptera, Braconidae, Bracon, Fergusobia, Nematoda, Sphaerulariidae.

Transactions uf the Rowel Seceteabs. Aust W996, 101 131-16.

BIOLOGY OF THE EUCALYPT GALL-FORMING FLY, FERGUSONINA
FLAVICORNIS MALLOCH (DIPTERA: FERGUSONINIDAE) AND ITS ASSOCIATED
HYMENOPTERANS IN SOUTH AUSTRALIA, WITH A DESCRIPTION OF A NEW

SPECIES OF BRACON (HYMENOPTERA: BRACONIDAE)

by G. S. Tavcor. A.D, Austin & K. A. Davies!

Summary

Tayo, O.S.. Austin, ALD. & Davies. KR, A (1996) Biology of jhe etedypr gall-forming Aye Fergus
fluvicornis Malloch (Diptera: Mergusoninidie) and iy assocated Ay mencpterans i South Austrabia. with a
description of a new species of Brace (Hymenoptera Beaconidue). drain it, Sac. 8. Avast. 12004). 131-146, 29
November, 1996,

Calls initiateal by the asseciahon ol Fereusonind avicgends Malloch (Unseeta; Diptera; Ferzusonmidae) and
Fergwobia sp. (Nematoda: Sphacrulanidae) an Bicelyptes comuiideiy Delnh.at Goolwa. South Australia
Were Modoc duving a single, late summer generation for enivrecnee al flies and assuviued Hymenoptera.
The morphology of (he galls is deseabed forthe frstiime. A fowlot 12 hymenoptenin species was reared from
sully. (wie (he number previously recorded (ron, beryuyorind/Renawobia walls There wads a strong positive
correlate bernveen emergence of Ties, combined totals of Tigs andowasps and gall size, Brecon fergawertins
sp nu\. a probable primary purasitond at & fawicventy larvae, iy deseribed, Nores onthe biology und Gesonony

of dich hymenopterin speciosdre presented along with an dlusirated key tether meanticunan

Key Woks:

Aiealipins, wallslorming, Ferswserine, Diptera. Gerausoninidie, parasitic Hymenoptera,
M e ° f ] — d

Braconidae, Kroc Mergisebia. Nematoda, Sphieritfariidie,

Introduction

The biology of gall-issoeiated Hymenoptera is
fundamentally complex and it is offen extremely
difficult to determine the tue relationships of species
(Boutek 1988: Gough & MeMahon 1988: Naumann
M91: Schénrogee er a/, 1996), For mstanee, galls
inmuy contam a range of biolowigally different
Hymenoptera, viz, primary parasitoids. facullaive
and obligute fyperparasitoids and imquilines. The
biology of the Latter group is particularly. difficult to

uoravel. Inquilines are species thal five mide galls a

other vecidogenie insects und are phylophugous for
all or some ol their larval development. They often
kill the primary gall-forming species atin early stige
of pall development and then increase (he size of the
oall, thereby making iLdilficult to determine whether
ag not the species concerned is indeed an inquiline or
the primary gall-former, Other inyuilines apparently
invade walls und feed Gn their tissues witheut
disrupting the primary cecidagente species,

One of the post complea gall assuciitions known
18 thal between Avreaseaine spp, (insect; Diptera:
Fergusoninidue) and Fergasebia spp. (Nematoda:
Tylenebida: Sphaerularitdae) in gulls on Myrtaceae
(Fisher & Nickle 1968; Harris 1982: Davies & Liyyd
1996), However. the only detailed study of the

| Department of Crop Protection, Waite Cumpus The
University of Adehide PMB 1 Cen Osniond Ss A\iist
506d

biology and phenology of Fereusening Malloch flies
is that of Currie (1947). whoa comunented on tte
possible pole of bymenopterans and other gall
myuilings i the seasonal (luetuarans of Thy
populations: This study showed that qumerots wisps
were associated with Ferensenine/Pereusohia lower
bud galls. Although Currie (1937) exanimed the
biology of this system, the uxonomic knowledge at
the time was limited and the biology of the species
involved was nol well understood.

This paper reports on uw study of the loa! galls of
Fereusonind flavieariiis Malloch associated with an
undescribed species of Fergusefid Curtie and the
associated guild of hymenopterans from gully on
Aucalyptiy camatdulenses Dehnh. at Goolwa, South
Austenlia Information 1% presented on gall
morphology, the phetiology of the fly and wasps, the
probable biolagy of the bymenopterin species
involved, their taxonomy, and ure relationship
between numbers of flies and wasps and wall size. A
new species of Bravon F. thought to be a primary
parasitoid of Fergusenita. is deseribed und an
iNustrated key to the 12 vallassoented hymenop-
lerinis is presented, ‘This irformation is compared
with Currie’s ()9R87) study and ts discussed iy
relation tothe inherent difficulties tn determining the
biology of all associated wasps,

Materials and Methods

Seven mature galls on an ornamental, +m high
cameldulensis sapling at Goohwa, (A5°3h'" S,

|32 Gos TAYLOR, AD AUSTIN & IA. DAVIES

134°46/ E) South Australia were caged wilh muslin
in-early March. 1995, preceding emerence of adult
fh flivicornis and other gall oceupants, and
periodically monitored until afler senescence of the
galland when Mies and wasps ceased lo emerge, Any
insects Lhat had emerged al each sampling time were
colleeted, identified and counted, Gully were dried
and weighed to compare gall mass with total insect
cihergences, Por observations on gull deyelopment
und morphology. young Iresh galls were sliced open
m water by a series of muisverse cuts, Abbreviations
used are: ANIC (Australian National Thseet
Collection, CSIRO, Canberra) and WARE (Duncan
Swain Insect Collection. Waite Cumpus. The
University of Adelaide), Voucher maternal of &
flavicernty has been lodged at ANIC und WARL und
material of ull hymenopteran species is Ueposived jn
WARE,

Results and Observations

Disrrthurion itd hase plant association

hersuyenina flavicernis, was the only dipleran
reared from un ormumentul specimen of &,
conkddulensiy ac Goolwa. Steir gulls bwve heen
collected by us from several other South Australiin
localities from naturally-oceurring E. ceaieldalensis
near Verdun and near Milang, and from this sume
host in the Adelaide suburbs of Hyule Park. Rose
Purk, Tusmore and Urrbrae. Ferguvenina flavicarnis
was described trom a single femile specimen frons
Sydney, NSW (Malloch 1925), Larvae. males and
galls have remained undeseribed to date. This is the
first host plant record for the species and first
description Ol the will (see below). We hive alse
collected all life history stages of AO flevicerniy ane
the associated, undescribed species al’ /ereusabra
whe plan to discuss these mia separate publication.

Gall morphology aid Jarnuition

The galls of A flavieornis are formed from
terminal teal buds, mature galls (Fig. 1) beme
bulbous structures, mostly ove an longitudinal
eross section, 18-50 mm in length by 12-20 nim in
diameter. Mean dry weight of mature galls was 2.22
+ 1.45 2 (range 0.89-5.16 2, 2 = 7). Galls appear to
consist of a single leat bud comprising, a number of
leaves, with the vente) surface of the oulerniost
forming the external face of the gall, The growth of
these outer leaves sometimes continues unalfeeted
beyond the gall proper, either running wong (he side
ofthe gall or emerging as normal leaf Gssue beyeud
its apex, The stem suppordag the gull usually
develops a bend just before its iaseruion in the base
of the gall. Transverse sections showed that the
galled tissui@ 1s Soft, Consisting OF purenchynatats
tissue, and thar A flavieernty larvae develop in

hserele cavilies, These are oval din longitudiial
oucine and distributed throughout the gall The
cavines are lined with plant cells which ae paler in
colour than the parenchyma cells between them.
Aerguyebia nematodes are found in the cavities with
the fy larvaies

bor the generation of /. flavicarnis which began in
late March, small, recently initiated galls were
observed, Four of (hese were collected und dixsecied
and the development of others was monitored during
the study. Pull gull size was. reached on the uree in
(wo to four weeks from pull initiation, The followin
observalions were made en the dissected galls. The
sroallest gall Ro mm in length contained 20 spindle-
shaped AL fhaicoriy cogs (Pi. 2) within whieh
developing lurvae could be seen. Tt contained many
parthenogenetic and juvenile nematodes and some
nematode eges. Al this stage of development na
eavilies were present, Another, 10) min long,
CoimmTcU sever) ees, a few first instar larvae OF
Fereivenina and many parthetiagehetic nenatodes
id eves, Some My kiryae were found within
cavilles, The small cavtlies being ringed with many
white cells, A 12 mm Jong gall contained Hirst and
secund iastur larvae of Ferenvonie, ull within
cavities, and muny parthenogenetic nemiutodes abe
cues. The lumest gall about 30 mn in length,
conlainedt first and second instar thy larvae within

Fiz. |. Hapvttial
(iii opi Feronsobia on Kucadyatus vronaladntensis at
¢

Cian, South Adsterdnn Seale bat = 28 on

Termin! deal bud gnll ot

BIOLOGY OF FERGUSONINA ANDY TTS ASSOCIATED HYMENOPTERA iat

Fie2. Bey ol Fergsoning favicoois from a terminal leat
bud gall from Eueelypruy cormealdulensty wl Goolwa,
South Australi, Seale bar = 0.25 mm,

well-defined cavities, none of which was ringed with
while cells. UL contained many parthenogenctic
female, juvenile and male nematodes and eggs, ‘The
three durger galls cach contiined brown ‘streaks’,
running from the centre to the outer edgcs,
upparently foriied by the darkeniig of intaet plant
cell walls. These observations on the process of gull
induchoo madieate that & flevicornis and nematodes
are always present together and that both Ay larvae
und hematodes continue (oO develop, thus increasing
the sive of (he cavities, We have not been able to
determine Whether gall initiation requires. both
species lo be presentor if only one is required, Given
that nematodes are present in the carly wall miaters
pllor to Hatching of fly eggs, it is possible that
initially, wall Uissue is produced in response to
nematode seeretions (see Discussion).

Rey and dorsal shield of Feveusouina Clavicarnis

The egg of FO flavieorniy is spindle-shaped and
tapers to a short, blunt spedicel’ at the micropylur
(anterior) end and to a seniewhat longer, tapered
process al the ather end (Fig. 2). The egy iy 0.65
SE 0.06 mm Jong by 0.26 + SE 0.03 mm wide (n=
4). 1tas hirger than the egg of 4 riehotsen’ Tonnair
which wus deseribed and iHustrted by Corrie
(1937),

Second and third instar larvae of most deseribed
species OF Fereusebid have av more or less elaborate
dorsal shield (Curie 1937) In A flaniearnis it
consists of a selerotised modification to the second
and jhird thoracic segments und first ubdeminal
segment comprising a plate with sometimes three,

Call weight (g)

Pig. 3. Relotionship between tou iuiitber ol emerences
per ull lor (ial Perguseaniit flavins alone, and (by Fe.
flavicomiy plus ussociied Pimenoptene versus wall
weight (in grant) reared between & Marvel und 30 April
1995 rom Auealyniy cameleon ay Conlwa Seuh
Australia

mostly four and rarely five anteriorly projecting
prongs aising from the third thoracic segment, 'The
structure Of the shield is similar to that depicted lor
Ke lockharti Tonnoir by Currie (1937), Ths finetion ts
imphicaled in feeding and the enlargement of gall
cavities (see Discussion). Indeed many pellets of
phant material were found i the gall cavities often
adhering to the dorsal shield. Puecal material within
the gall cavities was not observed,

Emergence and Phenology of Fergusonina ate
associated Hymenaplera

Galls collected in late February 1995 contained
only hurval stages oF FB flavicornéy but pupae were
presenl soon alter, Galls were cuged in the field on 5
Murch (day O) and the first adult £) flaviearnts had
emerged by day 3. Adult flies continued to emerge
for about the bext 40 days, peaking around day ES
(23 March) (Fig. 4). The 370 adult A flavicerits
that emerged trom the caved galls had o sex ratio
slightly based in favour ot males (1. 1.2). Further,
the number of Thes that emerged had a positive
rehationship with gall size (measured as dry gall
weight) (Pig. 3. line b), However, the regression
against gall size was steeper for flies plus associated
Hymenoprera (Vig. 3, compuring lines a and b), and
the correlation slightly better (°° = Ob versus 0.87).
compared with (es alone. The reasou for this ts nor
clear bul, presumably, gall size is a reasonable
estimate Of available food and so for 4 given gall
there must be a limit to the number ob ttes a ecu
support. Hewever, all galls surveyed (Table 1)
contained some Hymenoptera, and these individuals,
whether they are phytophigous. parasitic on F
flavicorniy, or both Ciaquilines) will have displaced
or killed Mies that otherwise would have conmmpleted
development and emerged. Hence, the steeper slope
of the regression line for flies plus Llymenaptera
represents what is a closer estimate for the carrying

ay

per d.

Emergences

134 G. S. TAYLOR, A. D. AUSTIN & K. A. DAVIES

a Fergusonina flavicornis Malloch € Eurytoma spp.

0.5
0.4
0.3

f Megastigmus spp.

0.5 2 Bootanelleus sp,

C Cirrospilus sp.

68 12 17 23 | 13 30
March April 1995

68 12 17 23 | 13 30
March April 1995

Fig. 4. Number of emergences per day for Fergusonina flavicornis (4a) and the six most abundant Hymenoptera (4b-4¢)
reared from galls between 5 March (day 0) and 30 April 1995 (day 56) on Eucalyptus camaldulensis at Goolwa, South
Australia. The standard errors around the means are different for each species because the number of galls in which they
occurred was also different (see Table 1).

BIOLOGY OF FERGUSONINA AND TTS ASSOCTATED HYMENOPTERA

138

Tabhh 1) Vitel emervences fram Marc -30 April, $999 of Perausonina lhavicornis and asseciaicd Aymenoplenur spp.
jr Fergusonina/Pertusobia gal/y on Buealyptus camaldiilensis at Goolwa, S. Aust.

Gull number

Species - | 2 3 4 5 6 7 Total
Foreusonina flasicnrnis Malloch 1 LA¥ES An 44 44 {RO BR 370
Brecon (ergusonints ap. Woy. a (7 0 ‘) ] WW | 3)
Poeciliciyptis spp a | a | 2 2 0 G
Cirroypilus sp, 6 7 | | 2\ 37 13 KY
Aindevits sp. } | 0) Y 5 Is h 44
Pediohius sp. a 0 i] J if) ( a 1
Eueviane spp. i iI 3 WW 4 25 3 71
Crelvevba higher te Ashmead () 5 | 2 2 5 () W
Ditopinalella sp () () () ‘) 4 3 2 Q
Messiiaanns spp. 6 5 2 19 12 30 1 Th
Bauranellens spy 2 () () \7 2 i? it) 33
47 7 64 MM 130 28 AT

Tolil parasitoids 33

capacihy of a gall ins terms of food resources.
compared with that for flies alone,

The composition of the hymenopteran species thut
emerged from caged galls varied in all eases, except
between galls S and © whieh yielded the same
species (Table 1), Calls contained an averase of
seven species (range = 4-9), with no gall containing
all species (note (hatin Table | species for each af
Poectlocryprs, Eurvloma and Meevastienus have
heen combined), The biology of the individual
hymenopteran species in this guild ts diseussed
below. OF these. several were reared in safficient
numbers (> 30 individtails) to examine their patterh
of emergence (Pie. dbh-2), Brecon ferguyeninus sp.
nov, Cirrospiliy sp. und Megasiignims spp. all had a
peak in theiremergenee prior to thatot fe flavicaenis.
For B. fersisoninus, this peak oeeurred ut the
beginning GF the study (day 3) and virtually all
individuals had emerged prior to the peak of
Jlavicernis on about day TS, while Cirraspiltis sp.
andl Meveastigmus spp. continued to emerge from

walls until day 56, albeit in low numbers for most af

this time. Buderus sp. and Eveyrona spp, bad a oroad
range of emergence times, beginning prior to the
maximum in 2 favicon (diy 1S), peaking alaubout
day 27 and continuing to, or almost to. diy 56 when
the study was lerininated, In contrast, Bootiuiellens
sp. did not start emerging from walls until 21 days
after the peak m 72 flaweornix, Emergence of this
species reached a maximum about day 39 and some
individuals were Still emerging from galls up to day
56, OF the other hymenopterun species that were
reared in jaw numbers (<30 individuals), the
emergence ol Cy miarocineta (days 1-27) straddled
the peak Tor / flavicornis, Poecilocrypius spp. (6
individuals from days 23-39) and Ainvpinetella sp.
(9 inchividils from days 26-56) emerged welbalier /
Jlavicornis, while the single speeimen at Pectiobiirs

sp. emerged between days 13 and 18 (18-23 March).

Taxonomy dnd biclog af Hymenoptera associated
wot Ferguson

During the study J2 species of Hymenoplera were
reared from Fergusonine/Ferguyabia walls on 2.
cannidulensiv. This section describes some aspects
of the biology of the species concerned und
speculates on other aspects of their association.
based on information available for other related
species. Notes are provided on their taxonomy,
including the description of a new species, and an
iNustrated key is provided to identify the 12 species,

Family Bracoutdae
Bracon fergusoninus sp. now.
(FICS 5-8)

Holotype: ¥. South Australia, Goolwa, 6-8.i11.1995_
G. Taylor & K. Davies, reared Irom Fergusenine
galls on Avealvprus sp. (ANIC).

Parawpess & 2S, 18 Ga, same data as holotype
except some with dules as lollows 9-12.95, 13-
17.0095, 2-1-1985 (ANIC, WARI).

Female

Size: Length 24-2.6 min (excluding ovipusitor).
Colour: Head, scutuim, pronotum, legs ind anterior
Iwo-thirds of metasoma yellow-brown to orange:
brown; aplennae, ves. posterior parts of mesosomia
and metusoma, drstal half or hind legs, proximal, mid
and hind coxa wad oevipositor sheaths blick:
sometimes acca around acelli and posterior maryinol
scatum darker: wings evenly infuse:tte.

Head: In anterior view face hall! width af head: eves
moderately bulbous, hairless: face, vena and
mandibles evenly covered with long hairs; in dorsal
view vertex, dorsal temples and Jateral frons with

136 G. S. TAYLOR, A. D. AUSTIN & K. A. DAVIES

BIOLOGY OF PENG USONINA ANDES ASSOCIATED HYMENOPTERA a

loug hairs: in lateral view lower temple glabrous:
untenhue slightly longer than houdy; Tagelloneres:al]
Jonger thon wide,

Mesosoua: Souttim and seutcllam) smoowh, shiny.
with a few Jong sparse Talks on posterior parts: of
cueh rergite, seniund wilh sparse hairs along daleval
Hmrainund wlong Non. natiul indicated by faint
depressions Weare prominent Te posterrar tall
scutelhay sutuee slightly depressed) and curving,
posteriorly. comprising 1214 Toyeate pupetures:
dorsal and lateral seutellunt delimited by row of
longish huis: propodeuim with medial longitudinal
cari Wn posterior One-third, Liinthy depressed in
anmenor vnndine, with sparse long hairs in litera
pur in lateral wew preagtun) und mesapleuron
sencorh and Hrostly hairless: epienemial wea covered
with long hairs; metuplearon cavered with Tong
halts, smooth in dorsal ball. faintly ruywse in
postero-ventral parts legs evenly and densely setose
exoept for outer surfaces of eoxaey fore wing, will
(Mand 1-Rs+M faintly bowed, 2-R4 straight: 3-CUI
evenly curved: 2nd submurginul coll clougate. sides
(Rs and 2-My almost parallel, dst diseal cell
moderately clongule (1-RstMp 2.8 x length af incur.
Mensome Mostly smooth throughout, sclerotised
part oF TI slightly longer than wide, lateral margins
With long sparse hairs, posturo-ncdial lobe of TL
broad, Bunty longittidinally stiyte along posterior
murgin. delimited by shallow buleeal hiraws which
are pereurrent lo base of Th: T2 with broad medial
loneitudinal carina in anterior half weth a few strive
oither side, postero-mudial ures very fuutly taised
and tridueulars; T2 and TS subequal in leneth und
comprising about one-third length of metasomea, with
sparse long hare i) lateral parts: T3-TS about three
quarters leneth oF T2-T3. with tainsverse row af long
hairs and at literal mareins; ovipositor aid: sheaths
slishily longer (han body, sheaths with even covering
of long hairs, Slivhtly Jonger than diaieter of
sheaths.

Mele

Diflering from female as follows: Length 2.32.6
ming badly generally narrower and mare clonzate,
pariioulurly posterior mesosoma and TL; scape and
ofvo proximal one-quarter oF antennae brown, lees
with sane coluur pattern but yellowery wings
Hacrower,

Comments

This isa very large genus in Australia of whieh few
spevies have heen described, Prmurily, members of

Piws 58, Brecon fereusouiius sp. now, 25. Dorsal habitus (not showing ovipesitary 6. Silhouette of dorsal body show-

the venus are parisine on lepidopteran larvae bul
they lave also been recorded trom Coleoptera.
Diplera and lew ling pergid suwfhes (Quieke
IOBK: Quieke & Ingram 1993; Atistin & Paulas
1989), Other than one unsubstamiisted record oF a
Brieonk species being revved fram uw gall-forming
homopleran (Chadwick & Nikitin 1976), this ts the
first record Of an Australian species being reared
Troma plint gall. Studies onthe biology oF numerous
specres Overseas and from Australia indicate that all
Members Of the genus are solitary primary
eclopurasiloids, Give) (His, did that A fergaventinty
was reared in moderalely hinge ninnbers form five oF
seven walls ducing the project (Table |). (his speeres
ismost bkely to be parasite i Feggisenii hirvie
Previously, Cierig (1937) reported nating an
unknown braconid frou galls of Peewee in Ure
Canberra area, und unis may be the sume species us
deeribed here. However. there is apparently ni
volicher material evailuble from Currie’s study tthe
ANIC or elsewhere, Further, Currie (1937) lyund
this braconid “lo feed indiscriminately on gall Ussues
and Ty lorvaw” We have been unable to conlinn
Curie’s observations hut. i this species isu
faculative feeder on gall tissue, Th will be only the
fourth record of phytophaey. ii (he Braconidae in the
world (Austin & Dangerfield inpub.), and (he frst
for the subliamily Braeoninae

Bracon fersuseniius diflers trom other described
Austtalian Braces species in the form of the mnedial
propodeum, shape of TL, sculptiring on T2. length
of the ovipositor. and ly colour pattern. Jt is easily
identified fron other parasitoids wssoeiated with
Pepeusonina/Fergusebia “alls by its complete wing
vention (Fig. 7), eireular und depressed (eyclostome)
clypeus (hig. $8). elongate ovipositar (Pig. 6), ane
orange and black colour pattem,

Family lehneumonidie
Porcilacryprys spp.
(FIGS 815)

This genus belongs to an unusual sublomity. the
Lubeninae. in that il represents one Of rhe mose
plesiGmorphie groups of iehncumonids sind has a
Gondwanin distribution (Guuld 1983, 1984; Gauld
& Holloway 1986), Peecioeryptiws has heen
postulated to he phytophagous on the basis of the
structure of the mandible (Short 197%) and it bus
previously been reared from galls on Acactr amd
Eucalyptus (Gauld & Holloway 1986) and, more
yecently, from Banksio (Austin & Dangerfield

ing length of ovipositoy 7 Fore gad hind wings (vein eet arrowed in find wing), 8) Anterior view oF head showing cit

wular clypeus. Scale bars OS min 5; 1 Oman: Os cin &

138 GS. TAYLOR, A. D. AUSTIN & K. A. DAVIES

unpub.). However, only very few specimens have
ever been reared, as is the cuse here (i.e. six
specimens from four galls - Table 1), indicating that
the genus is more likely to be a parasitoid or
inquiline than a primary gall-former. Surprisingly.
two of the four known species of Poecilocryptus, P.
nigromaculatus Cameron and P. galliphagus Gauld

& Holloway, are here associated with Fereuy-
onina/Ferguyobia galls. Both species have previously
been collected from the eastern part of the continent,
Queensland to Tasmania. but both are recorded here
from South Australia for the first time, Poecilocryptes
nigromaculatus has been reared from anthribid and
chalcid galls on Acacia longifolia and criococeid

Figs 9-14. 9-11. Poecilocryptus nigromaculatas Cameron, 9. Dorsal habitus, ¢ (extent of black markings indicated by
shipping). LO. Silhouette of dorsal body of @ showing length of ovipositor. 11. Fore and hind wings of 3 (vein 2meu

arrowed in fore wing and vein r-m arrowed in hind wing). 12-14. Cirraypiluy sp, &. 12. Hind leg, 13, Fore wing. 14

Dorsal head and mesosoma (extent of metallic green markings indicated by stippling). Seale bars = 1.25 mm 9 115 0.5

mm 12-14.

BIOLOGY OF FERGUSONINA AND [TS ASSOCIATED HYMENOPTERA 139

Figs 15-20. 15, 16, Euderus sp. &. 15. Dorsal habitus (pr = pronotum). 16, Fore wing (postmarginal vein arrowed). 17,
20, Euryioma sp. 17. Fore wing (stigmal yein arrowed). 20, Dorsal head and mesosoma (pr = pronotum), 18, 19,

oO

Pediobius sp, 9. 18, Fore wing. 19, Dorsal head and mesosoma. Scale bar = 0.5 mm.

10 ths PAYEOH, A 1) AUSTIN & K, A, DAVIES

mills on Beales, While Po galliphaeis has beer
ussuciiled wilh unknown galls on Ey eelevarensis
und E. punviflond (Gauld & Holloway 1986).
Poeciloeryplay spp. aire distinclive: cornipured with
other parasitoids in the guild because of they laree
size (approximately LO fut excluding ovipositar)
wid wink yenation (Pig. Th). Although the two
spewies are very similar io each other they can be
separated by the colour of the hind lemur and length
Al the OVIpOsitar

Family Eulophicae
Cirrospilas sp.
(FIGS 12-)4)

Members ol this very large and taxonemiently
difficulé cosmopolitan genus are relavely small in
size und areootien brightly coloured with metallic
markings, us is the species here, Different species
have bee feared as primury parasitoids or as
hacullative or obligate hyperparusitoidts trom leat
mining and ealltorming insects. ‘Vhunilert and
Austin (1994) recorded Thur species ussaciated with
Ihe mines of PAvldereopheged on cueatypts i South
Austin whieh, in the case of the dominant species
Co muerginiveatus Giranlt, was shown to be a
hicullative byperparasito of Brecon and other
chaleidoids, The biology of the species here is
Uneleur and it could be either a privkiry parasitoid on
hergusoniue larvae or facnttalively ly perparasitic on
one or more ol the gallassociated Hymenoptera. I
Was The most abundant parasitoid encountered durijie
(he stiidy and it was recorded fram all seven salls
(Table 1). The species ts close to ©) aecipitiy Giraull
bul differs in the colour pattern of the dorsal
mesosomi. and this may represent a mew species.
Within the opurusitoll guild ussoertod with
Fergusonind/Ferusebia galls, it can be casily
denied based on the form of the fore wing
vention (Pig. 13), colour pattern and shape of the
body (Fin. 14),

Buderus sp.
(FIGS 15. 16)

Like Cirrespilus, this is 4 lige genus whose
Members are difTewlt to identity, Extratimital host

records indieate (hal Muderny ave parasitvids of

lepidopteran larvae or are hyperparasitoils,
particularly of Braconidae (Boutek 1988). The
association OF the species reared here from
Fergusonina/Pergusebia galls is (he first biological
information recorded for the genus in Australasia,
Given that species overseas have been feared (rom
Bracomdue, jk is possible that this species is
hyperparasifie on B. fergusoninus This proposal is

supported al least io part by the tee that bvderay sp.
was feared Thom galls thar also. yielded individuals ot
B. fersnyoninny CRible 1). and that no ather
hymenopterains in the sane galls ure Liree enous in
body size to provide sufficient (oad for the
development of Zuderis larvae (wiih the exeepnun
of Poecioervyprus, whieh was too rare), In the
parusitoil guild ussocioted with Pereysenntnd
Fergivobia pulls, Evclerus sp. 18 easily separated
fron) other species by is metallic blue-green colour
wonwite hedy (Fig. 15). mesosone with very Sine
Peticuhue-punerae sculpluring and fare wing will
postinargiul vein Whout as long as stigma vein (Fig.
10)

Pediobins sp,

(FIGS Lh. 19)

Members of they genus are Kiwi to he primary
and hyperparasitoids of eres, pupue and somenmes
lurvae of wirious inseets. matoly Lepidoptera,
Coleoptera, Diptera and Hymenoptera (Boutek
T9KS), Several species have been reared trom lear
feeding insets on enealypts (eas. Uribe fiers
Walker und Phvidcwaphae frageanth Riek -~ Austin
& Allen 1989: Thunlert & Austiy 1994) ahd various
leal-mining Diplera (Boutek T988), bul upparenily
none has heen reared front gall-associated hosts. The
species here was tecorded by only wsingle speeinen
(Table 1). Although ats biglogy is net known, (his
very low abundance would iidiesdte tal i ts
probably a hyperparasiloi on another hymenopterin
in the Fergusomtna/Fereosubia euild, Pediohuis sp.
can be eusily separied from the other parusitanls
recorded by its relatively short compuct body (Fig.
19), Black colour very Finely reticulate dorsal
mesosoma, und elongate postmarginal vei (Pig. 18>

Family Eurylormidae
ary toner Spp.
(PIGS 17, 20)

‘This is a very hinge genus in Australia contains
species with divergent hinlogies, Species can be
phytophagous, facultatively patasitie, obligatoerly
parasitic ob hyperparasiice (Boutek 1988). In
Australia, the majorily of species ure postulited to
fall into the Jalter two culegories, but the generic
classification is not well understood and species
belonging lo apparently closely related genera and/or
species groups are known (9 be either phytophagous
or parasitic, A number is knowh to he cetoparasitic
On gill-focming Mscets or hyperparasitic through
iehncumonid und braconid cocoons (Boutek 1988),
Currie (1937) reported one species of Enryvronna, &
“varirufipes”. do be phytophagous in /ereusondia
galls near Canberra (varindfipey is an unpublished.

RIOLOGY OF PERGENSOAMING ASDITS ASSOC TATED HS MENOPTERA I+]

maruseripl numne - see Dahiis }98o) Boucek lost).
Unfartinately. we have mol been able ty confirm the
hiolowy of the Species encountered in this study by
direet observuition. One of them (see below) ony be
Ihe Same species as pecdrded hy Currie (NBL ne
voucher material rs ayailible from his sity) and
therefore possibly be phytophagous (see Discussion),
ar they could be differentspecies und be parasite on
Fereqsoning larvae or on-one of the other hymenopterats
im the guild, possibly B. fevgisandiy sp. now. The
Species here was recorded from all ihe galls surweyed
aud was the third most abundant hymenopleriun wu
the Ferwivenina/lemoisobla guid (Table 1).
Lurviemea often display intraspecific morpho-
Fowieal virtability and ape therefore diffieult to
separate al the species level, The material reared here
varies slivhtly (size. colour. pilosity and sculplucing
and may represent more that one Species. The gents
can be cusily separated from other parasitoids i the
euild Biased on the shape of the pranotum, dorsal
seulpturing pattern (Pig, 20), wnd wing venation (bus,
17). The sexes ure brahly dimoerphie wilh niles
beine smaller, having w disproportionately sotaller
fetusonia, aid day mmoticul pilose antennae.

Fumily Pleromalrdie
Coelyvha nigracinem Ashmead
(FIGS 23, 24)

This species und all members of the genus. are
probably inquilines of hymenopleran aed dipleran
gall-formers on WWewei and Aucedypras (Boutek
DORK. Previously, C. riigroetuere Was been reared
Irom the vally of the phyluphagous preromutids
Trivhtlogaster and Perilampella (Noble 1940, 1941)
imd, dn the latter Gase. has been proved to be an
inquilipe, Les. i Kills the resident cecidoyenic wetsp
and forms its own cells to teed on the gall tissue. The
Species was orivinully deserbed frome maternal
reared from “agromyzid galls on Ereulypriey
corvmhose” (identified by Boutek (988) us
Fervusonina galls) in Sydney (Astinead 1900), anc
was subsequently recorded trom herenseorrntet
Kerensobie ails as an obligate pirasitonl in the
Cinberrvarea by Currie (1937). Specimens in ANIC
were collected from galls on BE. camalduleasiy al
Alice Springs, NTand 7. sieheri L. Johnson and fe.
coyvedeline Labill. at Bicheno. Tas. (1. DB. Naumann
pers. von. We hive been uimible to confirm: ibs
biology in this study, but clearly it is most likely to
he an amquiligg oF primary parisitoid of Fergasenind,
Cocloryha nigroeinene was feared only in mocerate
Numbers here bul i wis reconled from five of the
seven wally surveyed (Table 1) Wis apparently
(distributed along the cust coast of Australia from
north Queensland to Tasmania, but this seems to be
the first record of the species trom South Australia.

Covloevii nigroenienme cin be easily sepiurled troy
other puirisilowls assueiated with Merusaenmne?
rerensobig dalls by is Gleangate stigmal yeu. shor
oviposilor and distinctive bright Yellow body und
metasona wilh blick (raipsverse stripes (Fig, 24).

Ditrapinenedia sp,
(PIC 26)

Members of this gemus have previvusty heer
treated under the “Toiyindae of as miseneasierine
plefomalids, but mos recently they have heen placed
in thelr Own subhionyly, Ditnopiieiellinde, within the
Pievomalidae (Routek T988) Previously, they have
been roared Irom Apvomnerplia galls on /ucolyplits
dnd iinknown walls on Lucey, Acacia, Casuarina
(Boucek 1988) ane a number ol other plints, The
hinlogy of (he enn’ 1s poorly Kiown However
Cume (1937) reporied 2. reinyresstvenity Girl ty
be phytophious within Fereusooin/Tereusepie
solls in the Canherra area During Unis study, only
nine individuals were reared from three galls (Table
I) and. bused on these cuinbers, the species ts mone
likely lo be an inquiline pr parasitic on que or more
wull inhabitunts, We have tot been able postiively
to identity the species here as 2. compres theniris.
primarily due to morphological variability among
species in the genus (Boutek 1988), However it
canbe eusily distinguished from the other parasitoids
in the guild by its bright metallic green hoy,
laterally compressed metasoma. und caserted
oviposition (Pig. 20).

Fanly Soryovidie
Meg vtionis spp.
(FIGS 21,22. 35)

The Megistigminge are besh characterised by their
large. knub-like fore wig stignm und) elongate
ovipositor of the female. The genus Meuustigiiey is
large and Waxonomically difficult in Austria. with
more {han 40 described species. As well, its members
have diverse biologies in thal apparently closely
related spectes cay be phytophagous, cecidopenie,
inquilines or parasitic, although the detailed biology
of no Australian speetes has been confirmed (Bouck
JYS88), Various species have heen reared front dipteran
walls on Asterramtulasis, Apionorpha (Eniococeidae)
on Bucalyples, Bruchoplegus (Eurytomidae) galls on
Kremacinis, Trichilogamer (Pleromalidae) galls on
Avueta. as Well as unknown galls on numerous mulive
plants (Boutek 1988). Two species, ML quinymexetae
Girault and Megasigomy sp. were reared from
Fergusontna/Fergisobia galls in the Cunberre aren
by Currie (1947), and were reported by hin to be
phylophagous, We have been unable to confine
Curne’s identificahon ob these species, or Lo show

142 G_S. TAYLOR, A. D. AUSTIN & K. A. DAVIES

Figs 21-27, 21, 22, 25, Megastigmus sp. 2.21, Dorsal head and mesosoma (pr = pronotum), 22. Fore wing. 25. Lateral view
of metasoma and ovipositor. 23, 24, Coelocyba nigrocincta Ashmead © , 23, Lateral view of metasoma (extent of black
markings indicated by stippling). 24. Hind leg. 26. Ditropinotella sp.?. Lateral view of metasoma and ovipositor. 27,
Bootanellus sp. &. Anterior view of head. Scale bars= 0.5 mm 21-26: 0.25 mm 27.

OLOGY OF FERGUSONTINA AND ITS ASSOCIATED HYMENOPTERA

that they are indeed phytophagous. The number ot
individu obtained for the two species reared here
was pooled because of initial problems in separating
them tasxooumieally. Combibed. they were reared
from all seven walls surveyed and oecurred in large
hufbers (Table 1), being the second most abundant
genus present in the guild,

Mevastivmuiy can be easily distinguished fron the
other Fereusenina/hergusobia pall-inhabiting ysrien-
optera by their spat-like fore wing stigma (Piz, 22).
bright yellow colour with some dark inarkings, and
the length of the ovipositor (Fig, 23), From each other
Hicy Gun be separated by the colour of the metasonta.
The metasoma of one species is mostly dark whereas
in the other a is yellow with transverse black stripes.

Bontanelleus sp,
(FIG. 27)

This is a moderite-sized genus of (negastigmines
characterised by the presence of a medial (opth on

the lower clypeal margin and some metallic
colorution. There are 20) described species in

Australia of which three have been previously
associated Wilh seeds of Caseeriny and galls on
Cirrus and Mieracitrus (Boutek 1985), The hiolvgs
ol the species
Fergusohia galls is unknown, bul it was rearcd in
moderate mimbers from three of the seven galls
examined (fable |), Based on these numbers uid the
fact that it emerged well after all other gull
Whabiunts. this speetes 15 wiost probably a parasitoid
or inquiline. The speeies is very striking anil easy to
separate Fron the other Hvorenoptera associated with
Ferunsonina/Fergusobia galls based oi ais yellow
body with bright metllic green dorsal head jad
mesosoma, black dorsal metasoma and clypeus with
aomedial oth (hay. 27).

Key to the parasitoids reared from
Ferunsonina/Ferrusobia Galls
in South Australia

1. Fore wing with more Uh one efclosed cell,
ingderuely complete venation (Figs 7. 11h
prepeetus absent (Lehneumonoided) ccc cern
Fore wing with no more than one enclosed vell.
verition restricted la untertor margin ob wing
(Pigs 13, 16, 22): prepeetus ‘dln
(ChalgidGidéa aroun ui onitr dienes?
Fore wing ywith ye THicu present; hind wing
wilh vein rin mecting Rs afler Rs diverges from
Sce+R (Pig. 1!) (lehneumonidae) [lenuth about 10
nit without ovipesitor; ovipasitor shorter than
metusema (Pig. 10): elypeus nortmal: body
oninge yellow with black stripes on metason
and hind leas (Fi. 9] Baecioeryptu soc. oF

i)

reared here from Ferensovinas

6,

{),

O

1),

1a

Fore wing willl vein 2meu absent: hind wing with
vein rm meeting Rs hefore Rs diverges [ram
SctR (Pig. 7) (Braconidae) [length about 2.5 min
without ovipostlor: oyipositor almost as lone us
hody (Fig. bi clypeus circular and depressed
(Fig. $); orange in colour with corer
Mesos aidl fretasoma HNACK | oc ccseseereeen
Braco erenteaninus sp. NOY,
Hind coxa blac ky ovipositor about 2,5 x length of
hind Ubi. P onigromacilans Cameron
Hind coxa yellows ovipasitor about 2.2 8 Jength
of hind tibiie.. Ra sn
Foci soci "gallinhauns. Gyuld & Holloway
In dorsal view, pronoturm large and sub-
rechingular or broadly rounded (Pigs 20, 21) 5
In dorsal view, propotdny arrow and comprising
little if any of dorsal surfuce of mesosoma (Fixes
15. 19)... A
bore wine 2 Mitt Mengatc anginal vein (Fig, 17)
(Eurytamidue) [budy wl black, lees often brown
op orange-brown detally; female with dorsal
mesosoma Cowesely retieubile-punclate (Pig. 20),
oyipysiior short and virtually hidden under
THGLISOIND] occ ecceeseee er teereee ee cere ee PPI SDD,
Kove wing with lirge spotlike stigmal vein (Fig,
2): female with aoa exposed ovipositor
Fig 25) (Tern eels atensericscettasae oo
Body all yellow with some dark markings:
clypeus without a medial tooth (Mevastigzts)
Ventral und lateral durbleby af rhody yellow, dorsal
head and imetasonia bright metic green, dorsal
metasoma bluck: clypeus with medial tooth (Fig.
27) _Bauranellens sp.
Metusomut mostly dark dorsally. stripes or Striped
PATLEP THUISHNCE yc eueNitunitty sp.
Metusomna yellow with rransverse black stripes
nome scree Megeyiiginly sp. 2
Hind tarsi S-segmented (Fig, 24) (Pleromialtdite)
at)
12) (Eulophidae)

gy cealevdaglvacaverd sabi rrzet aaa tasie erat

TaRET SSR OSESER CT CSESEEy

Hind tarsi A: siturntenitesd (Fig.
Body briishd yellow: or yellow. row the rmisigecnn
depressed. with broad black tramsverse stipes
dorsally: ovipositor short, hardly pn from
posterior metasoma (Fiz. 23) —. veapidlndiee

Coelocyhu nigrouit te Ashmead

Body bright metallic green: metusoma
compressed; ovipostlor exsefted postertarty
(PIS. 26) secon enineditropinolelle Sp,
Body with disupcr bright colour pattern.
including metilic green markings on dorsal
mesosoma (Big, 14) and black transverse stripes
an Nicrasanya - 2. Cirmapilus sp
Body with uniform dark culuur

\44 GOS PAVEOR 41 AUSTIN & i AL DAVIES

1. Body Clongale (fig. TS). aietalhe blueseeeey pn
rolour, dorsal mesosoma wilh very line
betieuhue-punwture seulpluriog: Fore wing wath
Postural ver abodl us long ds stigmul vein
PETE PD secre ciarserecnersneecs ce Aeerees ST
Body relatively shork cunrpaet (hie, 1) ane
black in-eclour;, dorsal mesosoma with very fine
jeliculaie = seulplufing: tere wine willl
postivurginal Vein nich langer thay sting! vei
(ETE. PED setisreeconess rene csnnsanenecsssnnel Cel opines S77,

Discussion

The mechinisings by whieh the best plant cells ave
Ttuced lo prolilerate le fem a wall remain
problematic, Currie (1037) observed thar nematodes
were uctively feechay and causing vell prolilerdion
Of plant tissie around dre eves af the fly before they
hatched, "This suggests chat plvenllc ponatodes gre
tesponsible for wall initiation (presumably via
salivary seererions). This pot inconceivable. however
that a fruetion of oviposition Uuid whieh is injeetod
ints the plant by the fly along with the eges, cr
Mechanical duiyage by the oviposttar alone. abs
cHuses cell protiferation: 1 his been ohserved wilh
aher berwasenind/Pereusebia assochanons (Davies
unpuh.) chit galls in whieh Wy hirvae hawe died, but
in Which nematodes ure still present. remain ereen
uid viable, Cure (1937) reported that when infertile
Ny epes were deposited, galls were intiated. hut were
‘Wberted at an early stage’ while the nematades
persisted Tor some noumths but eventually died jut
He suggested that further gall eroweh wis
ultoibitable to Ty larvae. Whatever Ihe mechunisin,
xall growth for A flavicarmy was particularly rapid
indhar galls attained fill size ina mater of weeks
duty the late summer generation,

While no faecal material was observed in gall
cavities of A flavicerais, Curme (1937) observed Wal
faeces were “practically absent” in all bul one species
OF Fenuwentin. Te postulated thal nematodes. which
hy then occur within the fy cavity, feed on [ly feees
(hus keep the cavity tree of waste. Given the strong
styler of the nematodes, iis more likely thut they Feed
on plant cells, Microseopic examination of specimens
cleared in elycerol indicated the presence of ia rectum
umd anus in A flavicerniy larvae, Tt is possible,
however, that larvae may store, or produce mininul
umounts of exeretory products to avoid: fouling the
cavily. Hibs isso, i supgests iin uduptation sinvlur to
that found om the gall-lorming Cecidomyiidae
(Diptera), the larvae ofiat least some of which possess
a funetional anus but produce no noticeable waste
products (Gagne 1989), Perhups related lo (he storage
of wasle, Fergusenina puparia were observed anchored
within the gall vavity by means of a gelatinaus
substance atthe posterior end. It is conceivable that this

subshince represenls digestive waste vended just prior
to pupation, Unlike some cecidomiyiid galls (Caine
JYR9), Tonal colonisation of Fresh Fereuseitiind sills
was Hot observed. The fly larvae inay dee the dorsal
shield, will its anteriorly directed prongs, (o serape the
observed pellets of plant material front the wall of the
“awity, Corrie (1937) deserihed third instar (ly laryie as
“our down winner layers af the cavities for fond
The shield nay also Be used to-anehor (he kurvae Heane
spol within the cavily and to break down parts of the
cavity walls prior to pupation to facile emergence of
the adult Hy,

Ie is dineléur how due (Mies escape front the walks
wher cihefeinie frome puparit Dissection al gulls
revealed (hat by the fime of pupation, cavilies on the
vdges oF whe lh were separated trom the outside
only bya thin layer of epidermal cells, suggestini
(hat the larvae (uiinel near to te sutlace prior te
pupalion, Presumably, ail Mes are able to emerge
Wirouzh this thin dayer relatively gosily. Cywvities
deeper in the nidtiire cull are separated hy hon layers
Of PAPOUCLVHINGLS Lissue and sonmetinnes open mt
evel) other,

Vhe 12 species Ob Hymenvplera reared front lev
galls ol. Jlaviearnis during (his shuty cepresent iw ice
the jumber previously recorded from feruiverine
galls. Curne (1937) reared only six species from
flower galls ja the Canberra area. while Haris (1982)
recorded Tour species from: Fergaxontie galls tn Imnbia
Further, several species ate bere assecrwed with
hercusening walls Tar the fest ine, viz. Bearnellens
sp. Cirraspilus sp., Ludenus sp., Pediohiis sp. ind
wa species OF Poeedocnynus (EP mignneculalus and
P valliphagus), Although the composition of
hymenopleran species associated with individual galls
Clearly differs, the most common species encountered
inthis study, Comnpilus sp. Laeviema spp, and
Mevastinuias sppu were reared from all galls (Table
1), while B fereuseninus, Co nigrocineia anil
Muderis sy, were teared from more than 70% of
galls. The diflerenee indie size of the hyntenoptena
void between this study and that of Currie ()937)
May be wiributable to differences in the structure and
position of galls, Le, A flaviearnis forms leat gulls
while the species studied by Carrie formed bud gulls.
Also, the fact thal galls were caged inthe field duriag
This study onieans thar ull Hymenoptera were
collected. Whether ley were cominon ur rare. Th
dasess the composition of wasp species by dissecting
valls, us apparently undertaken by Currie for at
Feast pit of his stuely, is likely to dese to an ander
estynation ol the trae qumber present, Such dissections
mostly reveal larval stages thal ure offen difficult
fo identify: they aire solt-bodied and therefore are
easily damaged and there is a vreater chance ol
overlouking rare species.

The lew! pally caused by Jo flavicoriy appear, of

BIOLOGY OF FERGUSONINA AND TES ASSOCIATED [TY MENOPTERA 15

dissection, lo have a relauively homogeneous matrix
and, although some woody tissue ix apparent. the
Most importunl mortality agent is likely to he

purasitic wasps. given (hal as many individuals of

Hymenoptera us és flavicornis were reared From galls
(although not all species are necessarily parasitic -
see ubove), However, Currie (1937) attributed up to
60% of Fereusonina mortality ui bud gulls to the
inability oF My kowvae. prior io pupation, to tine!
through the woody gall tissue induced by
liymenopteran inquilines, resulGoe in their
enlombment withitl the gall. He reported that wasp
inguilines were found mainly walhln the outer
surtice of bud galls, but fly larvae deeper in the gull
were largely free of pamisites, He also reported that
upproximately the same number of wasps and [Mies
emerged from Teal galls on A. Starr TM.
(sic), and sugeested (hat there is a higher level of
parasitism in stem tip and leat ip galls than in flower
bud galls.

Iris clear trou) this stady andl as reported elsewhere
(ug, Bouceh 1988: Gough & McMahon Psa) that it
is very difficult to determine the biology oF gall-
associated Hymenoptera. Prymardy, this is hecause
their pre-adull stages are hidden within the gall, their
bidlowy may Change POM one sltige bo another (is ain
the case of inquilines whieh may Kill the resident
gall-former and then become phytophiagous), oF a
may be different from related speeies that do not
iihabit aalls. tots therefore nave to assume
highowreal iaus for wall-yssoeiwld waspy by

extrapolaing from other species or indirect
information such us cihergence time und/or number
of individuals present For such species. their
biology may be determined only hy direct und
detailed observations including their morphology
und life history. 1) Unis respeet, We Have been unable
to confirm the biology of any species feared in this
study, although we speculate above on whal some ol
these species may be doing. as a means of
stimulating further study. The comments made by
Currie (1937) that the species of Dirrapinarella,
Mewasneois and Eurylome he recorded from
Ferguyenina bud galls are phytophagous must be
iteatecl with some cuution, given that he presents
fille or no evidence to support this conclusion, and
at Teast the luller (wo genera are Kiown te contain
species Which are cither phylophagous Or parasitic,
Further, his reporting that @. migraecinone: is ik “true
parasite” ahd that the unknown braconid: (probably
Bracon) he reared was an inquiline is: ineonsistent
with previous studies on these bixa (see above),

Acknowledpments

We wish to think John LaSalle, Grice Grissell, and
David Wahl for assistance with identifying some of
the parasitoid species. Peter Cranston for identifying
F Jlavicoruis, Peter Kolesik for useful disctssions
and Paul Dangerfield for the line drawings. We thank
Lee Haller for aecess lo the bicld site.

References

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146 G.S. TAYLOR, A. D. AUSTIN & K, A, DAVIES

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& Hoitoway, G, A. (1986) Australian
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Harris. K. M. (1982) First record of Fergusoninidae
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& INGRAM, S. N. (1993) Braconine wasps. of
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R. See. 8. Aust. PUB, 99-113.

STUDIES ON THE SOIL-INHABITING TARDIGRADE,
MACROBIOTUS CF. PPEUDOHUFELANDI,
FROM SOUTH AUSTRALIA

By ALAN F. BIRD*

Summary

Bird, A. F. (1996) Studies on the soil-inhabiting tardigrade, Macrobiotus cf.
Pseudohufelandi, from South Australia. Trans. R. Soc. S. Aust. 120(4), 147-154, 29
November, 1996.

A tardigrade isolated from agricultural soils at Avon is the first member of this
phylum to be described from South Australia. Specimens were isolated from freshly-
collected soils and from soil that had been stored dry for three years.

Live and fixed specimens were examined under the light microscope and fixed,
stained and gold-coated specimens were examined using the scanning electron
microscope.

Key Words: Macrobiotus cf. pseudohufelandi, anhydrobiosis, microscopy,
tardigrades, birefringence, biocontrol.

Transactions af te Ravel Seer of 8. Mest (1996), 20(4), 14-14

STUDIES ON 'THE SOIL-INHABITING TARDIGRADE, MACROBIOTUS
CF. PSEUDOHUFELANDI, FROM SOUTH AUSTRALIA

by ALAN [. BIRD *

Summary
Bisb, AJR (1996) Studies on the soilunhabiting Gadivride. Macnebietin cl pyendohifetandi, tron South
Austrulia, Trans. RK Soe, §, der 120 (4), 147-154, 24 November, 1996,

A tarnhiwrade isolated from iuriculturad soils at Avon is the first member of this phyla to be described (ron
South Australia, Specimens were isokied from treshly-colleeted soils and Grom sort that hack heen-stored uy lor
Mmvec years,

Live ynd fixed speeiens were caumined under the light microscope and tied, stiined: and. gold-eoated
specimens were examined using the scvoning electron micraseope-

This turdigride, a stout cylindrical organism about 500 pm long by 150 pin wide with Lhe four pairs of suibby
legs ending 11 paired elaws, Tas heen assigned to the species Meervbiorus ch, preudohufelandion the basis al
the Morphology of the buceopharnygeal apparatus, claw shapes and ege processes. The: stylets are sliphtly
curved, sabre-shaped structires about 40 tim in length and exhibit marked birefringence tinder polarized light.
When these turdigrades are killed the stylets break down ane. disappear

A specimen of ©. cf pyendofufelanei was observed feuding on a nematode anda snTlicagh decrease if the

humber of nematodes inthe soil asthe number of pardigrades increases has been demonstrated,

Kiy Woros:
hioconttol,

Introduction

The tardigrades or water bears belong to a discrete
phylum, ‘Tardigrada, of cosmopolitan dist bution
from diverse habitats including marine. fresh water
and terrestrial environments, The majority are
{hought to tive in water films surrounding the
“leaves” of mosses and lichens. They are microscopic
(with udulls commonly ranging m length From 200 -
S00 ui), are plump and cylindrical in shape and have
four pairs of stubby legs ending in chiws, They iiy
or may not have eye spats.

In Austutlia tardigmides have been recorded from
Queensland, New South Wales, Victoria, Western
Austria and Tasmania but nor from South Absualia
(MeInnes |994).

In this paper b report upon the morphology and
sume uspects of the behaviour of a Gordigrade
isolated from agricultural sandy louwm sol from
Avon, South Australia,

Materials and Methods

Locality ane sail wpe

The tardigrades were wolated from a suil of sandy
loam fextiire classified uso solonized brown earth
(Australian sail grouping) or as an entisol (US soil
chissification). The locality was an experimental plot
on it farm at Avon (latitide 34° 14° S . longitude
(38°)9" By which was direct drilled and had a
wheat/Wheat rotaion.

2 Play font Road Mitcham S. Aust. 5062.

Macrohiors ef pyeudoliyelandi. anhydrobrosis. microscopy. tirdignades, birefringence,

Sail cores (did, 5 em, depth 10 cm) were
collected and mixed in a plastic baw. The sample
mostly used in these experiments was collected in
July 1993 and had been stored dey at room
lemperature for three years. However, freshly-
collected soil from the same site on 29. iii, 1996 prior
lo the autumn rans was alse used for comparison,

Lalreelion frei soil

After thorough mixing of the soil, 50 @ aliquots
were placed in a ousting apparatus. for three cys
(Yeates & Bird 1994). This procedure was replic uted
in quadruplicate aid, aller Hiree days, the collectiiig
jubes were removed and their contents allowed to
setue for 1 oh after whieh the supernatunt was
removed by suction to within 2.5 em ol the bottom of
the tube. This extraction procedure was used lor all
soils, Whether reshly-colleeted or stored,

Counting

The contents of cach tube were poured, after
vigorous shaking. into a counting chamber (Doncaster
1962). The tardigrades gravitated to the Nour of the
counting Chamber between the rings and were counted
under at dissecting microscope.

Light mlerascepy

The lardiznides were exanuned under bright field,
polarized light and differential interference contrast
(Nomarski) optics nsing a Vanox Olympus AHBT
researeh microscope.

Living tirdigrades were examined in distilled water
underacoverslip scaled at its edges with fail varnish,

14M AF BIRD

Speeimens were fixed by adding an equal volume
of boiling double strength FA 4:1 (20 mil 40%
formaldehyde and 2 ml glacial acetic acid in 78 il
distilled witer)y ina test tube toa shaken suspension
of the (ardigrades in distilled water, also in a test
lube. These specimens were processed to pure
elyceral by Seinhorst's (1959) method and mounted
in-antiydrous glycerol on slides sealed toa coverslip
by molten paraffin as described by De Macseneer &
D'Herde (1963) and then ringed with Entellan
(Merck). Both living and fixed material were
photographed using MHford Delta 400 film,

Scanning electran mieroscapy

For observations under the seanuing electron
microscope (SEM), the fixed material was washed
repeatedly in distilled water, post-fixed and stiuned
in 1% aqueous osmium tetroxide, wished repeatedly
again in distilled water, immersed in filtered freshly-
made suturated aqueous thiocarbohydrazide for 40
min followed by repeated washings in distilled water
and a repetition of the osmium fixation. ‘his
ostium-binding lechnique (Kelley ef al. 1973) was
followed by further washings in distilled water,

Specimens were freeze-dried by placui then
belween membrane Allers which were frozen rapidly
by placing them ina slurry of freon cooled by liquid
nitrogen. The fillers with attached tardigrades were
then transferred rapidly to a freeze drying machine
wd freeze-dried at -70" © over a period of three
days. This dried material was then mounted on a
tlass coverslip attached to an SEM stub and couted
with 30 nm of gold to enhance stability and
conduetivity, The tiaterial Was then exapined ind
pholographed in a Cumbridge S 250 Mk 3 SEM
operated at 20 kV using THford 120 roll film CeP4
Plus).

feeding experiments
Attempts i determine whether or not A et.

preudehufelind’ would teed on Riigectonia solani,
and thus possibly omplicating the tardigrade in
slippression Gf this plant pathogen in the field. were
made using RK. selan/ grown in coltore media in Petri
dishes. Specimens ol M. ef, preudehufelane? that had
been washed repeatedly in sterile distilled water by
cenvifugation inan effort to sarfice sterilize then,
were poured on to the fungal plates under aseptic
conditions in a laminar flow cabinet,

Results

Morphology

Both living and fixed specimens of the tuedigrade
isoluted Crom the Avon soil were examined und
measured. Measurements of the lengths und widths
of teh specimens euch of fiaed und living muterial
showed, ux might he expected, that some shrinkage
had oecurred in the fixed material, Measurements of
the living material were made only On specimens (hat
had their lips und mouth parts retracted since this
was the state Observed in all fixed or dead material,
The mean length of living specimens was S114 +
47.7 jim (range 428-580 yim, Table () and that of
fixed material was 423.2 + 48.3 pm (range 364-500
um), Similarly, the width of the living specimens wis
(54.7 + 15.3 wm (range (28-172 pun, ‘Table J)
compared with 131,24 13,3 um (range 108-148 pum),

The lateral view of the living tardigrade with its
mouth everted und showing one of egch of the four
pairs of stubby legs ending in claws (Pig. 1) logether
willh the charactertslic internal structure ol the
anterior region (Fig, 2) and the elaws at. the
extremities OF the fourth pair of legs (Pigs 3, 4) are
shown as viewed under the light microscope,

The fixed tardigrade in yentro-lateral view wath the
mouth region inverted bul showing all Tour pairs of
legs with their claws (Fig, 5) and the ulteaswueture of
the claws on the second leg (Fig, 6) are shown its
viewed under the SEM.

TAREE 1 Ateawnrerents of living Mucrobioius ef) pyeddobulehonai

Part mensured (um) Numbers measured

Length of whole specimen 10
Width of whole specimen Ww
“ position of stylet

support on buceal tube 8
Length of buccal mibe 8
Width of buccal tube K
Leqeth af pharyageal bulb a
Width of pharyngeal bulb 8
Length of macroplacoid (1) 8
Length of macroplicoid (2) 8
Length of microplacoid 4K
Length at 4th frit elay |

Mean sD Range
Sila +477 428-580
154.7 +753 128-172
Ki +/4 TO_S-82 5
M25 #12 4-42
S| +02 55,5
aT4 #21 45-40)
34.4 +13 Sa 4G
W758 +/ft 9-42
0.) ATS 5-7
ry +()2 2.5-3,0)
6.25 #29 4)

A TARDICGRADE FROM SOUTH AUSTRALIAN SOTL 1a

Fig. 1. Whole living Maembions ef. pseudohufelanet. Bright Held optics. Lateral view showing everted sucking mouth
(mi) eyespo fe) aid one of each of the four pairs of legs with terminal claws (arrows lubelled 1-4 from (root lo rear).
The fourth lew is postenor und subterminal, Scale bar = 100 pin,

Fig, 2. Everted head of living Maerobions ef. pyeudohujelundi, Beight Held optes showing mouth (7m), buceal (ube (b),
stylets (8), Stylet support (ss), pharynx (p), macroplacuids |.& 2 and microplacoids (arrowheads from top fo bottom).
Scale bar = 10 fim,

Fig 4. Posterior regton of Macrodiors ef, pretmlohafelandi, Bright feld yptics showing claws oF the 4th legs (arrows).
Seale bar = 10 yon,

Fig. 4. Single chiw from the 4th leg of Maeynbiory ef. preudvlufidand:, Now basal plite at base of claw Garrow), Seale
bar = 10 pm.

A, F. BIRD

iso

A TARDIGRADE FROM SOUTH AUSTRALIAN SOIL ial

Nig. 7. Living Macrebierny ef preadefufelandi viewed under polarized light, A, Whole specimen. Note birelnagence ol
Styles (5), gut contents dnd musele atiichments to lees Gurrow Heads), One of the iwo eyes (a) is also shown. BL Inset
showing aneye (hand the peonounved birefringence of the twostylers, Seale bars = 40 pn A, LO yim B

Although for general purposes bright freld opties
were Jourid to be more convenient than the other
systems used, stylet structure was much more
obvious in living tardigrades under Nomarski and
polarized Hight optics than under bright field. Stylets
were, however, clearly observed under all three
optical systems provided that the tairdigrade was
alive, In both dead and fixed material, the stylets lose
their integrily ind appear lo break down,

When viewed under polarized light (hig. 7) the
stylets exhibited marked birefringence imdicating a
regulitr structural orientition, The stylets are slightly
curved, sabre-shaped structures about 40) tim ain
lenvih (Pigs 2, 7B) When the tirdigrides were
subjected lo oxygen deprivation on sealed slides, the

stylet birefringence gradually disuppeared and was
entirely lost over a period of several hours as the
stylets broke down. Muscles and iWitestinal contents
exhibited birefringence Lo a lesser extent than the
stylets (Fig. 7). However. the museles and intestinal
contents retained their integrity and birelringenee
alter stylet break down. Stylet break down alse
occurred during fixation which accounts for their
absence in cumera fucida drawiegs of fixed material
in the literature. Thus stylets can only be observed ih
living macerial,

The buccal tube (Pig. 2. Table 1) is 40.25 pnt long
x 51 pm wide (percent ratio to the length of the
buccal tube (pt) = 12.67). The stylet supports (Fig. 2,
Table [) are mserted al 81% of the buceal tube

Fin Ss. Whole. fixed umd vold-coared speeimen ol Macrebivtas cf premdiiideland) SEM plotograph showing four pairs
of dems wilh claws Gierows. hibellea Ted fron front te veur). Scale har = D0 pin.

Fig. 6, SEM photogriph of @ 2nd les. Note two sets of double branched Gliws, The mder broach in each claw eurres bwo
vogessory points (sid) arrows) ‘The basal plite at the base of the ehiw is seen here ie a rounded thekening (birger

uerow). Savile Bare + jin

152 A. P. BIRD

Fig. 8 Crushed egg of Macrobiotus cl. pseudohufelandi
lying within the moulted cuticle (mc). Bright field optics
showing the “cooling tower’-shaped projections on the
surface of the egg shell (arrows). Scale bar = 10 tm.

length. The pharyngeal bulb is 37.4 pm long x 34.4
um wide, has apophyses, two pairs of rod-shaped
macroplacoids and a pair of microplacoids. The
macroplacoids differ in length, the first being 10.75
uum Jong (pt = 26.71) and the second 6.0 um (pt
14.91); the microplacoids are 2.9 um long (pt
7.21). The claws (Figs 3, 4, 5, 6, Table 1) have
smooth basal plates (Dastych & Alberti 1990) and
well-developed accessory points on their inner
branches. The length of the fourth foot claw is 8.25
uum (pt = 20.50). The egg, enveloped by the shed
cuticle, (Fig. 8) has numerous “cooling tower’-
shaped projections on its surface which protrude
about 5 tim.

Taxonomy

This tardigrade was, using available keys and on
comparison of measurements, determined as
Macrobiotus hufelandi Schultze, 1834. This species
is cosmopolitan (Schuster & Grigarick 1965).
However, measurements and slides were sent to a
tardigrade taxonomist (S. Claxton) who kindly
examined the material and considers that this
tardigrade most closely resembles Macrobiotus
pseudohufelandi \haros, 1966 and should be
assigned tentatively to this species (S. Claxton pers.
comm.). Accordingly, the Avon tardigrade has been
identified as Macrobiotus ef. pseudohufelandi
subject to further deliberations by taxonomists,

Behaviour

Specimens of M. cf. pseudohufelandi were placed
in Petri dishes containing the plant pathogenic
fungus Rhizoctonia solani. Although some of the
tardigrades lived for several weeks, they were not
observed to feed, grow or reproduce on the fungus.
Furthermore, no significant differences were
observed in the logarithmically transformed numbers
of this tardigrade isolated from soil that exhibited
suppression of this fungus compared with the
tardigrade numbers, logarithmically transformed,
isolated from soil that did not exhibit suppression of
the fungus (7 < 0.05) (Table 2 ). However, there does
appear to be a significant inverse linear relationship
between the numbers of M. cf. pseudohufelandi in
the soil and the numbers of nematodes (p < 0.001).
Furthermore, a specimen has been obtained alive
from the soil that was in the process of feeding on a
nematode (Fig. 9). In soil that had been stored dry for
three years, there was a marked decline in the
numbers of nematodes (4 + 2.9 /50 g soil) compared
with the number of tardigrades (34 + 5.8 /50 g soil).

Discussion

The first recorded observation of a tardigrade was
by Goeze (1773) (cited by Nelson & Higgins 1990)
who referred to them as “little water bears” (Kleiner

TABLE 2. The relationships between tardigrades and nematodes in Rhizoctonia-suppressive and Rhizoctonia-nen-

suppressive soils

Soils Tardigrades Nematodes Tardigrades Nematodes
Mean Mean log mean log mean
counts* SD counts* SD counts’ counts!
Suppressive 19 +91 401 + 108 2.81 5.97
Non-
suppressive 32 +48 227 + 149 3.44 5.25
*SED = 65.2 1 SED = 0.346

‘LSD =0.75

A TARDIGRADE FROM SOUTH AUSTRALIAN SOUL, 153

Bip, 9 plead rewion ofa living Macrobietays ef. preadeliife-
fendi, Phatagraphed inthe process of feeding ona nemi-
fade, right Weld ophes showing mouth (mb siid nem
Wide (a). Seale bar = Di,

Wasser Beir), Three years later, Spallianzani (1776)
teited by Nelson & Higgins 1990) called them “slow
steppers” or /t Tandivrady whieh gave rise to the
phylum name used today, Beeause of their
chamelensue morphology, the randigeades are
recognized today as belonging toa discrete phylum,
‘They probably evalved more than 500 million yours
weo in the Cambrian pertod when there was an
explosive diversification of eukuryoue organinnts.
Although the Avon tardignide clearly belongs ta (he
hufeland grouping tn the genus Macrobrius
(Bertulani & Rebecchi 1993) its specific layonomic
identity a subject to tumher deliberation.. This
turdimrade fas similar measurements to a population
ol Me) pyeudohifeldnd? that includes. specimens
fron WA (S. Claxton pers. comm). Thus, despite the
fact (hat hey have “cooling tower’- shaped eee shell
projections that are more like (hose ibustrated for AZ,
lnfelandi (Nelsoo & Higgins 19890) than Those
ilustrited for M. pyemdoafelandi by Ramuzccou &
Maticci (1983), which have much broader bases, they
are tentatively assigned tod, of, pyejidalifedand (Ss.
Claxton pers, comm).

The ability of tardigrades jo survive tm a wide
Tange of environments ancl their world wide dispersal
most be due in no snl way to their ability to enter
into dnuunhydrobiote stated finetion that they share
with some nematodes und rotifers. | have shower that

M. cf. prendohufeland? can survive for at least three
years in dry soil maintained al room temperature.
This was the maximum time tested and W seems
likely that these creatures could survive for much
longer under these conditions, since i has been
reported (Keilin 1959) that survival times of up to 10
years can occur. Indeed, recovery. but not survival.
hus. been reported to ovcur after 120 years of
anhydrobiosis (Franchescht 1946 cited by Crowe
1971),

Anhydrobiosis is widespread in Phylum
Turdigrada and it is thought thar the disucchande
irehelose functions to protect these organisms since
i accumulates within them us they ure exposed to
desiccation (Westh & Ramldv 1991). Io this respect
tardigrades resemble Whose nematodes that exhibit
anhydrobiosis (Madin & Crowe 1975). In their
morphology, of course, they are completely different
and this is reflected in their different behaviour when
observed in water under a coverstip, Por example, Af
of. pseeudahufelandi iy able to push its way through
an air bubble, This isa capability that | have never
observed in a nemiutode,

The composition of the hwo sabre-shaped stylets
does nolappear to have heen studied in detail and
they are absen) from camera lucida drawings becuuse
they break down and Jisappedr in fixatives.
Sintilarly. they break down on the patoral death of
the animals cuused by anoxia on sealed slides, I
scems strange thut structures whieh), ty the course of
predation and feeding. cun peneteue both plant and
aninoal tissues, should be so fragile,

In the living state, the stylets are readily: observed
under all optical systems. However, when viewed

under polarized fight they exhibit marked
birefringence indicating a regular structural

orienkiion, ft has been mentioned in the literature
thal the stylets are calcareous (KRaestier 1968) so
their birefringence is probably crystalline and their
break down under uhoxie conditions might be due to
an increased internal acidity leading to the
dissolution of calcareous structures such as the
stylets. Clearly such an hypothesis requires further
testing,

Macrabiony el. pseadohufelind? was not observed
tw feed on the plant pathogenic fungus Rhizectonia
solani and t conclude dit the iudigrade whieh 1
have found in large numbers im dgricultaral soils in
South Australia probably does pot feed on this
fimgus in (he field but preys on gther small sou
organisnis such as nematodes,

The ability of tardigrades ta prey on nematodes has
been recognized lor some ume, Jn 1969 Sayre
showed that the tardigrade Mypsihins muvrapy could
be culttired using the free-living nematode
Panagrellay redivivuy as prey (Sayre 1469).
Vurtheemore, Sayre (1969) showed that He aryreps

was able to feed on the plant parasitic nematodes
Meloidogyne incognita and Dilylenchus dipsaci.
Suyre (1969) concluded that “under certain
circumstances, (turdigrades may) give some control
ind these need to he investigated”. A major draw
back to this work was that it was conducted in a
Moist environment Using Moss as a substrate, This is
a farcry trom the normal environment of the plant
parasitic nematodes that Sayre used in his
experiments.

The tardigrade that | have reported upon in this
paper comes from ou tue agneultural soil
environment where it has to survive extremely harsh
and variable conditions. Under these conditions, it
may have the capacity to reduce nemutode
populidions although the devline to the mumbers ot
nematodes compared with (the number of tardigrades
may reflect the anhydrobiotic capabilities of the
tardizrades compared with those of the mixed
nematode populasion rather than predation by the

154 AOE BIRD

tardigrades. However, M. ch, pwendo/ufelandi may
be an effective biocontrol agent. although Lite ts
known of its nematode food preferences and a more
quantilulive assessment of its potential for bineantrol
of plant parasitic nematodes is needed.

Acknowledgments

L wish lo thank Angela Reid CSIRO Biometrics
Unit for the statistical analyses, Soils Division
CSIRO for avcommodation, facilities and expertise
including that of Stuart MeClure for the STEM and
John Coppi for developing and printing the films. |
thank Sandra Claxton for advice on tirdigraide
taxonomy and Jean Bird for constructive eriticisin of
the manuseript, |am gratetul for a grant from the
Australian Biological Resourees Study whieh
provided facilites that made this research possible. |
thank Robin Manley on whose farm the tardigrades
were first discavered for his kind cooperation.

References

Brntonast, Ro k& Reakecut, L. (1993) A revision at the
Macrohioius hufeland? group (Vardignada, Mucrobiotidae),
Wilh some observations on the taxonomic characters oF
cutardigrades, Zoal. Seripla 22, 127-152.

Crown, JH, (1971) Anhydrobiesis: an unsolved problem,
Amer Nat (5, 563-573.

DaAstych, Ho & ALBERTI, Co (1990) Regdeseriplion of
Macrobiotus xeraphilus (Dastych, 1978) comb. noy.. with
some phylogenetic notes (Mirdigrada, Macrobiotidac )-
Mit humb. cook Mus. last, &7, 1ST-169,

De Marsenecr, J. & D'Hirbk, J. (1963) Meéthodes
utilisées pour Perude des anguillules libres di sol Revie
de PAgriculime, Bruvelles V6, 441-447.

Doneasrer, ©, 0.11962) A counting dish lor nematodes.
Nemrtlotowiod 7, V44-330,

Kakst ake, A. (1968). “Invertebrate Zoology” Vol OE Globn
Wiley & Sons Ine., New York),

Rettis, D. (1959) The problem of anaubiosis or litent bile:
history and current concepts. Proc, Ke. Sec. Leiden, B
150, |49-19},

Ke bhy ROL DeRKER, ROALE & BUTEMINK, IG, (1973)
Ligund-mediited osmium binding: its application. in
coiling biological specimens for scanning eleetron
microscopy. J Cnrustriw Res 48, 254-254.

Mctnans, So} (1994) Zoopeographic distribution of
lerresurial/freshwitter tardigrades from eurrenr literature
A. Nat. Hist. 28, 257-352,

MAbin. KoA, Oo & Crowe, ho HE 11975) Anhydrobiosis in
nematodes, curbohydnie and lipid metabolism during
dehydration, J, Bap. Zoe! 193, 321-330,

Nicson, DLR, & Hiacins, K.P (1990) Tintigrada pp, 193-
419 In Dindal. DAL. (Ed) “Sol Biology Guide™ (lohn
Wiley & Sons Iie.. New York).

Ramaccon, G. & Mauectl, W. (O83) fh Phylum
Turdigrada, Terza Edizidne, Mem. dy Mal. helrabiol. Ad,
1-10)2.

SAYRE. RM. (1969) A inethod for culturing it preditceous
uirdigeade on the nematode Punaureddes redivivas. Drany.
Amen Micvse. 8&8, 266-274,

Senbstin, BR, O. & Geicarick, Ay A. (1965) Tirdigrada
trom Western North Ainerica, Clit Cal Pal. Zool. 76.
|-7,

Semuorsr. JW, (1959) A rupid method for the transfer of
nemiulodes from fixative to anhydrous glycerin
Nemuatalogica 4, 67-69.

Wrstit, Bo & RAMLOV, BL (1997) Trehalose aeeumohiticns in
the tardigrnide deenvbieny canmifer during anhydrabiosiy.
J, Exp, Zool, 258. 303-33).

Yeates. GW. & Birt, AoE (1994) Some observations
bn the influence of agricultural practices an the
Hematnde foune of some South Aistalian soils,
Funiton, appl. Nemaiol. U7, V3a-145,

RHOPALOMYIA GOODENIAE, A NEW SPECIES OF
CECIDOMYIIDAE, (DIPTERA) DAMAGING GOODENIA
LUNATA (GOODENIACEAE) IN INLAND AUSTRALIA

By PETER KOLESIK*

Summary

Kolesik, P. (1996) Rhopalomyia goodeniae, a new species of Cecidomyiidae
(Diptera) damaging Goodenia lunata (Goodeniaceae) in inland Australia. Trans. R.
Soc. S. Aust. 120(4), 155-160, 29 November, 1996.

A new gall midge species, Rhopalomyia goodeniae, is described from stem galls on
Goodenia lunata (Goodeniaceae) from the Lake Eyre region, South Australia.
Detailed descriptions of the larva, pupa, male, female and infestation symptoms are
given. Plants infested by this gall midge are dwarfed and develop few or no flowers.
Key Words: Cecidomyiidae, Rhopalomyia goodeniae sp. nov., Goodenia lunata, Lake
Eyre, South Australia.

Fratisdetions af he Raval Society ofS Anyi. (1996). 124), 155-100

RHOPALOMYTA GOODENIAE, A NEW SPECLES OF CECIDOMYTDAE (DIPTERA)
DAMAGING GOODENIA LUNATA (GOODENLACEAE) IN INLAND AUSTRALIA

by PETER KOLESIK=

Summary

Konrsik. P1Y96) Ahapelomyra voodeniae. a new species of Cecdomyidae (Diptera) damaging Goarlerta
Janata (Gondeniacese) in Toland Australian, (rams, See 8. Anse, P2004), 155-100, 29 November 1996

A new pall midge species, Ahapelomvia goedeuae. is desevibedt from stem galls on Choodenta tueoa
(Cioodenieeue) fram the Luke Eyre region. South Aastra Detailed dexenptioas of the hiya: pupa, mile.
ferale and iafestaran aymptours are given. Plants infested by this gallinidge ure dwarled and develop few ar

nu flowers

Rey Worns, Ceciomyitdae. Riupdlonmyle goadeniee sp, nov, Geedenia tian, Lake Eyre, South Ausiatia,

Introduction

The insect family Cecidomyiidie is pourly knawn
im Australia and until now, has been unrecorded from
the inland regions of fhe continent, “The species
described here was found galling stern ol Gorden

Junie J. Black (Goodettaveac) in the vicinity of

Lake Eyre South Australia in October, 1993.

Sulf (or hairy) goodenia, Gecdenia lineata, isa
perennial herb 5 - 20 cm tigh with TS nun long
yellow flowers. It grows in clay soils along
walerenurses and in sandy soils in centr Australia.
Flowerme oceurs throughout most of the ver but
peaks between September and November or after
Nooding or beayy raiotall (Cunmmghan ef af TY8d:
Cooke L986).

The new wall inidge species described below is
placed in Rhapalonyia, a worldwide genus previously
known in Australie froin only ane species, &-
californica Felt. introduced (o control Bacehuariy
fedimifolid (Asteraceae), an American ornamental
plant turned pest on Australian pastures (Mcruyden et
a/, (983; Gagné 1989a).

Materials and Methods

Plants ol Geadena sp. manvesting stem
malformations cuused hy Ceemlomyndue larviue were
collected at William Creek (SQ kot west of Like
Eyre), during a South Australian Museuns collecting
inp, on 22 October 1993. The few lowers produced
by these plants shrank in the course of drying
precluding later authoritative identifieation of the
species. New plants were sampled into 70% ethanol
from the same population on 24 August 1995 and
idenuilied by D. E. Symon, State Herbarium of South
Australia. as G. fianata, The galls collected on the

F Department of Horticulture, Viticuldure and Oenology
Universityoof Adelaide PMB LT Glen Osmond S. Aust.
SO.

firs Ovcasion Were processed in one af two ways. A
sinwll nuinber was cut open und the kirvae preserved
in 70% ethanol, A larger number of galls was kept in
plastic bags and the kirvae were reared to adults,
Pupation took place within the galls. Plastic bags
Were examined daily and emerged adults preserved
together wilh their pupal skins in 70% ethanol,
Cumida dalsum maunts of type specimens for
mieroscopic examinution were prepiured acearding to
the tevhnique ouilined by Kolesik (1995). The type
series and other material retained tn 70% ethanol dre
deposited in the South Australian Museum, Adelaide
ISAMA| «and the Australian National Inseet
Collection, Canherru [ANIC]. Drivd and preserved
(70% ethanol) sumples of infested and uninfested
plants are deposited in the State Herbarium of South
Australia, Adelitide |SHSA].

Genus Rhapalomvia Ribsaamen, 1892

Rhopalonvia Riibsaumen, 1892: 370

Wpe spectes: Oligetraphis janacelivola Karsch,
1879: VIL Jber. westl, ProyVer Wiss. Kunst: 27 (des
Kictfer. 1896: 89)

Rhopelomyia is a Worldwide sends comprising
species of Oligalrophini with one- or two-seemented
palpi, simple or toothed tarsal chliws, elongate but
entire eighth female abdominal tergite und complelely
setulose gonostylus, The species described bere shares
with Rhepalonvia all the ubove characters with the
exception of having three- or four-segmented palpi.
As the number gf palpal segments is a derived
character and varies within several Oligolrophini
genera it does not prechide the new species Prem
Rhopalumyia.

Rhopalomyia goodeniae sp nov.
(FIGS I-15)

Holoivpes 3, William Creek, South Australia [23°

156

P. KOLESIK

Tilly

16:0:'o|

TEE

Figs l-6. Male of Rhopalomyia goodeniae sp. nov. 1, Genitalia in dorsal view. 2, Sixth flagellomere. 3, Gonocoxites,
parameres and aedeagus in ventral yiew. 4. Head in frontal yiew. 5, Wing. 6. Tarsal claw with empodium. Seale bars =

100 pm 1,3, 4, 50 pm 2. 6; 500 pm 5,

A NEW SPRCTES.OF CECIDOMYIIDAL PROM GUOQDENIA LUNATA a7

45'S. 1467 20° BY, 24.57,1993, P. Kolesik, reared
from larva trom: stem gall of Greedenioe funeta J.
Black, sampled 22.4.1993, 121328 [SAMA],

Paralypess 24 4, V2, 2 pupal skins [SAMA]. 1 d-
12, 2 pupal skins [ANIC], same data; 3 larvae
SAMA], 3 larvae |ANIC]. collected with holotype.

Oher materia? 2 34 09 [SAMA same data: 35
larvae [SAMA], gall [SHSA - AD99511278],
vollecled with holotype,

He scvipnian
Male (Pigs 1-6)

Colour: sclerotized parts al body brown, non-
sclerotized pans of abdoriten orange Wing: total
length 2.7 min (2.5 -2,8, 0 = 4). width 1.0 nim (1-0 -
Lo; RA al distal chd narrower, foore weakly
selerolized and slightly curved posteriorly, joi C
unteriorly lo apex: KI joimog C near wing mid-
length: Se cell strongly sclerotized and together with
KE and adjacent part of RS hearing scales,

Flagellomeres (3 in number, first wind second
ised, with nodes longer thu necks, third to fifteenth
with nodes and necks about same Jeneth, circumtihe
comprising one (rinsverse and one lougitodinal
band, Palpus four or three-segmented with well
developed palpiger in both cases, Tirsal chav simple,
rounded beyond imid-length, empodin as long ds
claws,

Abdonren: all lergiles with pale of setae i anterior
vurners. tergites | - Vio with single setal row
posteriorly and a low serie laterally tergites VIP and
VILL With a few setae scattered in centre: sternites LT -
VITE with pair of setac anteriarly, row of setae
posteriorly and fields of sete both huerally and
contrally

Genitaliy gonocoxites free ventrally. cylindrical,
willy small apicoverttal lobe, setose dnd setulose;
vonustylus stualed dorso-caudiilly of somocox te.
slightly tapering towards (he apex, wilh apical booth
comprising, strong Ghiw and a few firm bristles,
setase dorsally and setulose throughoul: cere
bilohed. with severil stout Setue on cach lobe.
setulose; hypoproct bilobed, with seta on each lobe
selulose; parumeres Clasping acdeagus along their
full leneth, setilose. apically bearing four (a hive
lure, sctose papillue: aedeagus robust, conical.

Femelle (Fiys 79, 11)

Wings towl lengus LO mim (17 © 2). = 2) width
O48 mm (0.7 - 0.9); RS steight and equally strong,
along full lengih as Opposed to being nayawer and
curved posteriorly in mule, Tergite Vifwith setal pow
posteriorly, tergite VIE with single pair of seuie
wuiteriorly. selerotization of both tergiles in shape of
loner “Y". Ovipositor protrusible, cere) fused) inte

stngle. terminal lamella, triangular in dorso-ventral
view, Setose and setulose: hypoproct trapezoid: in
dorsoventral view, beitring [wo setae posteriorly.
selulose. Other characters as in male,

Prpa (Pig. V2)

Colour: won-stlerotized paris of abdomen orange,
the rest brown, ‘Total length 2.7 mm (2.7 -2.8. n= 44.
Integument of abdominal segments covered with
spiculue. Cephalic selerite with two swelligs as
Jong as antennal horns, 30 pir (25 - 36). Cephahe
poirot pupiliae with strong sete. 148 pum (144-152),
Frons on cach side. one of twa lower facial papillae
with seta and one of tree lateral facial papillae with
seta. Prothoravie spirucke 92 - 93 uin mlength, with
trachea reaching its apex, Abdominal segments | -
VIL with pair of setose ventral papillae. two pairs of
setase pleural papillae. pair of setose and two pairs of
aselose dorsal papillae, Abdominal segments H- V I
dorsally wilh field of strong spines on anterior halt,
Abdominal segments VHT aod EX with pair of selose
ventral pupillae, Uwo pairs of setuse pleural papiiiae
and pair of setose dorsal pupillae. Facial papillae
with setae 5-6 pm, papillae on abdominal sediments
with setae 5 - 8 pm.

Larva (Pigs 10. 13, 14)

Colours orange Total length 3.0 min (1.6 - 34. 9 =
6), Inlegument covered with dense spicilae, up ta 1a)
jum long. Head strongly sclerotized, 52 pi (49 - 56)
lone and SS pit (MS - 91) wide. posterolateral
upodemes 75 tm (70 - 79) fong. antennae 17-20 pm
Jone and 8 ~ 10 um wide at buse. Thoracic and finst
sever) ubdominal segments with pair of ventral
papillae. nwo puurs oF pleural papillae andl three pats
of dorsul pupillae. Thoiueie segments with puir of
sternal piipillie and two groups of three tuteral
papillae on each side of spatula, two of each with
selac, one without, Abdonmial segment VILE with
we pairs of ventral papillae, Avo pairs ol pleural
papillae and pair of dorsal papillae. all wilh setae,
Abdominal segment IX beuring four pairs at
tertninal papillae, all wath setae. but same papillic
lackiig i some specimens. Anos ventral. Setae on
sternul and kueral pupillac about | pe, on the other
papillae ¥ - 22 pm, Spatula 294 pon (245-4328) long,
with apical enlargernent (32 pm (116 - 150) 10 width
and 44pm (34 - 48) in length.

hifesratian svaiproaty (rig, 15)

This gall midge Speetes. deforms. the ster of
Goodenia ltd ino subghobular, hairy gulls, J-2
om in diameter and 1-15 eo in henht. Hines are 1-2
indy long, dense, grey. The gall consists of many
vlobular fo subglobular, Unek-walled cells will) one
larva in each cell. Infested plants develop few or nu

P. KOLESIK

158

A NEW SPECIES OF CECIDOMYIIDAE FROM GOODENIA LUNATA 159

Fig. 15, Gall of Rhopalomyia goodeniae sp. nov. on Goodenia lunata J. Black, Scale bar = 20 mm,

flowers. On 22 October 1993 at William Creek,
about 95% of all plants from a population
comprising some 1000 plants were infested,

Enymology
Derived from the generic name of the host plant.

Discussion

Currently, the Rhopalomyia genus comprises 86
Nearctic (Gagné 1989b), 48 Palaearctic (Skuhrava
1986), 9 Neotropical (Gagné 1994) and | Oriental
(Gagné 1973) species. No Rhvpalomyia have been
recorded from the Afrotropical Region (Harris
1980). Until now, only 2 immigrant species of
Rhopalomyia have been recorded from Australasian
and Oceanian Regions with R. californica having
been introduced to Australia from North America

and R. chrysanthemi Ahlberg to the Hawaiian Islands
and New Zealand from Europe (Gagné 1989a).
Rhopalomyia is a catchall genus with the bulk of its
species producing complex galls on Asteraceae.
Species that form galls on other plant families
exhibit some morphological differences and their
placement in Rhopalomyia needs restudying (Gagné
1989b). The new species described here is the first
native Australian species to be placed in
Rhopalomyia and the only gall midge known to
attack plants of the family Goodeniaceae, The
species does not breach the current concept of
Rhopalomyia, except that it has two or three palpal
segments as opposed to one or two segments in other
described members of the genus. Although this
discrepancy precludes identifying the new species as
Rhopalomyia using the most authoritative current
key to Cecidomyiidae (Gagné 1981), I find it
insufficient reason to erect a new genus until more ts
known about its native Australian congeners.

Figs 7-14, Rhopalomyia goodeniae sp. nov.. 7-9, 11 female, 10, 13, 14 larva. 12 pupa. 7. Posterior end of abdomen in
dorsal view. 8. Posterior end of abdomen in ventral view. 9. Sixth flagellomere. 10, Head capsule in dorsal view. 11.
Posterior end of ovipositor in lateral view (cerci shriveled in available specimen). 12. Anterior part in ventral view. 13.
Sternal spatula with adjacent papillae. 14, Two terminal segments in dorsal view. Scale bars = 500 pm 7, 8, 12: 50 pm

9-11; 100 pm 13, 14.

160 PKOLESIRK.

Rhopalomyia goadeniae sp. noy. differs in several
characters from R. califernica, its only Australian
congener redescribed by Gagné & Boldt (1995). In
R. goudeniae, the gonostylus ts straight and about the
same width throughout most of its length, the
papillae on the male parameres are large (1/5 - 1/2 of
paramere width), the number of palpal segments is
three to four and the antennal horns in the pupa are
minute and rounded, In contrast, R. californica has a
gonostylus convex at the posterior end; papillae on
the male parameres are minute (about 1/20. of
puramere width), there are one to two palpal
segments and the antennal horns in the pupa are
elongate and bifid in frontal view.

Acknowledgments

I thank Leta and Philip Gee South Australian
Outback Research William Creek for acting us
guides in the Luke Eyre region and re-collecting the
host plant specimens. | am grateful t0 Mark A,
Adams and Terence B, Reardon Division of Natural
Science South Australian Museum Adelaide for
organizing and leading the collecting trip, Dayid E,
Symon State Herbarium of South Australia Adelaide
courteously identified the host plant species, 7 thank
John D. Gray Department of Horticulture, Viticulture
und Oenology University of Adelaide and Raymond
J, Gagné Systematic Entomology Laboratory USDA
Washington DC USA for commenting on an early
draft of the manuscript.

References

Cooke. D. A. (1986) Family Goodeniaceae pp. 1383-1418 I
Jessop, J.P & Toelken. H. R. (Eds) “Flora of South
Australia, Part 1 (Polemoniaceae - Compositaey” (South
Australian Government Printing Division, Adelaide).

CuNNINGIIAM. G, M., MULHAM, W. E., Minriiorrr, PL. &
Lrigi, LH (lO81) “Plants of Western New South

‘ales (New South Wales Government Printing Office.
Sydney).

GAGSE, RJ, (1981) Cecidomyiidae pp. 257-292 In McAlpine:
J. oR, Peterson, B. ¥.. Shewell, Go E. Teskey, He J,
Vockeroth, J. R. & Wodd, D. M. (Eds) “Manual of Nearctic
Diptera 1° (Cynadian Government Publishing Centre.
Quebec).

(1989) Family Cecidomyiidae pp, 152-163 Jn
Evenhuis, N. L. (Ed.) “Catalog of the Diptera of the
Austalasian and Oceaman Regions” (Bishop Museurn
Press and EJ, Brill, Honolulu).

(1989b) “The Plant-Feeding Gall Midges of North
Amerlea™ (Comell University Press, thaca, New York),

(1994) “The Gall Midges of the Neotropical Region”
(Cornell University Press. Ithaca. New York).

& Bown. PE. (1995) The gull midges (Diprern;

Cecidomyiidae) of Beechwis spp. (Asteraveae) in’ the
United States. Prov. Enionwl. Soc. Wash, 97(4), 767-778,
Harris, K,M_(1980) 18, Family Cecidomyiidae pp, 238-251
Jn Crosskey. R. W. CRd.) “Catalogue of the Diptera of the
Afrotropical Region” (British Museum [Nittural History |,

London ).

kurernr. J.J. (1896) Neue Mittheilungen Gber Gallmiicken.
Wien. Ent. Zty 15, 85-105,

Kowesik. P.(1995) A new species of Eocincticornia Melt
(Diptera: Cecidomyiidae) on Eucalyptus fascicylase in
South Austraha J) Aust. Bay, Sue, 34, 147-152,

McFayprn, P_., DoNNeLLY. GP, & ‘TombLey, A. J. (1983)
Biological control of groundsel bush pp. 28-30 Jit Harvey,
G. J. (Edy “Austulian Weeds Research Newsletter” (The
Alan Fletcher Research Station).

RUBSAAMEN. E. H, (1892) Die Gallmuicken des Koniglichen
Museums fiir Naturkunde zu Berlin. Berl. Ent. 2 37, 314)-
411, pls. VUIEXVIIL,

SKULIRAVA, M. (1986) Family Cecidomyiidae pp, 72-297 Jn
Sods (Ed) “Catalogue of Palacaretie Diptera. Vol. <1,
Sctaridue - Anisopodidae” (Akadémiai KiadG, Budapest).

ROBUSTNEMA FOSTERI SP. NOV., GEN. NOV. (XYALIDAE,
MONHYSTERIDA, NEMATODA), A COMMON NEMATODE OF
MANGROVE MUDFLATS IN AUSTRALIA

By WARWICK L. NICHOLAS*

Summary

Nicholas, W. L. (1996) Robustnema fosteri sp. nov., gen. nov. (Xylidae,
Monhysterida, Nematoda), a common nematode of mangrove mudflats in Australia.
Trans. R. Soc. S. Aust. 120(4), 161-165, 29 November, 1996.

A new genus with a single species, Robustnema fosteri, collected from mangrove
mudflats is described. The circular amphids, six outer labial and six cephalic setae in
one ring, annulated cuticle, single testis and single ovary with posterior vulva, place
the genus in the Xyalidae. The buccal cavity is small, conical and unarmoured, the
lips low and simple. The cardia and ovary are distinctive. Reproductive females are
wide-bodied with a very large uterus filled with developing eggs and unhatched
juveniles.

Key Words: Taxonomy, Robustnema, Xyalidae, Monhysterida, Nematoda, Mangrove.

Trdiasactions af the Koval Sectery ofS. Aust. (1996), 120(4). 161-165

ROBUSTNEMA FOSTERI SP. NOV., GEN. NOV. (XYALIDAE, MONHYSTERIDA,
NEMATODA), A COMMON NEMATODE OF MANGROVE MUDBELATS IN
AUSTRALIA

by Warwick L. NICHOLAS*

Summary

Sicnonas, WL, (1996) Rebustnemne fosteri sp, noy., sen now. (Xyalidve, Monhysterida. Nematoda), a commen
Henatode of mumwrove miudtlats i Australia, Trevi, Ro See S. Adi 12004), 1616165, 29 November, 1996,

A few gents witlia single species. Rebusitema foster’, collected from mangrove mudthats ix described, The
circular umphids.six outer labial and six cephalig setae ia one ning, unnulated cuticle, single testis and single
ovary with posterior vulva, pluce the genus inthe Xyalidue, The bucenl cavity is small, conical and wnarnmdred.
the lips low and simple, The cardia and ovary are distinctive. Reproduetive females are wide-bodied with a Very
large urerus filled with developing exes cud unhatched juveniles.

Key Worbs: Tixonomy, Aehtenienica. Xyalidac, Monhysrerndia, Nematoda, Mingroye.
y ) t

Introduction

A new spewes of nematode that as common in
Mangrove Mudflats in south cast Queensland und
New South Wales ts deserhed, Che possession of
outer labial and cephalie sensiflae ia a single ring,
cireuku aunphids. an outstretched single gonad in
both sexes, a posterior vulva. and) an aonulated
citicle place the new species im the Nyulidie
(Monhysterida), Th was previously misidentified as
Hilipjever sp. (odd and Nicholas L986; Nicholas er
ah WY9L) because of a resemblance to #, cructs
Blome and Schrage 1985. However. more recent
close jnspection has shown that the repraduchive
organs are quile different and this nemtode has been
pluced in anew genus,

Materials and Methods

Type speciiiens Were collected at low tide from
estuarine mud close to mangroves al Pine River
Estuary, whielr apens into Moreton Bay north of
Brisbane. Additional specimens have been collected
from mungroves in New South Wales, Nematodes
were lixedain the mud with 5% formulin immediately
after collection und were recovered from the mud by
uw combination of sieving and centrifugation in
aqueous colloidal sifica (Ludox. Du Pont de
Nemours) with specific gravity adjusted ta 1.2. The
mud was thoroughly dispersed in tip water,
centrifuged at GO00 ¢ lor 7 minutes, the supernatant
discarded, the vesidve re-suspended in colloidal
silica, centrifuged gina at 6000 & for 7 minutes, the
supernatant passed through a nylon sieve with 60 pm

* Division of Botany and Zoology, Austin National University
Canberr ACT 0200.

inesh, wid pematodes retained by the filer back-
washed inlo a petri dish, Specimens of the new
species were picked up with am eyebrow hair
(moutted ob a suck) Under a dissecting: mierosecape
dnd transterred ty 5% aqueous glycerol. After the
water had been allowed to evaporate at 40° C the
nematodes were transferred to fresh anhydrous
glycerol aad mounted on slides with cover ships
supported by glitss beuds (bathing and ringed with
glyceel (Gurr),

Measnrements, given in ym, Were inade from
drawings of 12 specimens using a camera Jucida,
Type specimens are in The South Austrahian
Museum (SAMA) and their numbers in the
Australian Helminth Collection (AHC) are given.

Robustnema sen. ney,

Six inner labral papillae. six ourer labial setae and
six cephalic setae in one ring; circulin amphids. Six
simple low lips, dnarmoured bueeal cavity, cuticle
wonulated. Cardia conver cap to anlernor pair of
intestinal cells. Single gonad in each sex, vulva close
to anus, uterus becomes capacious sac holding
developing cggs and juveniles.

Robustnema fosteri sp, nov.
(FIGS 1-9)

Holouwpe: 3. Pine River Estuary Queensland,
AO.viil. L986, SAMA AHC27695,
Measurements ¢ Table 1.
Deseription of Holotype male

‘Typical nematode form (Fig. 3), curved ventrally in
anal region. cuticle uniformly annulated, Tail
proximally conical. postertad 20% narrow, almost
cylindrical (Fig, 6), Body sctue restricted to five

16?

TAWLE L. Meustirements af Robustiema fostert spn ay

Mates

W. L. NICHOLAS

Sen Females
‘TWpe Holotype Parsitypes n=S Paratypes n=t
Meun sp Range Mean SD Runge
Lonuth }252 1242 29,28 196-1273 }379 178 JU89-1A59
Mix. width 7\ Os 2.80 65-7) Hi4 29 71-161
Cephalhe setae leneth 3,1 53 (149s 17-5. 64 1.06 5.5-4.2
Amphidl diam, 5,9 70 1,05 5.8-8.6 5.2 O55 43-59
Bueeal cavity width a) ta 0.55 1-12 (2.5 1.97 li-14
Bueenl cavity depo ld LK 2.49 16-20 ITB 2.33 {4-20
Head to nerve ring 106 rehe 215 54111 ue) 17.8 K2-| 29)
Pharyox length 22 224 hd 200-254 Daa 27,0 (92. 268
Head to vulva 1OO8 70 805-1220
Head to anus 1040 M47 20,0) 1018-1078 1164 152 905-1304
Tail length 226 1Q7 Va] IKS-220 206 3 [84-226
Width wt anus 57 56 5.27 A9-62 Os S13 54-77
Spicule, ure length 62 At 4.73 5462
De Man's a 17.6 Is 0.680 176-196 j25 2.09 YO-15,3
ae) 5.4 5.6 (374 S.-6.0 3M thou Ae 7
€ 5,3 6A (463 6 1-6.7 67 1.60 57S
v 40 35 (420 Vea. | 0,43 LO-88
V % ") 4.00) TAKS

pamed, ventral sete. evenly spaced between anus
und tailuip, + pm long, plus three terminal 4 yin long
setae. Six stout lips prehed over buceal cavity
forming shallow dome arising from very short
parallel-sided region, demarcated from cervical
region by strong annular groove (Fig. 1). Sie minute
inner labial papillae on lips; six short outer labial and
six equally short slightly wider cephalic -setac
inserted side by side at base af lips; amphids circular,
situated at level of buccal cavity. Bueeal cavity small.
simple, conical, without teeth, cuticular ridges or
denticles, Pharynx cylindrical without muscular
bulb, cardia with about 12 small cells forming
vonvex cap to anterior pair of intestinal cells (Fig. 4).
No renette cell, “exeretory’ duct or pore. Intestinal
cells large, paired. with prominent nuclei and
evanular cytoplasm, rectum very short, about 30 pum
long. Testis single. outstretched, to lett of intestine,
extending anteriorly almost to cardia, long seminal
vesicle filled with rounded spermatozoa, extending
posteriorly a little beyond mid-body, long vas
deferens, very short ejaculatory duet (Fig. 3). Short
strong spicules, are length 62 um (hig. 7), with
cupilulum, shallow angular bend hallway along,
extreme tip forms narrow peg; gubernaculum two
slightly curved rods; two very small pupilliform pre-
anal supplements. 46 and 77 pm anterior to anus
(Fig. 6). Three posi-anal caudal glands,

Paratypes, From Pine River Estuwry Queensland,
O.vill. JY86, SAMA AHC27060- 27665,

Meusurements; Table |,

Five d 2 essentially similar to holotype. The tip ol
the tail is not necessarily bent dorsally as itis in the
holotype. Pre-anal supplements very difficult to find
in some males when they do not protrude in profile.
Five 2S of much stouter build than males, but not
alLarched dorsally as in paratype illustrated in Fig. 2.
Uterus forms a large thin-walled sac occupying more
than half the body length. The uterus is filled with
developing eggs anteriorly and fully-formed
unhatched juveniles posteriorly (Pig. 5). There may
he as muny as 28 juveniles and embryonated eggs.
The ege shell is not rigid and it accommodates the
shapes of the developing embryos, which are ubout
30-40 um long. Juveniles. curled within the thin
flexible shell are 150-200 yim long. Three lo six
granular cells, probably unfertilised ooeytes, lic
along the anterior dorsal walbot the uterus.

Seunning electron microscopy (Figs 8,9) confirms
that there are six cephalic setae, inserted beside the
corresponding outer labial setae, The cephahe setae
are wider at their bases than the ouler labial setae.
Figures 8 und 9 show that the six lips are deeply
incised and that the head is hexagonal when viewed
en face

Figs.1-7. Rabusinema foxteri sp. nov, |. Male head, 2. Gravid female, 3, Enite mate. 4 Cardia. 5, Sra) portion af annie
lated cuticle and uterus containing unhatched juveniles. 6. Male dail. spreules, caudal glands, setie and pre-anal supple
ments, 7 Apicules and gubernaciuny. 1.4.0 and 7 holotype male: 2. 3, and 5 partiype mate and female AHC27661

ROBUSTNEMA FOSTERI SP. NOV., GEN, NOV. 163

ee LT 5 ee 2S5um
10 ees SL
Ses 10 7 25,
4 es 10um

W.L. NICHOLAS

Tiga, 8 nning electron m raphs of head of Rebusineme fastert sp. oy. ols outer labial seta. cs cephalic seta, am
wophid, ip inner labial papilla. Ip lip

ROBUSTNEMA FOSTERI SP NOV., GEN, NOV. 165

Differential diagnosis

The presence of six cephalic setae in the second
ring of sensilla, while not unique, is unusual in
Xyalidae as is the small size of the unarmoured
bueeal cavity and low profile of the lips. The shape
of the cardia and ovarian development are quite
distinctive within the Xyalidae. Together these
characters justify generic status.

Habitat
Mangrove mudflats.

Distribution

Pine River estuary, opening into Moreton Bay
Queensland; Fullerton Cove opening to the Hunter
River estuary New South Wales and the Clyde River
estuary New South Wales.

Etymology

Generic name from the strongly built body,
especially the very stout-bodied reproductive female:
specific name after a colleague.

Discussion

The female reproductive system is unusual and its
development warrants further study. Only a
discontinuous developmental series has been
observed. In immature females, the outstretched
ovary 1s filled with small cells, presumably oogonia.
Alter fertilisation, large eggs in early cleavage stages
appear amongst the small cells. By this time the
ovary extends forwards almost to the level of the
cardia. Later the ovary is largely transformed into a
capacious uterus filled with developing eggs and,
towards the posterior, unhatched juveniles. Some
large granular cells, with single nuclei, lic along the
dorsal wall of the uterus, These large cells are
probably unfertilised oocytes and oyarian
development can be described as hologonic rather
than showing the much more usual telogonic
development, No spermatozoa were recognised
within the female gonad.

Acknowledgments

1 am grateful to The Australian Biological
Resources Study for travelling expenses,

References

Brome , D. & Scurace, M, (1985) Freilebende Nematoden
aus Antarktis. Verdff Inst. Meeresforseh, Bremerh, 212,
71-96.

Hoppa, M. & Nicnionas, W. L. (1986) Temporal changes in
hittoral meiofauna from the Hunter River Estuary. Aus’. J.
Mar. Freshy, Res, 37, 729-741.

NicHowas, W. L., ELek, A, C., Stewart, A.C. & MARPLES,
T. G. (1991) The nematode fauna of a temperate
Australian mangrove mudflat; population density,
diversity and distribution. Hydrobiologia 209, 13-27

A REDESCRIPTION OF ASPERSENTIS ZANCHLORHYNCHI
(JOHNSTON & BEST, 1937) COMB. NOV.
(HETERACANTHOCEPHALIDAE:ACANTHOCEPHALA)

By LESLEY R. SMALES*

Summary

Smales, L. R. (1996) A redescription of Aspersentis zanchlorhynchi (Johnston &
Best, 1937) comb. noy. (Heteracanthocephalidae: Acanthocephala) Trans. R. Soc. S.
Aust. 120(4), 167-171, 29 November, 1996.

Aspersentis zanchlorhynchi (Johnston & Best, 1937) comb. nov., occurring in
Zanchlorhynchus spinifer, is redescribed from specimens collected off-shore from
Macquarie and Heard Islands between 1986 and 1990. Aspersentis zanchlorhynchi
can be distinguished from all other species in the genus by having a cylindrical-
shaped trunk, and proboscis armature of 14-16 rows of 10-12 hooks of which both the
larger ventral and smaller dorsal hooks have roots. An analysis of the literature
indicates that the genus Aspersentis comprises four species A. austrinus, A. johni, A.
minor and A. zanchlorhynchi.

Key Words: Acanthocephala, Aspersentis, Antarctic, fish hosts.

Treinsadrions af the Raval Soeteny af S. Alive, (1996). LAK A), 167-171.

A REDESCRIPTION OF ASPERSENTIS ZANCHLORHYNCHI
(JOTINSTON & BEST, 1937) COMB. NOV.
(HETERACANTHOCEPHALIDAE: ACANTHOCEPHALA)

by Lesity R. SMALES*

Summary

SMAnES, LAR (1996) A redescrption of Aspenvends sancllurinent (Johnston & Best) 1937) comb. ney,
(Heteneanthocephalidae: Acunthocephila) Trams. Ro bee, 8, Angi $20 (4.067171 29 November, 199A.
Aspersentis came hlerivacin Wohnston d& Best. 1937) comb. nov. occurring in Zaneiloriviehtis spiniier, is
mudeseeihed Hon specimens collected off-shore fom Macquarie and Heard Tslands between 18a and 1990.
Axperseutis canchlarhynch’ can be distinguished Crom all other species in Ure genus by luvin a eylindncal
shaped (rurik, and proboseis armature of (4-)6- rows of 10-12 hooks of which both the huwger ventral and sniler
Hors hooks five roots, Anainalysis of the Gteratune indicus tharthe genus Aspenenrs comprises lourspegies

A qusseinis |, gato’ A miner and A. sancilorivicdi.

Key Wokbs: Adanthovephala. Aspersenis, Antiretic fish hosts

Introduction

Avanthocephalan material, collucted by the
Australisiin Antwetlc Expedition (AAE) at 19tl-
1914, was the subject of a report by Johnston & Best
(1937). In that report they described a new species,
Lehinechynchuy sanchlorhyncht. occurring in the
stomach of a seorpaenid lish, Zanehlorhvachus
ypoafer Gtinther, from’ Macquarie Island, Since ieir
description was based on aw single female with iis
proboscis nor fully everted. Johnston & Best (1937)
indicated that the examination of additional
specimens woulil be required to confirm the species,
its description and its laxonormie position,

Subsequently, lwo immature specimens (one male,
one female) were found in 4. spiniler collected ut
Macquarie Island during the British Australian and
New Zealand Antaretic Research Expedition
(BANZARE) of 1929-31, and identified as A,
suarlilorhynchi by Edmonds (1957).

More ucaithocephalans were tound when
members of the Australian National Antarctic
Research Expeditions (ANARE) of LY86 - [99D
collected Z. sptaifer trom Muaequane and Heard
Ishinds, Examination of this material has allowed a
more complele description of the aeunthocephalan to
be prepared. These specimens, whilst conforming to
the general deseriptions of Johnston & Best (1937)
und Edmonds (1957) bad asymmetrical proboseis
armature and) spines oon the trunk, features
characteristic of the Aspersentinae, The siznificunce
of these morphological data are considered in this
paper and an analysis of the current status of ibe
pends Aspersentiy 5 piven,

“University of Central Queenstind, Rockhampton Old
1702,

Materials and Methods,

Thirteen Zanchlorhynehus \pinifer collected from
Macquarie Island witlers (84° 33°75. 158° 53" E) ane
one from the Heard Island shelf (trawled hetween
51? 34h and 53° 30'S, 72° and 78° 00'E) were fixed
in 10% Tommalin bulferad with exeess sodiun
tetraborate. Fish were then examined under ut
dissecting Microscope and yoy acunthocephalians
found were stored in 70% ethanol prior bo
oxumination, cither as temporary wer mounts, after
cleuring in becehwood creosote. or as permanent
preparations, afler staining in Crenacher’s cuarmine
alum. delydriting through aeraded series of ethanol,
clearing in xylene and mounting in Canada balsam,

Measurements of (0 mates and 10 females were
made with the did oF an ocular micrometer, drawing
tube and measuring wheel and ave given in pny unless
otherwise shiled, with the range followed by the
mean in parentheses. Figures were drawn with the
aid of a drawing tube. All specimens dive been
deposited in the Queensland Museunt (QM)

Systematics

Order Palacacunthocephiala Meyer, 1931

Family Heteracanthocephalidae Petrochenko, 1956
Sublumily Aspersentinae Golvun, 1960

Genus Asperseuty yin Cleuve, 1929

Type species Aypersenity anstrinuy Van Cleave,
1920,

Aspersentis zanchlorhynehs (Johnsion & Best,
1937) comb, noy
(MGS 1-7)

Synonvin Evhinorhvnchis sdnchlorhvnehi
Johnston & Best, 1937 pp. 12-13; Edmonds, 1957 p
96, Es, sanchlorhyneh? Zdzitowieckt, Y86 pp, SY,
102, fable tf,

o>
ao

LR, SMALES

=

SSSA J

SS

=
mK

SF RV TEP I?
= HOS

GS Mit Maa

AN
Pit

i

Figs 1-7 Aspersentis zanchlorhynchi comb. noy, Fig. |. Proboscis armature, one row of dorsal hooks. Fig. 2. Proboscis
armature, one row of ventral hooks, Fig. 3. Proboscis, dorsal view. Fig. 4. Male. Fig. 5. Fenvale, proboscis not fully evert-
ed, showing distribution of trunk armature. Fig. 6, Posterior end, female, Fig. 7. Egg. Scale bars = 50 pm, 1, 2: 150 pm.
3, 100 um. 6; 500 pm, 4, 5: 25 pm, 7.

A REDESCRIPTION OF ASPERSENTIS 2NNCHLORHYNCHT 164

Material examined

From Zanclilorlivnchuy spiniter. 126 99, 170
aod. Macquarie Island, 6.1).86. 12.vi.86, 6.x11.56;
G211324-G211335, 19 Heard Island, tt.yi.90;
G211325

Revived deyeription

‘Vrunk cylindrical. Proboscis long. cylindrical. set
at angle to trink (Pigs 4, 5) Proboseis armuture
siinilac in bollr sexes, 14-16 paws of 10-12 hooks
(Fig, 3); dorsal rows of hooks (Pie, 1) soniewhiat
smaller thin ventral rows (Pig 2): all hooks with
roots, Neck Short unarmed, truteated, Spines tiny,
embedded in cuticle in both sexes, encircling anterior
cndhel trunk toa level about halfway down proboscis
receplacle, then extending down literal truak to
posterior (Pig. 5). Proboseis reeepiacle double-
walled, imserted at base of proboscis; ganglion
phiced Neat posterior end, Lemnisei flat. longer than
proboscis receptacle when fully extended,

Male: Trink 3.46 (4.2) 1m long by 360-680
(480) wide, Proboscis, not fully extended fii most
specimens, 650 long by 215 wide (n=l), Largest
dorsal hooks Sed and tthe in row, 33-63. largest
venti hooks. 3rd and 4th in row 76-85, Neek |3a0-
195 (145) long by 140-260 (175) wide. Proboscis
receptacle 615-995 (745) long, Lennise’ 740 <1 300
(930) long. Testes ovoid, tandemly placed; anterit
testis 455-985 (675) long by TRO - 300 (263) wide,
posterior testis 490-715 (635) long by 195-445 (280)
wide. Cement glands, sis, pear-shaped. Mule
aperture lenin,

Female: ‘Trank 5.3-16 (10)mm long by 390-765
(500) wide, Proboseis, not fully extended, longer
than 900, width 200. Largest dorsal hooks, 3rd avd
4ih in row. 50-545 largest ventral hooks, 3rdund 4th

in row 80-83. Neck 182-227 (200) long by 175-260
(205) wide. Proboscis receptacle 810-1300 (1160)
long: lenmiser 925-1940 long (n=2), Pentale aperture
terminal (Piz. 6). Bggs embryonated. with
prolonpgations af middle shell 75-90 (87) long by Th
wide (Pir. 7).

Host! Zunchlarhvnichiuy spinijer Giinther,

Location: stomach, intestine.

Locality: Macquarie Phan,

Type specimens: Holotype female. South Australian
Museum V 2200.

Remarks

Mibough a large nuntber al specimens wis
vollected, none of them had been relaxed and
extended prior 10 fixation, which made then difficult
to study. Comparison of the material from this study
wilh the deseriplions of Johnston & Best (147) and
Edmonds (1957) showed that all the pnaterial
colleeted from #4. spinifer was the sume species, tn
Muny specimens the leninisci were vs described by
Johnston & Best (1937) that is, short abd irregular
and reaching uboul one-third the length of the
proboscis receptacle but in ihe more relaxed
specimens. the lenmiser were flit and extended
heyond the proboseis receptacle. The cylindrical
shape ol the proboscis und its armature, be 16 rows
of 10-12 hooks, were observed in the three
specimens described by Johmston & Best (1937) and
Rdmonds (1957) but in none of them had the
proboscis extended far enough to describe the
morphological details of the proboscis hooks. The
asvinnetry of fhe armature, ventral hooks bemg
Jarger than dorsal ones, could be seen only in those

TAWLL L. A compertsanaffeniale body measurements af Asypersentis wustinitts Van Cleave, (929 (aiken from
Zdcitawieck MYST) AL miner Belmunes & Sates, 1992 and A. vanehlorhyneh otnsten & Best, 1937), Measurements

in mm.

AL QUStris
South Shetlands
4.93-6,42 (S74)
116-1.79 (1.a9)
0.5 0.66 (0.59)

(runk length
trunk width
proboscis lengthy
proboseps width 029-042 (41)
dorsal hook length (maximum) 0.05-40.064 (0.060)
ventral hook length (maximum) G1 19-0. E37 (0.426)
neck leneth (17 4.23 (ON 2y
ewe (.060-0,088

; 4 0,019-0,025
hook disposition

South Georgia
6Y4-K 54 (7.15)
1,092.00 (173)
(07-073 (O70)
(29-4, 35 (0.42)
1060-4650 (0,062) O.030-04135 (104132) 0.050-0,056
0132-0, 149 (0.140) 0,062-0,080 (0.065) -0.080-0,083
(W224 Sb (UL27)
(1.07 1-0.087

«® O.0270-0,025
13-16 rows Of 7-LL hooks/row

AL minge
Tasmania

Tas (8.

AL Sunelilochiynchs
Macquarie & Pedra ts
5.3-16.0, (10.0)

31-095 (O54) 0.39-0L76 (O50)

O.24-0,42 (28) 20,90

TOLL? (0.14) 1.20

0,120.25
(,068%-0.077

O.18-0,.23 (0.20)
().073-0,090 (087)

x OOI2O16 A OAS
14 rows of 7-9 14-16 rows of 10-12
hooks/row hooks/w

70 L.R, SMALES

specimens with the proboscis almost, or completely,
everted, This character is indicative of the genus
Aspersentis rather than the genus Echinorhynchus to
which the species was originally allotted, Somatic
armature, present in this species, is also found on
other species of Aspersentis (see Zdzitowiecki 1981,

1986) but not Echinorhynchus. Since tiny spines are
easily overlooked, as has occurred in) some
collections of A. austrinus (see Zdzitowiecki and
Rokosz 1986). it is not surprising that they were
undetected in the earlier studies.

Aspersentis zanchlorhynchi (Johnston & Best,
1937) comb. nov, can be distinguished from A,
austrinus Van Cleave, 1929 in having a more
cylindrical trunk, a longer proboscis, at least 650 in
males and 900 in females, compared with up to 630

in males and 720 in females, in the distribution of

trunk spination on the lateral unk as well as
encircling the anterior trunk, and less marked
asymmetry of the proboscis armature with both
ventral and dorsal hooks having roots. Aspersentis
sanchlorhynchi bas more hooks per row (10-12) than
does A. austrinus (7-11) on the proboscis (Table 1).

In comparison with A. miner Edmonds & Smales,
1992, females 3.2 mm, and A. jo/mi (Baylis, 1929)
Chandler, 1934, females 3.0 mm, A. canchlorhynehi
is much larger, females 10 mm, With LO-12 hooks
per row on the proboscis, A. canchlorhynchi has
more hooks than A. miner, 7-9, and fewer hooks than
A, jolni, 12-14. Aspersentis minor occurs. in
Rhombasolea tapirina trom Tasmanian waters
(Edmonds & Smales 1992) and A. jo/ini occurs in
Merluccius sp. around the Falkland Islands
(Yamaguti 1963). By contrast A. zanchlorhynechi
occurs in Z. spinifer from Macquarie and Heard
Islands, that is from) sub-Antarctic and Antarctic
waters.

Discussion

In a redescription of A. austrinus Zdzitowiecki
(1981) placed Rhadinorhynchus wheeleri Baylis,
1929, Aspersentis wheeleri Chandler, 1934 and
Aspersentis megarhynchus (Linstow, 1892) Golvan,
1960 nec Echinorhynchus megarhynchus Linstow,
1892 as synonyms of A. austrinus. He commented
that Linstow (1892) gave the number of proboscis
hook rows as 18 and described the trunk as unarmed
and that Linstow neither measured, described nor
drew the ventral and dorsal rows of hooks as having
different shapes and dimensions. All of these
characters are inconsistent with the genus
Aspersentis. Amin (1985), however, in his
classification of the Acanthocephala overlooked
Zdzitowiecki’s paper and followed Golyan (1960) in

listing A. megarhynchus (Linstow, 1892) with A.
austrinus as its synonym and A. johni (Baylis 1929)
as the only two valid species in the genus.

Zdzitowiecki & Rokosz (1986) re-evaluated the
validity of Heteracanthocephalus hureaui Dolltus,
1965 and concluded that it was either a synonym of
A. austrinus or, because of the wide range of number
of hooks per row, of A. johni. Zdzitowiecki (1986) in
his systematic review of Antarctic acanthocephalans
reaffirmed his conclusion that Echinorhynchus
mevarhynehus Linstow, 1892 did not belong in the
genus Ayperyentix, and listed H. hureaui as a
synonym of A. austrinus. Then Zdzitowiecki (1990),
when re-examining material previously designated
H. hureaui, stated that, “as was suggested earlier H,
hureaui is identical with A, austrinus = A,
megarhynchus”. Edmonds & Smales (1992) noted
the inconsistencies in the designation of A. austrinuy
by Zdzitowiecki (1981, 1986, 1987, 1990) and
indicated that EL megarhynchus as described by
Linstow (1892) lacked body spines and did not have
asymmetric proboscis hooks.

Thus, A, megarfiynchus as redescribed by Golvan
(1960), is not the same species as E. megarhynchus
Linstow, 1892 because it has asymmetric proboscis
armature and cuticular spines on the trunk, and 1s
now considered as A. austrinus. Since A. austrinis
was described in 1929 by Van Cleave while A.
megarhynchus was not established until 1960 by
Golvan, A. austrinus becomes the type species of the
genus Aspersentis with A, megarhynchuy us a
synonym. The other synonyms are A. wheeler
(Baylis, 1929), Rhadinerhynchus wheeleri (Baylis.
1929) and Heteracanthocephalus hureaui (Dollfus,
1965).

The genus Asperyentiy therefore now consists of
four valid species A. austrinus, A, johni, A. minor and
A, zanchlorhyncnt.

As to Echinorhynchus megarhynehus, Johnston &
Best (1937) suggested that it might be identical to
Leptorhynchoides debenhami (Lieper & Atkinson,
1914) Johnston & Best, 1937 now Metacanthocephalus
rennecki (Lieper & Atkinson, 1914) Zdzitowiecki,
1983. Echinorhynachus megarhynchus occurs in Notothenie
corriceps, one of the hosts of Metacanthocephalus

Johnstoni Zdzitowiecki, 1983, but not in Trematomus

bernachii the host of M. rennecki (sce Zdzitowiecki
1983). Moreover the original description of F.
megarhynchus by Linstow (1892) resembles that of
M. johnstoni in having a proboscis armature of 18
rows each of 6 hooks, the proboscis 0.45 mm long,
and apparently no neck. Of the other species
occurring in Antarctic fish hosts, Echinorhynchus
petroschenkoi (Rodjuk, 1984) Zdzitowiecki, 1989, is
a larger helminth than E. megarhynchus with a
longer proboscis, proboscis armature of 15-19 rows

A REDESCRIPTION OF ASPERSENTIS ZANCHLORHYNCHI V7

of 10-13 hooks, and does not occur in N. corrticepy
(see Zdzitowieckt 1989). Heterosentis heteracanthus
(Linstow, 1896) has body spines and proboscis
armature of only 10 rows of 4-5 hooks, with a
striking difference between the length of the first two
and the last three hooks (Zdzitoweicki 1984). These
characters suggest that EL megarhynachus is closer to
M. johustoni than any of the other acanthocephalan
species occurring in) Antarctic) fish, Direct

examination of specimens of M. johnsenti is needed
before a determination on the status of 2.
megarhynehus can be made.

Acknowledgments

My thanks to Dr R. Williams who collected the
parasites and Professor K. Rohde who made them
available.

References

Asin, ©, M, (1985) Classification pp. 27-72 de Crompton.

D. W. T. & Nickol, B. B. (Eds) “Biology of the
Acanthocephalu” (Cambridge University — Press.
Cambridge).

Epmonps. S. J. (1957) Acanthocephala, B.A.N.Z.A.R.E,
1929-31. Report ser BL 6, 91-98.

— & SMaALES. Le R. (1992) A new species of
Acanthocephala from the Greenback — Flounder,
Rhombosolea tapirina Giinther, 1862. Tras. RK. Soc. 8.
Aust, 116, 35-38,

Govan, Y. J. (1960) Le phylum Avanthocephala,
‘Troisigme note. La clusse des Palacacunthocephala
(Meyer. 1931), Annly, Parasitol. hum, comp, 35, 138-
1045.

Jounston, T. AL & Bust, FL W. (1937) Acanthocephala.
A.A.B. 1911-14. Report ser, C. 10. 5-20.

Linstow, O. VON (1892) Helminthen von Siidgeorgian.
Nach der Ausbeute der deutschen Station von 1882-
1883. Jechrb, Hamb. wrssen. Anst. 9 LO-77.

YAMaAGurl, S. (1963) “Systema Helminthum™ Vol. 2
Acanthovephala (John Wiley & Sons, New York).

Zpzrrowleckl. K, (L981) Redeseription of Aspersentiy
austrinus Van Cleave, 1929 (Acanthocephala). Acta
Parasitol. Pol, 28, 73-83.

O(N 983) Antaretic deanthocephalans of the
cenus Mefacanthocephalus. lbid. 28, 413-437.

— (984) Desersprion of Heferasentis
heteracanthuys (Linstow, 1896) from Antarctic fishes, and
remarks on the taxonomic stutus of Hererosentiy Van
Cleave. 1931 (Acanthocephala. Arythmacanthidae ). /hiel.
29. J 1-115.

— (1986) Acanthocephala of the Antarctic.
Polish Pol. Res, 7. 97-117.

(1987) Acanthocephalans of marine fishes in
the regions of South Georgia and South Orkneys
(Antarctic). Acta Parasitol. Pol, 30, 211-217.

(1989) A redescription of Echinarhynchus
petrotschenkoi (Rodjuk, 1984) comb. n. (Acanthocephala).
Ihid, 34, V73-180.

(1990) Re-examination of five Antaretic and
subantaretic digenean and acanthocephalan species from
Professor Svidat’s collection, (hid. 35. 31-30.

— — & Rokosz, B. (1986) Prevalence of
acanthocephalans in fishes of South Shetlands
(Antarctic). I Aspersentiy austrinus Van Cleave, 1929
and remarks on the validity of Heteraventhocephaluy
finreaui Dollfus, 1965. Thid. 30, 161-17),

SEASONAL ACTIVITY OF THE EARTHWORM,
GEMASCOLEX LATERALIS (MEGASCOLECIDAE), IN A
EUCALYPTUS WOODLAND IN SOUTH AUSTRALIA

BRIEF COMMUNICATION

Summary

The earthworm fauna of agricultural habitats, especially pastures, has been
extensively surveyed in south-eastern Australia in recent years'*. The fauna is
dominated by accidentally introduced species, particularly Lumbricidae from Europe
(e.g., Aporrectodea caliginosa, A. trapezoides and A. rosea). Native species are rare.
Very little is known of the biologies of native earthworms, either in urban,
agricultural or native habitats**'', This brief communication reports on the seasonal
activity of Gemascolex lateralis (Spencer 1892) (Megascolecidae), one of the most
common native species in South Australia, and offers one possible explanation for its
rarity in pastures.

Transactions of rhe Royal Seviety of 8, Aust. (1996), 9120/4). 173-174,

BRIEF COMMUNICATION

SEASONAL ACTIVITY OF THE EARTHWORM, GEMASCOLEX LATERALIS
(MEGASCOLECIDAE), IN A EUCALYPTUS WOODLAND IN SOUTH AUSTRALIA

The earthworm fauna of agricullural habitats, especially
pustures. has been extensively surveyed in south-eastern
Australia in recent years'4, The fauna is dominated by
accidentally introduced species, particularly Lumbricidie
from Europe (¢.2., Aporrectoded culiginasa, A, trapecoides
and A. rovect), Native species are rare. Very little is known
of the biologies of native earthworms, either in urban.
agriculural oor native habits"! This brief
communication reports on the seasonal activity of
Gemeascvolex lateraliy (Spencer 1892) (Megaseolecidie).
one of the most Common native species in South Australia,
and offers one possible explanation for its rarity in pastures.

Engelbrook National Trust Reserve is an open forest
(Eucalypiuis obliqua - E. baxter’) at Bridgewater in the
Mount Lofty Ranges, SA (350 m altitude. Grid reference
962224 on Sheet 6627-1, Echunga, [ ; 50000), The

understorey of the reserve is composed af shrubs such as
Banksia, Acacia, Hakea, Leprospermum and Hibbertia spp,
The soil is a yellow podzolic. Average annual rainfall is
1050 mm. Much of Engelbrook Reserve has been burnt by
bushfires in recent years but the sites reported here had not
been burnt for at least 35 years at (he Ome of the study.
Baker!? 4 seteighty pitfall traps (plastic jars. 9 cm diam.
9 vin deep) flush with the sail surface within Engelbrook
Reserve in March 1983. Forty traps were set on a north-
west slope and the other forty on # south-west slope. The
traps were set LO) im apart in two transects on each slope.
Each trap was covered by a ceramic tile (15 4 15 em), set
approximately 2 ci ahove the traps on three nails. These
tiles prevented rain and leaf litter from fouling the traps, No
preservative was added to the traps, The taps were
inspected weekly until March 1984 and the mvertebrates

120

60

|

50

40 -

Numbers of Earthworms

10

A M J J A
1983

-|100

Weekly Rainfall (mm)

i

i (ATT

S O NN, Bye dy <F’ ™

1984

big. 1. Numbers of Gemascole. lateralis tapped per week in Engelbrook Reserve, SA dunny 1983-54 (bars). Weekly
rainfalls, recorded nearby at Stirling Post Office. are included (line),

174 G.H. BAKER

thar were Cuught Were preserved for later counting,

One species of earthworm. G, lateralix, was trapped,
mostly in the summer months (Pig. 1). There was ne
discernible difference belween the catches on the two
slopes (t= 1.42, p > 0,05 for a paired t test comparing total
weekly catches for the two slopes), The dala for the two
groups Of traps have therefore been combined in Mig. 1.
There was no apparent relationship bewwveen the peak in

tapped earthworms in December and prevailing weather

(eu. nein) wt Chat lime (r= - 0.239, p > O05 Tor weekly
data for? November 1983 ta 21 March 1984),

The activities of carthworms, on and neur the soil surface,
ie usually associnted with cool, wet weather in temperate
and mediterranean habitats! In) particular, Abbott’
reported that the activity of native earthworms ina jarrah
forest in south-western WA was restricted te winter and
spring when soil nioistite was highest, Tis therefore
suirpeisig thatthe surkiwe activity of G. dareraliy peaked in
summer at Engelbrook Reserve when Weather was at its
hottest and driest. However, Linwson!! found Go hireralts
Throughout (he yearin the surface lavers of the sail (0-5 cn
deep) ina Eucolyius Woodland near Cape Jervis, SA, She
argued that, because of the predominance of clitellite adults

‘Baker, GIL, Buckerfield, J.C., Grey-Gardner, R,,
Merry, R. & Doube, B.M. (1992) Soil Biol, Biochem, 24,
1389-14995.

‘Baker, GAUL, Barrett. VJ., Williams, P.M.L., Carter, PJ.
& Buckertield, J.C. (1993) “Distribution and abundance ol
Gurthworms in south-eastern Australia and their influence
on the burial of lime” pp. 345-348 fn Corey. §.A., Dall, Dud.
& Milne, W.M. (Eds) “Pest Control and Sustainable
Agriculture” (CSIRO, Melbourne).

‘Baker, GIL, Thumlert, T.A.. Meisel, L.S., Carter, Pu.
& Kilpin, G.P. (1996) Soil Biol. Biochem. In Press
“Baker, GLH. (1996) “The ccology, management and
benefits of earthworms in agricultural soils. with particular
reference to southern Australia Ja Edwards, CA, (Eu.)
“Earthworm Ecology’ (Soil and Water Conservation
Society of Amertea, Ankeny), In Press.

‘Wood, TG, (1974) J. Anim, Bent, 43, 87-106.

"Abbott, L (1985) Aust. J. Soil Kes, 23, 264-270.
"Abbott, E. (1985) (bid, 23, 271-290.

al this me. G. lateralis reproduced during. the hotter,
warmer months of the year in this woodland, She did not
find cocoons (at uny fine) to verily this conclusion, Tn
addition, Lawson could not find G. fareraliy ina nearby
pasture. Baker er af." did find small numbers of GF, faverdtis
in other pastures in the Mount Lofty Ranges. lit these
pastures, G fafereliy was present in the top LO ent of soil
from autumn to spring bul retreated lo greater depths in the
simmer inonths, Most worms were found in. patch within
one pasture near a clump of Lucalvprus wees and: taller
lows,

Why G. leferaliy is present. indeed most active, on the
soil surfuce in native woodlands during summer is not al all
clear. Apparently, the shelter provided by the above-ground
vegehition in woodlinds provides sufficient moisture ane
cool temperatures to enable such behaviown, But in open
pastures, Ue seems Likely that the hick of similar vegetation
would prevent this summer surface activiry and henee
reduce the abundance of G) fafendlis. (itis lo survive in
pastures at all through summer. G. keeralis must retreat
from surface layers to greater depths where conditions wre
cooler and moister as do the more abundant. introdueve
lumibrierds 6,

‘Van Praagh, B. (1992) Soil Biol. Biochem, 24, 1363.
1367,

‘Baker, G.H., Barrett, V.J., Grey-Gardner, R. &
Buckerfield, J.C. (1993) Trans, R. Soc, S. Aust. 117. 47
53.

“Lawson, L.M, (1993) “The Distribution and Abundance
of Native and Introduced Barthworms in an Area of Pasture
und Native Vegetation near Cape Jervis, South Australia”
PhD Thesis, Flinders University of South Australia.
"Abbott, E(1994) Aust. J. Soil Res, 32. 117-126.
Baker, G.H. (1986) Trans. RK. Soe. S. Aust. 110, 43-48,
Baker, GH. (198%) Aust. Ent. Maw. 15, 127-139.

Mhee, RYE. (1985) “Eurthworms, Their Ecology ancl
Relationships with Soils and Land Use" (Academic Press,
Sydney).

“Bouche, MB, (1972) “Lombriciens de France. Eeologic
et Systematique” (INRA, Paris).

"Baker, G.H,, Grey-Gardner, R. & Buckertfield, J.C,
(1992) Aust. J, Ecol 17, 177-188.

G.H, BAKER, Division of Entomology, CSIRO PMB 2 Glen Osmond $.Aust. 5064.

GASTRO-INTESTINAL PARASITES OF FERAL CATS
IN THE NORTHERN TERRITORY

BRIEF COMMUNICATION

Summary

The feral cat, Felis catus, is well established as a predator in Australia and feeds on
insects, fish, amphibians, birds, reptiles and native and introduced mammals’. Several
surveys of the gastro-intestinal parasites of feral cats have been carried out in south-
eastern Australia”**”, the species and prevalence of the parasites encountered varying
between states, depending on available food sources and climate. In this study we
present information on the gastro-intestinal parasites recovered from feral cats
collected from the Northern Territory, a region from which only limited data currently
exist.

Trevmactiony af de Bayal Ancien of S Anse (900), E2004, EPS bo:

BRIRF COMMUNICATION

GA

STRO-INTESTINAL PARASITES OF FERAL CATS

IN THE NORTHERN TERRITORY.

The feral cat. Felir cettiy. ie well established asa predinton
in Austiilia and Tees on insects, fish, ampbibiins, birds,
reptiles qnd onutive und infrodieed miaamals! Several
suVeVvs OF The wastro-intestinal parasites of feral cabs hive
been eairried out in southeastern Australie, the species
dnd prevalence of the parasites encountered varying
beoween stutes, depending on available food sourees and
climale. WWothis stiely we present information on the eststra-
intestinal parasites recovered fram feral cats collected (on
the Northern Verritory. a region from which only tnmteal
ibiln euerenitly GN esL.

Stall from the Parks and Wildlife Commissien of the
Northern Territory Gapped and/or shot (88 feral ests tor
this stody between 1991 and 1993, Twenty-two cts
orivinited Cron Witarrka (Rings Canyon) Nutinasil Park
souhewestal Awe Splines (24° 20 and 25° 20'S. 130° 30!
(192° 45' Ey. 25 from an area norih of the MacDonnell
Ranges team Glen Helen Gorge to Yarnbah und Alcoote
Stultons (22) 30% 23° a5 8, 133" ale 134 S57 Rs
neue Mount Davidson ad the Taoamt Desert Wildlife
Saneruury, north-west of Alice Springs (20° 20 to 205 95°
8, O25 to 1312 53° RB), 8 7a the Davenport Ranges fram
Murray Downs Station vit Barrow Creek (20° 50" to 21°
OOS, (34 108 to ade 28! ba ST from Lake Nish
(Alpurrvlolan) (20° 20% to 21° 105S, 137° 30% Tas" 10!
(on the Barkly Tablelands. (6 from the sorthen Burkly
‘Tablelands (17° 50) ta (O° 45'S, 134 00 to £978 OO) and
10 from Marrakai fo Katherine (12) 35" to b4) 3058. bal”
Jt FAP? 20) south-east a Ditewin,

Vheestumaehs of the cats were opened so (bat undigested
contents could be identified and then the entire gastto-
intestinal tracts wer preserved in LO%e lormalin or 70%
ethanol. Phe preserved material was transported 19 Adelaide

Trpce 1. Prevedence ef itestaral helainth pomeastites: found
mm 18S feral eats from the Northern Territory,

Parisive Preyalenee (% ) Ahundance
(mean)

Acanthocephuta
Oneicoly pamaosiome — 63-4 fb 9oy | aun
Nematoda
Abbreviata tastaspiowla Aa 1 46 (121
Aneviestome canmuunt 05 3
Aim vlestomet tabaeforme J 2.8 b= teh
Cyarliospirurca das vuriiliy 44 1. 1A)
Physaloptery praepurialis 4.4 1-51 (So
Toxocura eati 0 4-15 (9)
Cestoda
Spiramnetra erinacvd lg 1-25 (m1
THenia utente fnrnis 47 U

1-747)

where pirisites were remoyed and counted usin
dissecting microscope and biter ater they had been cleéured
in fietophonal. identified employing a corpo
imwroscupe. When present rectal samples were excunined
for protozou using ventritigal (oration i sutured MgsQ,
soliton

Many of the leliiaths found (bible 1) faye been
reporled previously in surveys ol feral cats fron New South
Wales, Vietorii and “Tasmania (Anevlastame lihaelvrme.
Cyuthespione dasyirudis, Teyecere can. Sprremeira
erinaees, Taenia taerieeformis) although tere bre
sionifivant differences in previkenee between stiles. The
principal difference henween this survey and previous ones
is the very low prevalence of 7 ocefh in the Northern
Territory. Only 40% of the eats examined Nid ne parisnes.

Acanthocephilan parasites aecurrimg in feral cal
populations elsewhere have been referred lo ae Queieala
ap 2 Ph Selmide’ identified the species as Onereele
pamaayion. Both the dings. Canis fanifaris dinge, anal
fora) cataet as delinitive toss, with hurval sluges cceurring
under the skin of a variety Of pusserme birds! Oncteeder
pormitestom) Was tie most commonly detested purasrte a
this study and was ise present it hirge nunabers tn many oF
the eats. Although worms were Mound with their prabosees
deeply enbedied into the intestinal maeos. the ussociated
pabmlogival reaelivns were liniled toca inllammtory
infillrale around (he proboscis, detected wher Histilog teat
sections Were euiinged. "THis punisite wees nar reeavercd
from (9 eats inthe area south of Darwin but was presentain
animals [rool aril ares af The Northern Territory, The
large Numbers al QO. pomettastond Hound: i cats suggests
that birds constitute significant dentof their diet although
a dack of dakioon abundance of (his parunite if puratenic
hosts prevents more detuiled conclasions from being deawn

Pliyselapera pracputialis wid Mibreviaiee haistespicite
have heen reported from feral cats and dows from che
Northern Territory’ ola single occasion, The evidence bere
conlirms that 2 prepitieiy ist rekitively common parisite
of feral cats in eential Australin, Abbreviand hastaspieule ys
normally a parasite Ob weirania Faards* hut, apparently, will
develop alse in the stomach af cats Ryan founb a
“Plivsalunerd sp. present in cats in New South Wiles but
al Wal fine dismissed ef as ain ineidentil parasite Te my
have heen vA. famespeudy bur the lack of deposited
specimens mikes thimnpossible (o confirm this hypathesis,

Anevlostanid tubdeforme is generally considered te be
the common Teokwarn of domestic curs bur records to
dite sugeest (hat it po uncemmon in feral cats except hn
thease collected in the yvieiity ol Sydney?. AL dabuetorme
Was Widely distributed in the Northern Termlory, occurring
nioxt Wequently im cats fron the Kings Canyan area. the
MacDonnell Ranges und Murtay Downs Station, Infections
vonststed of small qumbers of worms (rin nubber =
3h).A single cat was found infected with AL ean. a
parasite more commonly found in canids in Northern
Australia! this cat harboured only Hive adult worms.

An immature specimen ol Cratlostamu spinige runt wats

76 M.G. O'CALLAGHAN

found in the stomach oF a cat collected in Kings Canyon, G,
spinigerum has been found sporadically in feral eats in
Australia’ ? but appears to be uncommon,

The identity of Teenia taeniaeformiy was confirmed by
counting and measuring the large and small rostellar hooks
from U3 cestode scoleces und comparing the data with those
provided by Verster!. Spiromerra erinacei was detected in
all Of the cats Collected in the area Lo the south of Darwin
but in only cizht cats (rom the other areas of the Northern
Territory. This may be due fo the faet that the first
intermediate host of this parasite is a fresh-waler vrustacean
of the genus Cyclops! and the pools of fresh water
necessary for its transmission are less Frequent in arid areas.
Parasites sueh as Dipylidinm caninun, Uneinaria
venocephala and Cyvlicespirara felineus. present in other
surveys, Were nol found,

Facea! cauminutions revealed two species of coceidia,
baspora felix in O60 of 146 cats and 7 rived in 6.90%.
Both are common parasites of cats and ean cause disease in
younger animals, The majorily of the cats examined were
adults and this could explain the Jow prevalenee of
coccidiin infections (including the absence of Tavepsleene
gondii) io this and other surveys’,

Additional nematodes. Lehimonenia canenin, Rictularia
carstaient and Wararisirongylas clenott were rarely found
im their presence was presumubly the result af the
ingestion of native manimals und reptiles which are the
normal hosts of these parasites. Similarly. the imestion of
birds would explain the presence of female nematodes o
Aprocta Sp. in the stomachs of (Vo cats, Other parasites

ones, BR. (1983) “Feral Cat? po 489 Jn Strahan, R. (Ed)
Complete Book of Australian Mammals” (Angus &
Robertson, Australia).

‘Ryan, G. E. (1976) Aust Vet, J. $2, 224-227,

‘Coman, B. J. (1972) fhid. 48, 133-136.

'Gvegory, G.G. & Munday, B. L. (1976) Ibid. 82, 317-320.
“Coman, B. J., Jones, E. H. & Driesen, M.A. (1981) bad,
57, 324-327.

‘Schmidt, G. D. (1983) J. Parasitol. 69(2), 397-399,
‘Barton, M. A. & McEwan, D. R. (1993) Aust, Vel. J,
7(7). 270.

collected from intestinal contents and faeces but obviously
related to the ingestion of rodent hosts were Denieder sp,
und Pyorergarés sp. Xenopsvila vexabtis, aw flea found
commonly on Ratty villosivyvimuy was found in the
stomach of cats from the Lake Nash area and the mite.
Laelups hapalati, a parasite of Notaries spp. wits found in
cals from the Tanami Desert area. Another accidental
parasite recovered was Sypacia obvelatia an oxyuricd
Parasitic me the cuecum) of rodents.

The information presented here identifies a greater
Variety of parasites occurring in feral cats in the Northern
Territory than previous studies have found, It identities ?
praepatialis a a Common parasite and demonstrates that
ills are Frequently infected with A. hesrespicnds. Te alsa
confirnis Hat the feral cat preys on mative mimmials, birds
and reptiles, This is particularly so in the ease of birds. with
the large muimbers of GQ. ponafosioni suggesting frequent
predation.

This work was begun by Murray Barton, when cniployed
atthe Arid Zone Research Institute, Alice Springs und was
conducted on material collected by the stall of the
Conservation Commission of the Northern Territory, Alice
Springs, We wish to thank Mr Barton and all of the
individual officers who kindly collected and preserved the
viscent of the cats and David Gibson for his help with the
HINSC rip,

Representative specimens of all of the helminths
collected have been depesited with the Australian
Heminthalogical Collection, South Australian Museum,
Registration numbers AIC 40181 to 30231

‘Jones, H. 1. (1983) Aust. J. Zool. 31, 285-248,

“Prescott, C. Wy. (1984) University Of Sydney,
Postgraduate Foundation in Veterinary Science, Review Nao,
24. Parasitic Diseases of the Cacin Aust.

"Verster, A, (1969) Onderstepoort J. Vet. Res. 3601), 3-58.
'Dunsmore, J. D. & Shaw, S. E. (1990) University of
Sydney, Postgraduate Foundation in Veterinary Science,
Review No. 31. Clinical Parasitology of Dogs.
“Beveridge, 1., Presidente, P. J. A. & Arundel, J. H,
(1978) Aust. Vet. J. 54, 46.

M. G. O'CALLAGHAN, Primury Industries (South Australia), GPO Box 167) Adelaide S, Aust. S001 and [.
BEVERIDGE, The University of Melbourne, Veterinary Clinical Centre Princes Highway Werribee Vic. 3030,

PREDATION OF EARTHWORMS BY THE LAND PLANARIAN,
AUSTRALOPLANA SANGUINEA (MOSELEY) VAR. ALBA
(DENDY) SENSU JONES, 1981 (TRICLADIDA: GEOPLANIDAE)

BRIEF COMMUNICATION

Summary

Earthworms can significantly reduce soil degradation and improve pasture and crop
production’. Land planarians (terrestrial flatworms) can, however, greatly reduce the
abundance of earthworms in agricultural fields. For example, a New Zealand
planarian, Artioposthia triangulata (Dendy), has been accidentally introduced into the
British Isles where it now feeds voraciously on local earthworm populations. The
implications for crop and pasture yields and the abundance of wildlife that feed on
earthworms in the British Isles, are of major concern’**. Another planarian,
Australoplana sanguinea (Moseley) var. alba (Dendy) sensu Jones, 1981, has also
been accidentally introduced to the British Isles, either from Australia or New
Zealand**°. Relatively little is known of its biology, in particular its feeding habits,
except that it will feed on “earthworms”.

Cramartions at tie Reval Seerary ofS. Awse 119964, EE,

BRIEF COMMUNICATION

77-178

PREDATION OF EARTHWORMS BY THLE LAND PLANARIAN, AUSTRALOPLANA
SANGUINEA (MOSELEY) VAR, ALBA (DENDY) SENSU JONES, 1981
(TRICLADIDA; GEOPLANIDAE)

Lathwoniits can significintly reduce sort daeradanon and
improve pasture qin crap peeduction! Land: plunarians
(ermestril Thtwornish cin, however, greally rediee (he
“Dondanee oF carthiworms in agricultiinal fells. For
example. a New Zealand pling. Ariepe iin
rrimenkie (enly bas been aecidentally muroduced into
(he British hlesowhere Th iow feeds Volaciousty Gn lien)
carthiwernt) populates: ‘The joplicaiions fer erep: anc
pasture vields sind (he abundance of waldtife hat feed on
eartinwarms om the British Isles, afG of major coment
Another phinurian. Adstralaplang sencuned (Moseley) var
adn) (Dendy) sense domes, 1984, bits also been aecidentally
Introdeed lo the Brtaly tshes, eiiier fon) Austtulia or New
Zoalind Relatively lwle is. known oP ats biology. on
pucticudal ity feeding: habits, excepe that i} will feed on
“gurthavorms

Austulaplaid species are widespread and. trequently
eneougtered fn taineigeditied habitat, on easter
Australia’’. Recently, A, sunwuined var athe has heen
found to be neither’, sai@uined nor A. cle but isin tae a
Wew genus tad spowes (bh. Winsor pers cami), Pats
jiawware Ss onty Kuews oecurrence i Australia is at
Kineswood, South Australia. although it probably oeeuny mn
diher trban sad disturbed habitats.

The dorsal surfiee of A. seria var, cll te mostly a
untarn pade Ohanige-hrowi colour with a deeper brawn
peach anterior psits ventral surface is white, A, deeneaiedt
vir, dba aves Commonly found by one af us (MT) in
moist habitats, such as under bricks. rocks and pieces al
wood und. sometimes. id the top LO em of seit ina subi ban
pardeo in Kingswood, Ufed on garden earthworms when
offered them fy plustic jars, but Hot OW Other imvertebriles
such as Snails. sligs. millipedes, shaters Or eurwigs.

This paper documents five earthwornt species upon
which A, Sanguine var afb wall feed tinder laboratory
Conditions and provides Some dats on dts reproduction and
early development The wark forms part of a broader study
aimed al improved ymunaeement al earthworis as a
resource in AduTfecastern Australia,

Purstly, ais adult Wy savauined va, alba lapprax. 6-8 ont
Jonw) Were collected from the Kingswood garden in
October 1904. These plavurians were plteed ia pans tn
(bree opaque, Closed plastic coplainers (wan 9 era; Acialit
(Oocm) and maintained ut t°C. The bolum af eich
contitiner was covered with 1.5 em of givden sol which
was Kepl anoist hy sprying cewulaly with water, Pive
exohie curthworm species, Aperrectudeu calfeinose
(Savianyh ay arsed (Savigny), A, mreapecetdes (Duges) und
A. lane (Ue) (Lurmbricidges dnd Microscalen dibius
(Pletcher) LAcaothoilrifidae) were offered to the planuriuns
us food, These garlhworm spevies, apiel fhony A. hinge
which is Widespread mm and was ohmined from northern
‘Tasmania, are commonly found in gardens and aerica}iural
Nields iy South Australia and Victoria "! Two ineividiials of
ub particular cavihwarm species. Were miroduved vate each

COntioter LEN thie Chabwornps. ie CArrhworn Sper ies
at a time, Procdition was assessed over a period of Ff ob
Banhworrn species Were changed in the containers when
predation aueuned, All species of earthworm were killed
and eaten by A. saagined yar alba, The phanarians gto alt
Or prot oh aa carthwort), AW parts of the body Were
Auiehedl wantely head, kil and midsection. [iy some eae.
where oTly the [ail was euler. (he earworm sorvived. tn
contrast, carmbworns diwovs die when partilly eaion hy A,
Hreingldane.

Seeondly, seven adil a, sage var adhe, Tron Ue
sume Kingswood carded, were placed 1) separate plastic
combines. us desevibed tbove. Two adult Apateeernidia
vifemose were added to vayh container, The nurabers of
missing oF purtilly cater carthworns were recwded
approximately every 2) oh lor 3a 0) Burthworms were
replived [complete or pactial predation bud oeeurred, Two
planiriaos dived during the osperimefy, horh a} iay 28. On
average, GarthwWorms Were altacked aiee evry 5S dpe
phivanan tranpe | - 12 dy. This ts stnilar ia the tate at
which A. miata iliwked another carlhwor species,
Piyenia fetid Savieny) (14 alicks per plandriin por week
weer.

Muring the first experiment. the sty phunundns deposited
(ive cee capsules between them. In the second experiment,
the sever plananans deposited [7 eve capsules (4-0, 1.
2 3 and 3 wichh. The capsules avere spherical to oval oa
shape ald purged tom 3 - 4 mm in diameter They were
miurgerered i colour und were deposited either oy the soil
surface or afew mor below it bul were not athiched to the
sou) surtuee hy slime ws Has been roted hv Terrace &
Baker’ for Crenaplima coeclea (Moseley). ‘The vey
capsules were tainnioed at 13°C unopaque, closed. pkistic
comtuiners (jam, Fem hewht Seni) foe up. te 144d.
Foartven of the capsules hatched, These suecessfil capsules
were deposited berween 3) Oviober and 13 December 1994
and all hatched during February (995. Mean hatehiig finite
wus 87d trange 71 - 101 d). Forty one jayenile plinurians
were rented from The capsules. | - 5 from each capsule
(mean = 4). Upon emergence, these juveniles were pale
yellow in colour with a reddish untecior hp. The main bady
volour darkened to either ¢ grey or pale orange = brown over
a3. 11d period. The juvenile planurians were provided
wilh juvenile A, culigiresa as food (ad fib) and began
feeding on them after dn venue of 10d (range 3-22 cd).
Right of (he javenile planarians reached wut sive three
months after emergence, These eight indiyaiduals were then
prouped ia One conkuoner, and, afer a further month,
produced three egg capsales heiween then

Introduced lumbricid: earthworms are generally mich
Hore ablindany thin native speeics in Urban dnd agricultural
sells in southern “Australi!!! Native earthworms sre
presumed (oO be nore abundant iy undisturbed habitats,
where invasion by intradgeed species is believed to he
lioredh) Whether or not . aineuirea var afb) preys

178

upon native earthworm species or significantly influences
the abundance of cither introduced or native species in
disturbed or undisturbed sites in the field has yet to be
determined. However, given the numbers of earthworms
eaten in the laboratory experiments ( | worm / planarian / 5

‘Lee, K. BE, (1985) “Earthworms. Their Ecology and
Relationships with Soils and Land Use” (Academic Press,
Sydney).

*Blackshaw, R. P. & Stewart, V. 1. (1992) Agric. Zool.
Rey. 5, 201-219.

‘Blackshaw, R. P. (1995) Acta Zool. Fennica 196, 107-110.
‘Boag, B., Yeates, G. W., Johns, P. M., Neilson, R.,
Palmer, L. F. & Legg, R. K. (1995) /bid. 196, 212-214.
‘Jones, H. D. (1981) J. Nat. Hist. 15, 837-843.
’Anderson, R. (1986) Ir, Nat. J, 22, 141-146.

’Winsor, L. (1977) Proc. R. Soc, Vic. 89, 137-146.
‘Winsor, L. (1979) Vic. Nat. 96, 155-161.

"Baker, G, H. (1996) The ecology, management and benefits of

d), the influence of A. sanguinea on earthworm abundance
could be significant.

We thank Don Terrace for help with the collection of the
planarians and Leigh Winsor for assistance with
identifications.

earthworms in agricultural soils, with particular reference to
southern Australia /1 Edwards, C.A. (Ed.). “Earthworm Ecology”
(Soil and Water Conservation Society, Ankeny). In Press.
Baker, G. H., Thumlert, T, A., Meisel, L. S., Carter, P. J.
& Kilpin, G. P. (1996) Soil Biol. Biochem. In Press,
''Terrace, 'T. E. & Baker, G. H. (1994) J. Aust. Ent. Soc.
33, 371-372.

"Wood, T, G, (1974) J, Anim, Ecol. 43, 87-106.

Abbott, F. (1985) Aust. J. Soil Res. 23, 263-270.
"Lawson, L. M. (1993) “The distribution and abundance
of native and introduced earthworms in an area of pasture
and native vegetation near Cape Jervis, South Australia”
PhD Thesis, Flinders University of South Australia, Unpub.

T. E. TERRACE, Division of Soils and G. H. BAKER, Division of Entomology, CSIRO PMB 2 Glen Osmond

5. Aust. 5064.

MASS FROG MORTALITY AT TWO LOCALITIES
IN SOUTH AUSTRALIA

BRIEF COMMUNICATION

Summary

In recent years much attention has been given to the alarming decline in amphibian
populations worldwide. Australia is no exception to this with severe reductions in
population size recorded for several species'*”. Suggested reasons for apparent
declines vary widely, with habitat alteration and fragmentation, acid rain, introduction
of exotic animals, toxicants, increased UV and pathogens being implicated in several
instances**®. It is possible that such factors also act in concert”. For example, habitat
alteration may sufficiently stress the frogs so that their immune systems become
compromised, making them more susceptible to infection’. It has also been suggested
that such declines may only be a small part of a naturally occurring cycle of
population flux’.

Fromnonias of te Row dando ah ay Mie (199, 12004 179,

BRIEF COMMUNICATION

MASS FROG MORTALITY AT TWO LOCALITIES IN SOUTH AUSTRALIA

I) decen years WUch avention bas been piven to the
alarmmg declitie TH amphibiin papulitions werkwrle,
Austialis ty no exeeprion ti this wilh severe redichons in
population Size recended (or sever species! 7. Sueeested
reasdns for iipparent declines vary widely. with habit
allerunin aod) fragmentation. uci rain ytroduetion of
exote aunts, losieants, laereased: (Vo and) pathogeris
hele implicated in severabinstinees! "Tis possible tat
stivh factors alsa yeh in convene. For example. babar
alteration may sufficiently stress (he lroas se: (ht (hele
TWintuhe systems bevonie compramised, making them more
susceptible (a infeetion®. Th Has alo heen sugested: pit
suich declines may only be a sryall pat of a qmatraeily
Hecurine evel of PopHhAtlone TUK

Ty the field the Observation @P severil tla Pros
simutiineousty ts inusial as they degrade quickly alter
Jeu. Henee the presence of MUI ple carcasses Tso Si of
mass Wordilily Wea Short dine frame for frags this world
probably te less Than tire days). There have heen recent
reporls ob such a phegomenun frome several eegtors
Tnehiding Switverlund. Hawaii England! and Australia,
Here wi report Hass Mog minutes at two loculires in
South: Australia

Vhe first tnendent took pluce on 20 Muy 1992. Rain ut
Nuriootpa. South Australia (larrude 34° 29° 5. fongitude
(AOU EP) Filled yo dam ong vinevard loa depih of
approximately OS it Numerous burrowing frogs
(Neola lio picts) emerged sink vn the following dey,
nrimy were Lound dead at the edges of (he dard One ob as
(Mi TS) vasited the site on 4 May L992 cind recovered 32
dead specimens. AL were at the edge OF ple water whieh
had sready receded (a depth of O,3 em, Some fad Tae
Spawi but other dead, gravid Wndividials were lneated, The
Spawn Tid dried vulaind if could net be deterinine Lt te
ova hud been ferrlised,

Nhe scomilineoiis mtite OF the death of the fags
indicited the presenee ef a pollutant. According to the
laudewners the only chemical used ap rhe site was spol
opriving of the herbiode Rotiidoep Cienulietired by
Monaiinta andl eonsiating at etypltosite and a dispersaney to
conol tiistles, Spraying hud heen undertaken
approx nomely Uiree (onthe previously Gin Pebruary Tua)
This pemod should hive been adequate for the deerdauen
ail the herbicide,

Tyler Mya, (1994) Alyten #450.

“Richards, S.., WeDonald, KK. R. & Alford, RL AY |) 905)
Poortte Conservation Biology B.GO-77,

Vial, be L, & Saylon be (9S) The situs of amphibyia
pepultionss a compilation antl analysis Workii
document LIUICM/SSC Dectiting Arplibiin Tsk Fores.
Worl) Comservation) Craton.

"Wyman, Reb. (1990), Conservattiin Biology 4. 450.357
Cirey, GC. & Bryant, ©. 4, (1995) Environmental Healthy
Perspectives WS Supyph 4. 14-17

‘Laurance, Wo 71, MeDonald. KOR. & Speare, K,
Conservation Biglogy, ty Press.

‘Carvy, ©. (1993) [bid 7 395-362.

The second sie of mass moriulity was ait Parahind Pt
Springs, 27 kin northeast of Arkardohe Homestead ye tie
northem Flinders Ranges (OM 199606 on Yudnirmutini
6737-1. 150000 Lopogmaplie hap) This is a georherinally
heated body of water through whieh radon gis bubbles,
Walter emerges ar GO® Coanth bats a padiowetivily, of
approximmeely JOO pCu/l After sulfivient taiilall Tesh
willer cin alse he found upstream al the hot spring. On ¥
December 1994 nine deal frogs were discovered by C.W,
Tovttine atthe hot source al the spring anh by. herr state.
were considered to hitve heen dead for fess than two diy.
Seven tineliuding one juveniles avere the spotted wrtss (nog,
Linnodynasies dagnemensy. anid We ole Iwo were
representatives of the red tree frog, Litetia enbelli. Their
adVineed state ou deeay preeluded) 3 meanimelul
myestigulten by pathologists,

Despite extensive searches, me line Lined estes
laymanionsss were discovered al the site. whereas Gitepla
ribella was beard calling but not seen, In cooler parts of the
sapling deawstreay. Were were both larval aid acti
frowets (Crimea riperiol and ne dead frogs were loud
These phservations were made the day after rhe fest
oceusin What fain had fallen tor one maath,

Mass mortilines were again recorded on & May l96 a
the same she, Rleven dem, eparie were found both sl the
hot setiree Gnd opstrGa re ob TL A isalafed: pools of coaler
water (24) CO. Vhese ineloded both adetty and juveniles.
Several living durval, juvenile and adult Co riperre and L,
fasnmuiensts were also observed af the sie on this dite.
Qucasionil sightings of deuwd frogs floating in the hot pauls
have been made repulwdy over the hist 20 yaurs CD, Sprige
pers somo)

The presence of such a large noniber of dead frogs is
Sligpestive al a palllagenico cuuse. Phis occurrence canna
be altribuced readily tooany dorm of anthropogenic
ince ferenee. ind While high water femperunire may have
vontibuted tothe mass corhility. pes dilikely to have been
the single cause of so many deaths within such ashore fine.
AlLhouth Tavestization lite amphibian pathogens ts siihin
its anfaney they have heen previously linked with) trop
inertilities both in Australi’ and abrevstel!

Simon Stemborner, Advi Bradford. Steven Walker und
Rebeecou Short are thanked fur then assistance mnie Oelel.

'Pechmann, J. U. K., Seott, Do E.. Semlitseh. KR. D.,
Caldwell, . P. Vili Ly. & Whitfield Gibbons, J. (1991)
Science 253. 892-995,

‘Wroglog (7492) Newsletter of the TECN/SSC Tusk Foree
oy Deeliing Amphibians, June 1992 (2)
"Green, DAB, (1904) Report produced for the Disease ind
Pathology Warking Group. Declining Aniphibian
Populations Tasidforce.

Drury, S. E. N, Gough, KR. iE. & Cunningham, A. A.
(1995) Veterinary Roconl (47, 72-73,

‘Cullen, B, R.. Owens, L, & Whittington, Rf. (1995)
Diseases of Aquithe Organisms 24, §3-42,

MICHAEL LOPYLER and CRAIG RO WILLIAMS, Department of Zoology, University of Adelude 8. Aust

S05,

ROYAL SOCIETY OF SOUTH AUSTRALIA INCORPORATED

Patron:

HIS EXCELLENCY THE GOVERNOR OF SOUTH AUSTRALIA
SIR ERIC NEAL, AC, CVO

OFFICERS FOR 1996-97

President:
T. C. R. WHITE, BSc, BSc(For), PhD

Vice-Presidents:
M. DAVIES, BSc, MSc, PhD
M. A. J. WILLIAMS, BA(Hons), MA, PhD, DSc

Secretary: Treasurer:
O. W. WIEBKIN, BSc, PhD J. H. BRADBURY, BSc, MSc
Editor: Assistant Editor:
J. BIRD, BSc N. F ALLEY, BA(Hons), MA, PhD
Librarian: Programme Secretary:
J. A. ROURKE, BA(Hons) A. F. BIRD, BSc, MSc, PhD, DSc

GradDip (Inf Studies)

Minute Secretary: Membership Secretary:
G. A. CROOK, BSc
Members of Council:
P. KOLESIK, BSc, PhD S. BARKER, BSc (Hons), PhD
N. A. LOCKET, MA, BM, BCh, PhD, DO M. J. WRIGHT, RDA
R. D. SHARRAD, BSc(Hons), PhD, DipT(Sec)

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