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VOL. 119, PARTS 1 & 2
31 MAY, 1995

Contents

Transactions of the

Royal Society of South
Australia

Incorporated

Bourman, R. P. A review of laterite studies in southern South Australia - -
Shiel, R. J. & Dickson, J. A. Cladocera recorded from Australia - - - -
Kolesik, P. Skusemyia allocasuarinae, a new genus and species of Cecidomyiidae
(Diptera) damaging lateral branch buds of drooping sheoak,
Allocasuarina verticillata in Australia - - - - -
Nicholas, W. L. & Stewart, A. C. New genera, species and a new subfamily of Xyalidae
(Nematoda: Monhysterida) from ocean beaches in Australia and
Pe a a ee a ee es
Bradbury, J. H. & Williams, W. D. A new genus and species of crangonyctoid SERIES
(Crustacea) from Western Australian fresh waters - -
Fuller, M. K. & Jenkins, R. J. F. Arrowipora fromensis, a new genus and species of
tabulate-like coral from the Early Cambrian Moorowie Formation,
Flinders Ranges, South Australia - - - - - -
Molina Ballesteros, E., Campbell, E. M., Bourne, J. A. & Twidale, C. R. Character
and interpretation of the regolith exposed at Point Drummond, west
coast of Eyre Peninsula, South Australia - - - -
Smales, L. R. A revision of the genus Tikusnema (Nematoda: Acuarioidea) with the
description of a new species from the false water-rat Xeromys myoides
from Queensland - - - - - - - - -

Brief Communications:
Smales, L. R. Mastophorus muris (Nematoda: Spirocercidae) from the Musky Rat-
kangaroo, Hypsiprymnodon moschatus - = = = 3

Vilizzi, L. & Walker, K. F. Otoliths as potential indicators of age in common ae Gee
carpio L. (Cyprinidae: Teleostei) - -

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ih)

TRANSACTIONS OF THE

ROYAL SOCIETY
OF SOUTH AUSTRALIA

INCORPORATED

VOL. 119, PART 1

TRANSACTIONS OF THE
ROYAL SOCIETY OF SOUTH AUSTRALIA INC.

CONTENTS, VOL. 119, 1995

PARTS | & 2, 31 MAY, 1995

Bourman, R. P. A review of laterite studies in southern South Australia - - -
Shiel, R. J. & Dickson, J. A. Cladocera recorded from Australia- - - - -
Kolesik, P. Skusemyia allocasuarinae, a new genus and species of Cecidomyiidae (Diptera)
damaging lateral branch buds of drooping sheoak, Allocasuarina verticillata
in Australia - - - - - - - - - - -
Nicholas, W. L. & Stewart, A. C. New genera, species and a new subfamily of Xyalidae
(Nematoda: Monhysterida) from ocean beaches in Australia and Thailand
Bradbury, J. H. & Williams, M. D. A new genus and species of Eraneonscwid amphiged
(Crustacea) from Western Australian fresh waters = - -
Fuller, M. K. & Jenkins, R. J. F. Arrowipora fromensis, a new genus and species of tabulate-
like coral from the Early Cambrian Moorowie Formation, Flinders Ranges,
South Australia = - - . - - - - - - -
Molina Ballesteros, E., Campbell, E. M., Bourne, J. A. & Twidale, C. R. Character and
interpretation of the regolith exposed at Point Drummond, west coast of
Eyre Peninsula, South Australia - - - - = = -
Smales, L. R. A revision of the genus 7ikusnema (Nematoda: Acuarioidea) with the description
of a new species from the false water-rat Xeromys myoides from Queensland

Brief Communications:

Smales, L. R. Mastophorus muris (Nematoda: Spirocercidae) from the aise Rat- kangaroo,
Hypsiprymnodon moschatus - - - - - -

Vilizzi, L. & Walker, K. F. Otoliths as potential indicators of age in common carp, Cyprinus
carpio L. (Cyprinidae: Teleostei) — - - - - z > »

PARTS 3 & 4, 30 NOVEMBER, 1995

Li, Q. & McGowran, B. Comments on some southern Australian foraminifera and description
of the new genus Parredicta - - - - - - - -
Cappo, M. The population biology of the temperate reef fish annie iin niga in an
artificial reef environment - - - -
Schluter, C., Bye, J. A. T. & Harbison, P. The most vigorous South Australian tide = -
Bird, A. F. Studies on Eutobrilus heptapapillatus (Nematoda: Tobrilidae) the predominant
nematode inhabiting the bottoms of Lakes Albert and Alexandrina, South
Australia - - - - - - - - - - -
Skerratt, L. F., Beveridge, I. & Durette-Desset, M. -C. Distribution of species of
trichostrongyloid nematode parasites in the small intestine of the bush rat,
Rattus fuscipes - - - - - - - - - -
Barker, S. Eight new species of Australian Buprestidae (Insecta: Coleoptera) - - -
Richards, S. J., Burton, T. C., Cunningham, M. J. & Dennis, A. J. A new species of
Callulops from New Guinea and comments on the status of C. humicola
comptus (Zweifel) (Anura: Microhylidae: Asterophryinae) - -
Stewart, A. C. & Nicholas, W. L. Manunema pectenophora sp. nov. (Peresianidae,
Leptolaimina), a nematode possessing unusual male supplementary
organs - - - - - - - - - - - -
Kolesik, P. Asphondylia dodonaeae, a new species of Cecidomyiidae (Diptera) damaging leaves
and branches of hop-bush, Dodonaea viscosa (Sapindaceae) in Australia
Kolesik, P. Contarinia bursariae, a new species of Cecidomyiidae (Diptera) infesting fruits
of sweet bursaria, Bursaria spinosa (Pittosporaceae) in Australia -
Williams, D, J. & Brookes, H, M. A review of the scale insect subtribe Andaspidina
(Hemiptera: Coccoidea: Diaspididae) and a new genus, Notandaspis, for
two Australian species - - - - - - - - -
Connolly, R. M. Diet of juvenile King George whiting Sillaginodes punctata (Pisces:
Sillaginidae) in the Barker Inlet - Port River estuary, South Australia -

Insert ta Transactions of the Royal Society of South Australia, Vol. 119, parts 3 & 4, 30 November, 1995

113
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149

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163

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19]

A REVIEW OF LATERITE STUDIES IN
SOUTHERN SOUTH AUSTRALIA

By ROBERT P. BOURMAN*

Summary

Bourman, R. P. (1995) A review of laterite studies in southern South Australia. Trans.
R. Soc. S. Aust. 119(1), 1-28, 31 May, 1995.

Studies of laterite in southern South Australia are reviewed to throw light on the
nature of laterite, its genesis, classification, its relationships to substrate materials and
constraining sediments, its use as a morpho-stratigraphic marker and palaeoclimatic
indicator, its relationships to deep weathering, and the timing of lateritisation.
Evolving views of laterite as a rock unit, as an iron-rich horizon and as a weathering
product are traced and processes attributed to laterite formation viz., capillarity,
leaching, combinations of water table movements, leaching and capillarity, wetting
and drying processes, weathering transformations of materials rich in ferrous iron, and
as a lacustrine deposit are assessed. Fundamental to theories of laterite genesis are the
roles of relative and absolute accumulation of iron and aluminium minerals.

Key Words: laterite, laterite profiles, ferricrete, polygenetic profiles, peneplains, deep
weathering, palaeoclimatic indicators, multicyclic landscapes, morpho-strati graphic
markers,

Transacrions of tt: Royal Society of S, Aust. 1998), WAU), 28.

A REVIEW OF LATERITE STUDIES IN SOUTHERN SOUTH AUSTRALIA

by ROBERT P. BOURMAN*

Summary

Bourman, R. F995) A review of laterite studies in southera South Australia. Tras. R. See S. Aust U9).
1-28, 31 May, 1995.

Studies of kverite i southern South Australia ure reviewed! to (hrow light on (he nature of Jaterte, ils genesis,
classification, its relationships to substrate materials ind constraming sediments, its ise as a morpha-stradigraphie
marker and palacoclinnaitic indicator, its relationships to deep weathering, and the (ming of lateritisanon. Evolving
views of laterite as at rock amit. as.an iron-rich horizon and as-a weathering product are duced iind processes
attributed i laterne formation viz,, capillarity, leaching. combinatons of waler lable movenients, leaching and
capillurity, wetting and. drying processes, weathering transformations of materials rich in ferrous iron, and as
a lacustrine deposit.are assessed, Fundamenta, to theories of laterite genesis are the roles of relative and absolute
accumulation of iran and aluniiniuin minerals,

In Southern South Australia interpretations of lindscape evolution have depended heavily on recognition of
parts of an original normal laterite profile, consisting of a pallid, bleached zone successively overlain by a mottled
yong and latente, a ferruginous and/or alumioous crust, ‘Chis profile has been associaled with formation on a
penepluin surface under a humid, bul seasonally dey, (ropical climate. The possible preservation oC a pristine
lulerite surface of ereat anhquity in the modern landseape On uplifted peneplains has been entertained by some
workers, but questioned by others. Alternatives ta this approach are provided by stratigraphic inyestigalions of
polypenetic profiles, and a continual weathering model of laterite formation (hat results in lateral Variability in
the distribution of pallid, mouted and daterite materials ord surface initially with regular Wpogruphy. Laterpretations
of lateritiged landscapes include diflerential dissection of a complete laterite profile on an uplifted peneplain surface,
mullicyelic landscapes successively lateritised and the formation and Jateritisalion of high level surtaves during uplift.

Evidence of laterite formation under non-teopical conditions questions the climate-laterite correlation as does
the lack of reliable minerals as climatic indicators of laterifsation. Furthermore, the recognition of laleritisation
oveurring throughour the Mesozoic and Cainozoic restricts lhe usefulness of Iilerite asa palaeoctimiatic and morpho-
straligraphic marker,

Key Worps: laterite, laterite profiles. ferricrete, polygenetic profiles, peneplains, deep weathering,

pakieochimane mdicators, mulsicyelic landscapes, morpho-straigraphic markers

Introduction

There iy a long history of research on materials called

laterite in South Australia. The general distribution of

lulwrilic materials in southern South Australia is shown
in Fig.t. Specifically, these materials include
ferruginous and aluminous crusts. yaniably described
as ‘ortsteim, “Ferruginous duricrust (Lang 1965),
‘durierust’ (Woolnough 1927), ‘ironstone’ (Teale 1918),
‘ironstone cappings’ (Segnit 1937), “indurated zones)
‘ironstone gravels” (Prescot! 1931) and ‘fetricrete’
(Firman 9674; Bourman 1969'; Milnes et ul, 1985),

© School of Human und Environmental Sciences, University
of South Australia, Holbrooks Road, Onderdale, South
Australia, 5032.
BotkMan, RP (1969) Landform Studies near Victor
Harbour, B.A. (Huns) thesis, The University of Adelaide
(unpubl).

weathered bedrock. sediments and soils, variably
lerruginised, mottled and/or bleached. This paper
summarises and critically comments on these
definitions, Issues addressed include the diversity of
interpretations concerning the nature of laterite,
processes of laterite development, laterite profiles, (he
topographic and climatic requirements for its
formation, its age, and reconstructions of: and
interpretations drawn from, laleritie landscapes.

The tert ‘laterite’ has been in the scientific iterature
since the publivation of Buchanan (1807), David (1887)
discussed the origin of laterite in the New England
district of New South Wales. but did so without
reference to Buchanan's work, and the term did not
appear in.the South Australian literature until more than
100 years after its First usage (Teale 1918). Nevertheless.
features. subsequently regarded as laterne were
discussed by early workers under labels such as ‘Desert
Sandstone’ (Woolnough 1927) and “Upland Mincenes’
(Tate 1879),

With few exceptions, the majority of investigators
in South Australia have followed the view that laterite
formed as a result of intense chemical weathering in
a seasonally dry tropical climate on a peneplain
surface, largely during the Tertiary. These conditions
fayoured the development of a laterite profile
comprising a leached sandy A-horizon successively
underlain by laterite, a mottled zone and a pallid zone
resting on unweathered bedrock (Fig.2). Generally, the
present discontinuous distribution of these materials
has been ascribed to differential erosion following
tectonic uplift of the peneplain.

Corrobinnie
Depression

Cleve
Uplands

Neild ~
Glenville Hs

4
ad ee
wit

ROBERT P. BOURMAN

Definition of laterite

Laterite as a rock unit

Early studies of laterite in southern South Australia
were undertaken by geologists, who considered laterite
to be a rock or sedimentary unit and equated it with
‘Desert Sandstone’ (silcrete) or with terrestrial deposits
referred to as ‘Upland Miocenes. For example, Tate
(1879. p, lix) regarded ‘evenly-bedded sandrock,
mottled clayey sands and ironstone conglomerates,
occupying flat-topped localities in the Adelaide foothills
and within the ranges as ‘Upland Miocenes. He

Flinders

Mt :
Olinthus f
? /

ara \ ve “Blue Range

i -n rote 4

4) ” Turnby Bay.

NN
Flat

~ Pt Lincoln

Cape

AUSTRALIA

1000

Yallunda

Taylor

Wadella
Springs

Oy
XO

Cygnet - Snelling
Fault Scarp mee

Border, ae Parndana

Fleurieu
Peninsula

ipa fe

Banks

Kangaroo
Island

40 60 80 100 Km

Fig. |. Map showing the localities of lateriiic areas in southern South Australia referred to in the text.

Mig, 1A. (Opposite page). Inset in Figure 1.

*/ Ardrossan

(

By
me
ea
ee)
s
Ew
n
a
aa)
GO

LATERITE IN SOUTHERN SOUTH AUSTRALIA

Adelaide © ’

Hallett Cove

Witton Bluff

s
Myponga

Spring &
ount

Happy ~,
s Valley _.:

nf Kapund

ue = aM Gawler
“ & '® Highbury

‘
} & , Gun
| Emplacement

Mt Lofty

) Blackwood Mt

Barker

- Longwood a
Clarendon a
Meadows °,
e .
Macclestield

Mt Compass
Upper

Cz «~ Hindmarsh

Valley

A < Peeralilla
Hill

Green Hills
e

a Parawa®
e
ca

Wilson

Hill «A

Victor
Harbor

Bremer Basin \

Millendeljg Fault

“s. Harrogate

° 1
a’ Whalley Hill

on!
“> Lucernbrae

'
‘

*s~ Callington

a)
‘

‘

4 ROBERT P BOURMAN

loneinietsatey A lhyfiaian

Lager tle — otewed oy Vu tealir
varpurtrea Lap, peli,

Mottled cw

Titi tnt
Melnice Wealpes

PUN Bony

Fig, 2, Sketch of the pedogenic model of the normal ur
stayidard laterite profile incorporating a sandy. bleached A
horizon above a laterite horizon, successively underliin by
companion materials of mottled and bleached bedrock
(Stephens 1946). considered to fave developed on a
peneplain under humid tropical eanditions,

considered hen to be correlative bout with terrestrial
clays overlying fossiliferous limestone at Adelaide and
Tertiary terrestrial sediments borderimy the ML Lolly
Ranges. Later work has demonstrated that these
sediments vary in age from Pleistocene ta Eocene.
Furthermore, the limestone exposed in Adelaide 1s now
known lo be Late Pliocene (LLudbrook 1980) rather than
Miocene as assumed by Tate.

The Desert Sandstone in northern South Australia,
currently known as silcrete, was interpreted by Tate
(1879) ds an extensive lacustrine deposit contemporary
with fiver gravels and sands of the Upland Miocenes.
Thus silcrete and laterite were noi distinguished and
they were both considered to be sediments or rocks.
Whereas Tate (1879) equated sediments within the
fanges and on their flanks as Upland Miocenes, Benson
(1906) separated them into iwo groups, with an older
Miocene series cupping hills and a younger series
flanking the Western escarpment of the Mt Lofty
Ranges.

More recent papers have also consideréd laterite to
be a rock unit, For example, Major & Vitols (1973,
p. 46) described laterite on Kangaroo Island as a
‘massive rock composed of pebble-sized pisolites of
maghemite and limonite and fine-grained quartz sand
cemented by limonite. This crust, up to 1 m thick, was
overlain by white, fine-grained quartz sand and
underlain by mottled yellow and red clay or rocks of
the weathered Kanmantoo Group metasediments. The
crust occurred as boulders where ripped up by ploughs
and loose pisolites, mixed with white or yellow sand,
were recorded on the margins of the inland plateau.

2 Bourne, £ A. (1974) Chronology of denudation of
Northern Eyre Peninsula. M.A. thesis, The University of
Adelaide (anpubl,).

Lanerire as an jrat-rich harizoan

Many geological studies have been concerned with
laterite in only a very incidental fashion, and almost
any iron-rich horizon has been regarded as laterite (e.4,
Glaessner 1953a, Olliver 1964). At various locations
within and on the margins of the Mt Lolly Ranges,
Teruary sediments, variably weathered and
ferruginised, have been reported to centain laterite. For
example, at Happy Valley, Olliver (1964) described a
sequence of Eocene marine Blanche Point Marls and
North Maslin Sands overlain by Pliocene freshwater
sands and clays capped and preserved by a lateritre
horizon at about 200 m above sea level. The laterite
vonsisted of a band of iron-impregnated sandy
sediment. Similar occurrences were described tn many
sand quarries in Tertiary sands in the Adelaide region
by Harris & Olliver (i964) and Olliver & Weir (1967).

Linerite as a weathering product

The association of lateritic crusts with weathering,
profiles (Walther 1889; Maclaren 1906) was iniroduced
lo Australian studies by Simpson (1912) und Walther
(1915). They espoused the view that laterite formed as
iron dnd dluminium oxide effloresences were
transported if solution from the water table by
capillarity, Walther (915) assigned the term ‘laterite’
to the complete profile,

However, laterite in South Australia has most
commonly been considered to be an indurated
ferruginous horizon ina Weathering profile (Stephens
1946, Hallsworth & Costin 1953, Connah & Hubble
1960), which is quite different from the original laterite,
described by Buchanan (1807) as a low-level
sedimentary deposit consisting of massive, unstratified
iron-rich clay material, full of cavities and pores, which
hardened once cut into blocks and exposed to |he
atmosphere.

Lang (1965) followed Hallsworth & Castin (1953),
restricting the term ‘laterite’ to crusts associated with
well-differentiated profiles apparently formed by in situ
relative accumulation of iron oxides, “Ortsten’ was used
by Lang (1965) to describe crusts developed by laterally
derived ubsolute accumulations. Where ortstein crusts
formed above weathered profiles and simulated in vita
weathering profiles they were called ‘duricrusts. Lang
considered that laterites on the oldest surfaces were
only occasionally developed frojn materials
recognisable in the pallid zone, and he assumed tat
a discontinuous layer of Tertiary sediments overlay
older rocks throughout the lateritic area. Maud (1972)
also considered that only soils containing ironstones
overlying mottled and pallid zones should be regarded
as laterites. This definition has sometimes been
ascribed the descriptor “ue laterite (e.g, Bourne
TE),

LATERITE IN SOUTHERN SOUTH AUSTRALIA 5

Classificanan of laterite

There has. been litthe attempt to classify lateritic
matenals m South Australia. Teale (1918) used the term
‘ironstone’ to describe ferruginous materials, which
were noted to affect all materials except alluvium. They
were categorised into four main types: Joose,
concretionary gravels in deposits up to} m thick, iron-
cemented sands and gravels, ferruginised slates and
quarizites, and Jateritic ironstone forming hard sheets.

Laterite was categorised by Forrest (19694) as "fossil
or relict’, which referred to the complete normal laterite
profile, ‘truncated, where the upper horizon was
absent, ‘immature’, where the percentage of tron im the
cupping was low and the underlying bedrock was only
partially weathered, ‘derived’, when the cappiny, had

) Forresr, G. 1 (1969) Geomorphological evolution of the
Bremer Valley, B.A. Hons thesis, The University of
Adelaide (unpubl.),

4 BourmAn, R. P (1989) Investigations of ferricretes and
weathered zones in paris of soufhern and southeastern
Australia ~ A reassessment of the laterite concept. PD, |
thesis, The University of Adelaide (unpubl),

laterite crust.

been derived from the reworking of higher crusts: and
where this reworked capping rested on weathered.
bedrock, and. ‘terricrete’ where an iron-rich crust
incorporated partially-rounded quartz pebbles and
overlay fresh bedrock.

The use of the term ‘ferricrete. coined by Lamplugh
(1902) 10 deseribe a ferruginous conglomerate, has been
extended to apply to all iron-cemented and indurated
continuous horizons and crusts in preference to laterite
by some workers (e.g. Milnes et al. 1985. Bourman
19894). Perrierete was classified by Milnes ef al,
(1987) and Bourman (19894) as simple types, which
included ferruginised bedrock and clastic and organic
sediments, and complex. types such as pisolitic.
nodular, slabby and vermiform ferricrete, The diflerent
forms of ferricrete were noted to display differences
in micromorphology, mineralogy and chemistry that
reflect the nature of the parent material, environmental
conditions during iron impregnation and subsequent
transformations during landform evolution. Mottled
(Fig, 3) and bleached materials (Fig. 4) were regarded
as having developed independently of the ferricretes
by Bourman (19894).

6 ROBERT P. BOURMAN

Processes of laterite formation

It is necessary for the various potential processes
of laterite formation to be understood so that more
reliable interpretations of ages and relationships to
underlying companion materials can be provided. For
example, does laterite formation require a peneplain
surface, as many workers in South Australia have
claimed? Furthermore, with respect to pisoliths, is it
possible to distinguish formation in place from
transported origins? Many theories of laterite origins
have concentrated on vertical translocations of minerals
in the regolith that involve capillary rise, vertical
leaching and fluctuating water tables. However, Milnes
eral, (1985) and Bourman er al. (1987) demonstrated
that ferricrete development in southern South Australia
has been almost exclusively related to lateral physical
and chemical transport to, and accumulation of iron
and or aluminium minerals in, discrete preferred sites,

Capillarity
Teale (1918) favoured the role of capillarity in laterite
formation. He concluded that laterite formation

depended upon a ferruginous rock or subsoil for an

iron source, dissolution of iron, largely by organic
acids, and a hot season to ‘pump the iron salts’ to the
surface, causing oxidation and precipitation of limonite.

Laterite development by prolonged chemical
weathering during the late stages of the cycle of erosion
(Davis 1909, 1920), on a Miocene continental peneplain
with sluggish surface drainage, in a seasonally dry
tropical climate that encouraged capillary rise of iron
and aluminium in solution, was described by
Woolnough (1927), who had widespread experience of
duricrust in Australia. He considered the ‘Upland
Miocenes’ of South Australia to be ‘veritable Duricrust
albeit of somewhat aberrant type’ (p. 46). He noted
similarities between ferruginous cappings in the Mt
Lofty Ranges with examples in Western Australia, and
regarded some of the ferruginous materials on
highlands as ‘thoroughly typical lateritic crusts’ (p. 46)
and that the ferruginous surface of much of the ‘Mount
Lofty Plateau’ was underlain by highly decomposed
arenaceous rocks, similar to those related to ‘Duricrust’
Laterite, ‘Upland Miocenes’ and ‘Desert Sandstone’
were thus considered as contemporary and equivalent
duricrusts, resting on weathered rock materials
(Woolnough 1927).

Fig. 4. Bleached and kaolinised Precambrian Aldgate Sandstone exposed by quarrying at Longwood, in the South Mount

Lofty Ranges at 400 m above sea level. The depth of the section is 30 m.

LATERITE IN SOUTHERN SOUTIL AUSTRALIA 7

The capillary model of laterite lormation should
result in the reversal of soil A and B-horizons, with
the surface laterite being the illwyial B-horizou and the
underlying iron-depleted clay being the eluyadl A-
Honzon, Thus, this model requires. the eogenetic
formation of the complete laterite profile.

Leading

The interpretation of laterite as the B-horizon of i
tassi| podzolic soil was pursued by Preseoit (1931) in
view of evidence of the dominance of leaching of bases
in lmerite profiles, 45 opposed to capilary uplilt,
evaponttion and surface precipilalion of iryn und
Mumunum oxides. Tropical podzolisation became the
most pervasive view on laterite formation im Sout
Australia, with the laterite horizon being regarded as
a fossil Tuvial B-horizon where laterite occurs in preity
of andity.

Johns (196la) proposed that poorly drained soils on
Eyre Peninsula were Jeached (which appears ty be
contradictory). during presumed humid pluvial
conditions of the Pliocene, leading t0 the accumulation
of iron oxides and the ir ara formation of Lalerile,

Combinations of water table movements, leaching and
eapillarity

Both Whitehouse 0940) and Stephens (146)
conturred with the general padzolie ortyin of laterite
bul envisaged soufees of jron not only from the
overlying leached Ashorizon but also trom iron-
depleted, weathered bedrock by water lable Tuctuarions
and cupillury rise,

Hallsworth & Costin (1953) questioned that the upper
podzolised layers of southern Australian laterites
comprised parts of original laterite profiles, and
suggested that they resulted from intense leaching after
lateritisation. However, Prescott & Pendleton (1952)
had pointed out that, in spite of current semt-aridity.
relic podzolic soils with tronstone gravels in Western
Australia remain acid, m-emphasising their hypothesis
of the pedogenic origin of laterite,

The interpretation of laterite as the indurated, iron-
rich B-horizon of a fossil, podzalic soi) profile was
favoured by Stephens (1946), who proposed a dynamic
pedological model of soil Jormution, subsequent upon
dissection of the Jateritic regions in South Australia.
He regarded Jaterite us a pedogenic material and
suggested that ferruginous concretionary gravels
accumulated in the soil profile in the zone of oscillating
seasonal water table as a result of alternating reducing

> Ravine, F H_ (1959) The tegional geography of Kangaroo
Island, Pl, D. thesis, Australian National Dbdversit,
Canberra (unpubl, },

und oxidising conditions. He associated the water table
Nuctuations with a low teltef und a humid eluate.
Under these conditions the concretionary gravels were
assumed to form an indurated horizon by their
progressive enlargement and coalescence, Later uplift
and dissection of the landscape was postulated to
explain. the luterite mantling remnants of former
peneplains. A major contribujon (6 pedologival studies
wus made by Stephens (1946) who recognised the
influences of soil development on both the im sip
weathered bedrock and the eroded. transported debris.
This model proved to be very productive for other
pedologists (e-g. Northente 1946; Northcote & Tucker
48; Rix & Hutton 1953),

Stephens (1971) later moditied some of his ews on
laterite formavon when he investigated a possible co-
genetic relauonship between sicrete and laterite. He
considered that laterite formed by the aecumulation of
hydrated oxides, Kaolinisation of mottled and pallid
zones and the acidification of the whole profile, with
pronounced leaching losses of silica and bases, Luterite
was noted to form by hoth relative and absolute
accumulation bat he believed that relative accumulation
was predominant. He aisce concluded that although
laterite formation was associated with a fluctuating
water table. i was nol dependent on either pertect
planation surfaces or opica) climates. yiews that have
largely been ignured in the local literature.

Wenting and drying processey

Th opposition to Prescot & Pendleton (1952), Bauer
(19595) favoured the view that laterite may currently
be forming in southern Australia where the regolith
is affected by wetting and drying. He postulated that
under these conditions lerraus iron would iigrate
upward during waterlogging and conver fo stable ferric
iron in dry, oxygenating periods. He recognised a ready
source of iron from decomposing country rock und i
lemperalure regime warm enough to allow the
reduction, jnigration and oxidation of iron.

Weathering transformations vf materials rich in ferrous
vn

Mawson. (19072) described a large saucer-shaped
body of bog iron ore, with a maximum thickness ol
(Om, forming # flattish-epped hill about 200 m above
sea level at Wadella Springs on Eyre Peninsula. He
concluded that the deposit had originated from spring
waters, with iron sulphate haying derived from the
oxidation of underlying pyrite bodies. Thus Mawson,
without confusing the occurrence with laterite, had
observed and explained the fammanion ata distinetive
type of ferricete,

8 ROBERT P BOURMAN

The formation of ferruginous crusts, in such places
as the Telford and Murray basins, by weathering
transformations of minerals containing, ferrous ion
such as glauconite, siderite, chamosite and pyrite to
ferric iron minerals dominated by goethite has alsa
been recorded (Bourman 1989, Bourman ef al.
1995).

Lacustrine laterite

The view of ironstone formation as lacustrine (€.4.
Ferror IOI) or swamp deposits on a peneplain surface
close to sea level, with the water table close to the
vround surface was suggested lor South Australian
samples by Segnit (1937). He also noted the occurrence
of three types of fronstone cappings on high Icvel
ground and Slopes in the Mt Lofty Ranges Vesicular
ferricrete, fonned by iron oxide replacement of organic
material in former swamp environments, has been
recorded (Bourman 19894) in various landscape
posiuons in the Mt Lofty Ranges (Fig. 5) and on
Kangaroo Island.

Relative and ubsolute accumulation: jn situ versus
/ransported laterite

Luatertte formation by relative (d7 yftv) and absolute
(lateral) accumulation has long been recognised with
different workers attributing differing significance to
(hese processes. For example, Stephens (1971) attnbuted

laterite formation dominantly to relauve accumulation,
Crocker (1946) agreed with the in side formation of
some ferruginous gravels but considered that some
others have secondary origins, Milnes er a/ (1985) and
Bourmun ef a/. (1987) presented evidence of dominant
lateral transport in ferricrete and pisolith formation in
the Mi Lofty Ranges, although the possibility uf int vite
formation was not rejected (Fig. 6), Johns (1961a) also
conjectured that most of the sediment deposited on the
coastal plains and Central Basin of southern Eyre
Peninsula was material resorted from lhe uplands and
included pisolitic or massive ironstone gravels. Johns
beheved that during Jatentisation the previously
penepluined basement rocks underwent deep
weathering, ferruginisauien and kuvolinisution,
Lithologieal variations in the basement rocks were
thought to have had no influence on the final
weathering products.

Maud (1972), following d’Hoore (1954) proposed
absolute and relative sources of iron and aluminium
oxides for the formation of laterite, The accumulation
of iron and aluminium oxides was attributed either to
the renioval of silica and bases or their accumulation
from Outside sources Well-developed _ lateritic
ironstones on Permian glacigene sediments were
explained by the concentration of iron oxides from
lateral sources, Whereas thinner crusts on pre-Permian
rocks were ascribed to in situ weathering losses of silica
and bases (Maud |972), Maud (1972) believed that
following landscape rejuvenation and lowering of the
water table, the zones of iron-enrichment irreversibly

é —

Fig. 5. View of bulldozer excavation om Peeralilla Hill showifg terruginous crust of vesicular ferricrete and light-coloured
clays (bottom lef ot photograph) that melude calcite and barite. This deposit of ferricrete occurs on the summil surfitce
bul below (he level of surrounding hills, Borehole evidence indicates that this deposit is underlain by sandy sediments,
Excavation 2.5 m deep.

LATERITE IN SOUTHERN SOUTH AUSTRALIA 9

hardened into lateritic ironstones. Brock (1964°) also
agreed with d’Hoore (1954) that dissection of lateritic
terrain, accompanied by lateral water movement, may
have redeposited iron oxides on gentle slopes to form
cappings.

Wopfner (1972) carried out an analytical investigation
of mottled materials that in other contexts have been
referred to as lateritic. He discussed maghemite in
mottled Cainozoic sediments at Hallett Cove, where
both primary and reworked maghemite were identified.
Maghemite was reported from two locations: small
amounts (2%) of maghemite in conspicuous red
mottles, within medium grained white sandstone, were
used as evidence of in situ formation, whereas
maghemitic sub-rounded ironstone pebbles in a
conglomeratic horizon were considered to be indicators
of reworking. The profiles and crusts were considered
to be genetically related with the conglomerate forming
by reworking of an original in situ crust.

© Brock, E. J. (1964) The denudation chronology of Fleurieu
Peninsula. M.A, thesis, The University of Adelaide
(unpubl.).

Many soils associated with uplifted peneplains in
Australia have been noted to contain concretionary
ironstone gravels, attributed by Prescott (1934) to
former wetter periods when waterlogging of soils and
shallow water tables were more common than at
present. Chemical analyses of ferruginous gravels were
interpreted by Prescott (1934) to demonstrate the
concretionary character of the ironstone gravels.
However, many pisoliths in southern South Australia
appear to have formed dominantly by disintegration
of ferruginous materials such as mottles, followed by
physical transport and modifications in soils resulting
in increases in iron content as well as a mineralogy
dominated by hematite and maghemite (Milnes er al.
1987; Bourman er al. 1987), Transported pisoliths
typically are associated with stone lines, have different
chemical and mineralogical compositions to
surrounding matrix materials and display multiple
rinds. Milnes ef al. (1985) also considered that
ferricretes in southern South Australia, as well as
pisoliths, are dominantly remnants of iron impregnated
sediments, originally formed in former valley bottoms
and depressions,

Fig. 6, Road cutting on the Victor Harbor-Cape Jervis road west of the Waitpinga road, exposing bands rich in pisoliths,
at a depth of about 1 m, in ferruginous sandy sediments of probable Pliocene age and of aeolian origin. Other pisoliths
occur at the ground surface and in the upper soil mantle. The pisoliths at depth contain only goethite, whereas those
at the surface have higher iron contents and contain dominantly hematite and maghemite. Geological hammer 33 cm long.

1”) ROBERT P BOIRMAN

Although there 1s general agreement that terruginous
materials can form both by processes operating i sity
and those related to transportation, there jis beer
confusion in the use of the term fsiti. For example,
some workers have considered that ferrerete, formed
during landscupe dewynwasting. which involves both
vere) and lateral movement ot clasts, fotmed in site.
Such derrieretes may be benter regarded as residual witht
ia sith weathering applying more strictly only to
ovolumimous weathering (Bourman W94b),

Laterite profiles

The nermal laterite. profile

Throughout the South Australian htenttane, following
Stephens (1946), runs the thread of the normal laterite
profile, which bay influenced many palaeo climatic and
palaco-enyironmental reconstructions. Only rarely have
stuclies departed Jrom this model. The norrial lateritic
profile (Stephens 1946), was envisaged as essentially
a podzol with A, Band C horizons of cluviation,
Hluyidiocand weathering, with an accessory lalenle
honzon usually above a clayey B-horizon, Ovcasionally
several laterintic horizons were noted in one profile
Stephens believed that the normal laterite profile was
restricted to southern Australia, m Queenstand laterite
awwas thought to occur as an horizon in red-eurth proliles
(Arvin 1939: Whitehouse 1940), which vontined
silicified zones. suggesting the incomplete removal of
iissolved silica.

The model presented by Stephens his considerable
ment as emphasises the dynamic nature of landlorm
change and pedogenesis. However. its dependence on
the widespread occurrence of a former murnal laterite
profile related to former regional water tuble
Tluctuations, 1s unrealisuc and has led to siipliste
explanations of landscape development, Furtherninre
(here are various Objections (o the view thal (he onginal
laterite is the illuvial horizon of a fossil podzotic soil

Widespread Jateritic souls on the clevated pencplain
of Kangaroo Island, the Mt Lofty Ranges. Yorke
Peninsula and Eyre Peninsula were reported by
Crocker (1946), who obseryed that they contained
considerable percentages of logse and indurated
lateritic tronstone gravels. Some of these gravels were
considered to have formed in sine, but on dissected
imaginal slopes secondary origins for them were
suguested. Crocker (1946) followed the view of Prescott
(O31) that laterite is the fossil illuvial horizen of a
lfopical Pliocene podzohe soil, Thus he reiterated the
then current Lhoughts about laterite and further
promulgated. the association of laterite, peneplains,
(rapical climates and the Pliocene (or Tertiary}, thereby
setting the stage for jhe generation of circular

urcuments. Spriggs (946) concurred with Prescott (1931)
and Crocker (1946) vonceming laterite genesis,

The pedogenic orgin of luterite was promoted hy
Northente (1946) and Northcote & Tucker (1948)
These workers mupped and described a relic normal
laterite profile of Phoeenc age. the Eleanor Sand. on
the lateriGc plateau of Kangaroo (sland. Cracker (1946)
commented that latent residuals on Kangaroo Island
were covered by grey and white siliceous sands derived
Irom resorted A-honzons, originally developed on
Pleistocene coastal ealeareous sand duncs. However.
Northeot (1446) claimed thar the constant rate ol
course to fine sand throughout the protile indicated that
it had formed iy sita and that the surface lad oot
received accessions oF wind Hlown sand, Consequently
he regarded the Eleanor Sand as a relatively
undisturbed fossil soil of Pliocene age. with a lateritic
horizon developed i site and preserved Onan uplifted
peneplain. However, Muleahy (960) has stugestod that
such satidl ray TOL be fossil, but may have derived! from
lunerite destrucnion, thus yielding a similar grain size
analysis to that determined by Northcote. ‘Twidale
(1983) considered that this sandy A-horizon provides
evidence tor the preservation of an original Mesovzae
jateme prolile,

Rix & Hutton (953) regarded the suinmil surface
in the south Mt Lofty Ranges as a block-faulted,
uplifted and dissected peneplain. They follawed Sprigy
(946), vorsidering that by Farly Tertiary times
Precambrian rocks had been reduced ter 4 base surtice
and subsequently buried by a Tertiary lacustrine and
marine covermass. The soil pattern suggested to ther
that a further cycle of erosion had removed the grealer
part of the covermass. leaving isolated areas of varying
extents thereby creating a new peneplam, with
remnants of Tertiary deposits preserved in topognaphic
lows. They postulated laterinsation of soils on the
peneplain, prior to major faulting and dissection,
concurring with Whitehouse (1940) that there had been
comlemporanedus. laterite formation throughout!
Australia in the Pliocene. Residual literitic soils were
only mapped on hill summits and spurs 6a they
suggested that erosion had removed most of a larcritic
sandpluin following uplitt and dissection.

One soil, the Yaroona Gravelly Sand. was regurdel
#s an Original laterite profile and described as 3 massive
band of laterite 22-30 cm thick. containing water-
washed grits, gravels and sands, unconfurmably
overlying kaolinised Precambrian shales (Pip. 7). Rix
& Hutton (1953) deseribed other residual podzols un
the area as exhibiting normal profiles of ferruginuns,
motied und pallid zones, overlying unWweathered
country rock, These workers were strongly influenced
in their interpretations by the aormal laterite profile
mode! of Stephens (1946) and presented a convoluted
explanation of an anonialous laterite profile. preserved
in a road cut south of Clarendon, in order to agcount

LATERITE IN SOUTHERN SOUTH AUSTRALIA a

for a mottled zone overlying a laterite zone. The section
can also be interpreted as a geological sequence of
Precambrian rocks weathered in pre-Tertiary times,
and overlain by fluvial gravels, grits and sands of
Eocene age (Mawson 1953). Subsequently these
sediments were partially silicified and superficially
stained red by small amounts of iron oxides in
groundwaters. A thin grey soil with pisoliths occurs
at the surface. The above example demonstrates how
complex deductive arguments, within framework of the
model of the normal laterite profile, were used to
introduce events, for which there was no evidence, in
order to explain apparently aberrant observations,
Despite this, Rix & Hutton (1953) produced a detailed
soil map.

In southern South Australia the normal laterite
profile of Stephens (1946) has been given excessive
consideration, sometimes resulting in simplistic
interpretations of landscape development. This has
occurred despite observations indicating great

variability in lateritic weathering profiles and despite
the view of Stephens (1971) that the ‘normal profile’ is
the exception rather than the rule, Bauer (19595) and
Alley (1977) disagreed with the interpretation of laterite
as a tropical fossil soil profile developed on a
peneplain. However, their views have not been
generally accepted.

Polygenetic profiles - Alternatives to the normal laterite
profile

STRATIGRAPHIC APPROACH TO INVESTIGATIONS OF LATERITIC
MATERIALS

Firman (1967a, b, 1976, 1981, 1994) placed weathered
zones and palaeosols within a stratigraphic framework.
For example, he gave formal status to ferruginised
clastic sediments and bedrock weathering profiles
consisting of sesquioxides of iron and forming
ironstone crusts, by introducing the name Yallunda
Ferricrete. The Yallunda Ferricrete was reported to
exceed | m in thickness in its type area at high levels
over interfluves of the Lincoln Uplands (Eyre
Peninsula) and on remnants of old high surfaces
elsewhere. The term ‘ferricrete’ was used to describe
ferruginous layers and crusts both independent of, and
in association with, weathered profiles. Firman (1976)
interpreted the various zones of the so-called normal
laterite profile as having formed by different processes
at different times, with the profile as old as the initial
transformation of the parent material.

Ferricretes in various stratigraphic situations,
including ferricrete above and below Lower Tertiary
sediments in the Barossa Valley, as well as ferricretes

Fig. 7. Section in road cut near Clarendon, exposing the Yaroona Gravelly Sand of Rix & Hutton (1953), showing angular
unconformity with ferruginised pebbles, grits and sands of Eocene age overlying weathered, bleached and partly kaolinised
Precambrian meta-siltstones. Section is approximately 3 m high.

12 BERT & BOURMAN

in the highlands of the Mit Lotty Ranges and the Linealo
Uplinds were recorded by Firman (l987a). He alse
suggested thal there were equivalents of upland
lernieretes in the sedjmentary succession of the Murray
Basin. These included oolite siderite-neh sediments
and laterite in the Early Pliocene Bookpurnong Beds,
ys well as ferruginous beds and cappings in the Late
Plipcene Parills Sand near the Victoria-South Australia
border, Some ol the lerricretes, however, have resulted
from the relatively recent oxidation of pre-existing inon-
nich sediments containing glauconite and siderite and
cannot be used as reltable sor! stratigraphic markers.

Firman (1976) consideyed that between Permaitn und
Early Tertary times, some 200 Ma, the Mt Lofty
Ranges region was a land mass experiencing protonged
wealhering and erosion, so that by Barly Teruary noes
a subdued and deeply weathered landscape had
developed. Associated bleached profiles were
considered 9 haye onginated in the Mesozoic, A range
of different ages of weathering and lateritisation was
reported, Decomposed, bleached or mottled bedrock
underlying Eocene sediments was ascribed to pre-
Tertigry weathering, a laterite profile developed in
Bocene gravelly sands was used to indicate pos(-Hocene
weathering: silicifiec and ferruginous zones in Early
Pleistocene sediments, overlying older bleached zones
were argued to have equivalents in laterite protiles in
the adjoining uplands, and ferruginisation in
carbonaceaous and pyriuc Encene sediments was
wiribuled tu recent exposure and oxidation.

The work of Firman 1s significant in attempting In
establish stratigraphic ages for different weathering
ieatures. Nevertheless, correlating weathering phases
simply on shapes. sizes and colours of muottles may
be unreliable, as similar weathering patlerns occur m
profiles of different ages. Furthermore. Firman
observed modificaGon of Some profiles by later
weuthering, obscuring earlier weathering products.
Firman apparently took no wecount of local
environmental conditions, which may have favoured
synchronous bleaching in one area and imotling in
another Various questions remain unanswered, such
us What happened to the iron derived from the
bleaching of the Arckaringa Palacosol; where did the
iron for the development of the San Murino Palaeosol
come From; nd haw was it concentrated in diserete.
bul more-or-less uniformly distributed mottles witht
previously bleached bedrock?

POL YGINIIe PROFILES AND CONTINUAL WRATIIDRING Mant
op Ferricker FORM ation

Milnes ef af. (1985, 1987) and Bourman e eal, (1947)
combined investigations of the field relationships of
ferricretes and weathered zones with wicra-
morphological, chemical and mineralogical analyses
and questioned the former development of noroul

Jaierte profiles. These studies have suggested thar there
was complex reworking and continuous Weathering of
relic landscapes since the Early Mesozoic, and [hat
ferricretes are dominantly remaants of iret
impregnated sediments of ancient valleys or
Jepressivns. Some ferricretes may be the culmination
of processes beginning in the Mesozoic bul sill
proceeding, resulting in the repeated dissolution. break
up and neo-formation of lerricretes. as well as the
ongoing and current formation of mottles and bleached
7ones,

Some ferricretes may have developed as suggested
by MePurlane (976). who postulated fern¢rete
develupment by the surface aceumulation of
ferruginous materials during landscape downwasting,
the formation of gibbsitie-rich zones in near-surface
situations and continued development of ferricretes and
bleached zones. after uplifi. However, some other
features of her model do not fit the observations in
South Australia; there 7s evidence for some bicached
zones and mottled zones being older than the ferricretes
(Bourman 19894) and no evidence has been observed
ol progressive development of profiles, with honzons
having formed from progenitors resembling those
currently beneath therm, An extension of this model
is the continual weuthering hypothesis of Bourman
(989%, 19934), which propases. Ongoing epigenetic
transformations of ferricretes und weathered zones,
with rates of change influenced by climate and events
such as teetonism, sedimentary burial und submergence
beneath Jakes and the sea

Topographic requirements for laterite formation
Feneplain concept in laterite development

Many workers have associated laterite formation and
preservation wath penepluined surfaces. However,
Sprigg (1946) considered it unwise to associate laterite
formation with peneplanation, which implied formation
ower a very long peridd, since be believed thal
lateritisation secupied only a relatively shor rime span,
This interpretation has important implicanons for
landscape evolution as laterite formation would first
require the development of an extensive planation
surface, The peneplain concept of laterne development
begun early (Beason J8iL: Mawson J907b; Teale 1918:
Woolnuugh 1927), and bas persisted (Campana Av
Wilson 1954; Brock 19649; Ward 1966: Twidale 196b,
983, Maud 1972), For example, the summil surface
ofthe MU Gofty Ranges was interpreted as un Early
Tertiary differentially uplifted and dissected ‘peneplain,
surmounted by monadnocks such as Mt Lofty. Mt
Barker and Mt Gawler (Benson IIL). Tate (1879)
attributed the discontimuous distribution of the “Lipland
Miocenes’, separated by deep ravines, to extensive
denudation after uplift of the ranges,

LATERILE IN SOUTHERN SOUTH AUSTRALIA 12

Mawson (9076) extended the peneplain concept to
Kyre Peninsula where he deseribed peneplains in the
Port Lincoln are at about 00m ancl 6 m above sci
jevel. He equated inottled clay beds underlying the
lower surface with lreshwuater Miocene beds neat
Adelaide,

Some extremely perspicugious comments on the
nature and formation of ferruginous materials i the
south Mt Lofty Ranges were mathe by Teale (1918) and
his work represents the niost comprehensive, detailed
and objective discussion of iron -owides: mong all of
the early mvestigators, particularly oo classification
and theories of origin of ironstone crusts. ‘Teale (YTS)
inlerpreted the surumil surface of the ranges as
dislocated and eroded remnants of a former extensive
pencplamn Woolnough (1927) sugvested thal the
distribution of reninants of the terrestrial Upland
Miocenes agreed completely with the physiygraphic
vonditions postulated for duricrust formation ic, a
peneplain with sluggish drainage,

In the southern part of the Mt Lofty Ranges,
Campans & Wilson (954) described a planiton
surface at levels up to 420 m above sea level. as a pre-
Tertiary peneplain, uplifted during Termiary and
Quaternary times and deeply dissected by subsequent
cyeles of ctosion and Brock (19645) identilied
remnants Of an ancient lundsurface an the spine of
Fleusicl Peninsila covering on aren of 25 krm*. the
remnants Were described as having lithe relief und a
capping of the normal kilerite profile of Stephens
(1940). Ward (1966) also deserihed fal surlaces
preserved on ihe crests and gentle back slopes of the
western blocks of the Mt Lolty Ranges as relics of it
pre-deformational ML Lotry peneplain, mantled by deep
weathering and lalevite, Twidale (1968) described tie
summit surface of the Mt Lofty Ranges as a lalerny ined
peneplain, surmounted by a few residual remnants or
monadnoeks,

Today the tert “peneplain’ is rarely used in
geomorphic literature because, among other things, 1
carries with itan undemonsteable, Highly thearetical
mode of genesis. The terms ‘erosion surlice’ or
‘planation surface’ are preferted.

Irregular surfaces

Not all workers have considered peneplains
necessary for laterite formation. Tf penepliin surtaces
are necessary for laterite dormation, the unplieanou
is that latertte formation follows peneplinaunon,
although irregular surfaces suggest that laterite can
form during, and as a result of Jandsecape
downwasting. Some investigatory such ax Curmpana
(1955) have demonsirated great complexities ard
irregularities in Weathering and landscape evoly|ion,
Working, near Gawler in the north Mi Lofty Ranges.
Campana (1955) noted a leached lateritic soil overlying

eneisses, schists and Tertiary Nuviatile deposits resting
on it pre-Tertiary weathered erosion surface, He
reported gravels and coarse sands cemented by iron
oxides widiin the ‘Tertiary sediments. The mapping of
the Tertiary (Early Eocene) strata m this area indicated
thal deposition oceutred ina system of lakes gind rivers
on a weathered surface of rodersle rebel, above which
ridges of harder rocks projected, Campana (1955)
considered that the non-marine strata and older tocks
had been subjected to widespread lateritisation between
the Early Eocene and the Miocene, The sequence
outlined by Campana (1955) illustrates pre-Tertiary
weathering, and bleaehing of basement racks. the
deposition ol ‘Tertiary letrestrial sediments over a
dissected landscape. differential ferrupinisation ob
suilable host rocks aud Lhe likibitton of this by marine
submergence,

Bauer (19595) noted that repardless of elevauon, the
laterite profile. the Eleanor Sand, oeeurs on reas of
low reliel and poor drainage that would lave suited
periodic waterlogging and drying (Mig. &). Thus he
thought (hat toporraphy night buve been important in
assisting the formation of 4 distinctive soil in two
differing physiographic locations, that is, on a stepped
topography with fat treads. but nol mecessanily
peceplain.

Lung (1965) reported on soils und geomorpholugy
of the Yundi area within the south Mi Lofty Ranges,
His work represents a departure from that of many
eurlier workers us he invoked different types of
weathering, erosional and sedimentary mtluences to
explain the eurrent landscape and he envisaged
lateritisation and duricrust formation as procecding
over Jong periods of time, arn landscapes of variable
relicf und positions abave seu level, In the same area,
Maud (1972) noted Literitised surfaces occurring across
infilled glacial yalleys and correlated therm with fhe
summit surface despite their lower lindscupe posbons,
The gentle non-teetonic tnelinations of jronstone
coppings were reparded as original valley
morphologies, aid Mand (1972) concluded that the
original erosion surface was one of considerable: relief,

Bourman (19895, [994a) presented a model of
laterite formation involving an original landscape of
some rebel that provided lateral local cnvironmentil
variability This resultect in bleaching of higher parts
of the landscape and iron accumulation on plateau
margins, indepressions, swamps and valley bottoms.
Primary iron minerals mobilised in sub-surface zones
affected by water lable fuctuutions were concentrated
in hemaftitic notes. Landscape downwasting
concentrated dnd fragmented mottles at the sitface.
further weathering modified them, formed pisoliths.
cemented (rer lo Jor fertierete at the surface and
further modilied the ferricrele, Porhons of (ie sult
surface af the Mc Lofry Ranges have been continually
uffccted by Weathering and erosion since the Permian

crete = Miocene Jimestane

c ferricréte

Vermiform ferri

Pisoliti
Pisoliths

pia

ROBERT P, BOURMAN

salle This suggests ferricrete formation during landscape
ace. “S evolution rather than being dependent on the presence
ns of a planation surface, The continual weathering model

postulates ongoing variable weathering, interrupted by
tectomie activity, sedimentary bunal or marine or
lacustrine submergence,

Coastal dunes

Climatic conditions required for laterite formation

The vast majority of workers has equated laterite
formation with a hot, seasonally dry tropical climate
favouring the operation of intensive weathering
processes. Low topographic relief and tropical climates
were considered ideal for laterite formation, generating

o many circular arguments related to the laterite-tropical
climate-peneplain association,

A tropical climate with pronounced wet and dry
seasons, such as that of Darwin, was considered ideal
for the formation of laterite by Walther (1915). This
view has persisted. Stephens (1946), Sprizg (1946),
Crocker (1946) and Northcote (1946) associated laterite
formation with a pluvial period in the Pliocene und
Johns (196la) believed that low relief and high tropical
temperatures had favoured the removal of silica, with
seasonal oscillations of the water table leading to the
concentration of iron oxides, More recent workers such
as Bourne (19742), Daily er al. (1974), Twidale &
Bourne (1975a), McGowran (1979a), and Twidale
(1976b, 1983) also favoured torrid, tropical conditions
for Jateritisation. The tming of lateritisation has
commonly been associated with independent evidence
for tropical climates. For example, Twidale & Bourne
(1975a) noted that palaeontological considerations
favoured the Triassic as providing the most suitable
humid, tropical climatic conditions for the formation
of laterite in the Mt Lofty Ranges.

In marked contrast. Firman (1981) proposed different
climatic conditions for separate parts of the profiles,
For example, the bleached zone of the Arckaringa
Palaeosol was not considered to have been genetically
associated with younger ferruginous zones but to have
preceded the development of mottles and ferricretes,
He ascribed bleaching to early cool climates and
ferruginisation to tropical conditions.
uo Other workers such as Bauer (19595) and Campana

& Wilson (1954) considered that lateritic material might
be forming at present jn southern Australia so that
climate for lateritisation need be no different from that
of today. Maud (1972) also argued that the process of

Mt Stackdale
3

Mount Taylor Plain

130m
~
Kilometres

Mt Taylor

surtace:

Fig. &. Cross section of the Mount Taylor Plain, Kangaroo
Island, showing the relauonships of identical vermiform
ferricrete on a high pre-Miocene summil surface and a low
& post-Miocene surface,

wound

LATERITE IN SOUTHERN SOUTIT AUSTRALIA 1"

iron oxide-envichment of sediments is curreatly
proceeding. on broad valley floors in the Me Lofty
Ranges under current climatic conditions.
Furthermore, there is considerable evidence of miodern
iron mobilisation and precipitation in southern South
Australia (c.g. Bourman 19899, Ferguson ey al. 1984).
«o thet a hemi tropicul climute need not be a pre
requisate (or bleaching and iron enrichment,

The role of climatic influences in the formation of
ferruginous and siliceous duricrusts was. examined by
Alley (977), who provided evidence that both laterite
and silerete formed together, for at least some time
during the Tertiary, on identical strata and under similar
climatic conditions, suggesting that.some other factor(s)
must have controlled the processes of weathering. Only
the base levels of erosion differed hetween the silereted
and [ateritised surfaces and stlerete developed on a
surtace, the drainaye of which towed sluggishhy int)
Tertiary lakes. Pulynological data were interpreted by
Alley (1977) to demonstrate that the Kovene appeared
to huve heen warm and temperate with a very high
rainfall and that the Miocene was similar, with perhaps
Slightly warmer lemperalures and a slightly lower
precipitation. The concentration of silica at the
landsurfaee Was attributed to high alkalinity, slow
proundwater movement und a high water table chase
to the lakes. Alley (1977) concluded (hat laterjte arid
silorele covexisted for part of the Eurly Cainozoie in
adjacent drainage basins, Consequently. laterite and
silerete were got thought to form through the
mechanism proposed by Stephens (1971) which invelved
the formation of silerete by deposition of silice in Ury
aones aller faving been derived trom Jateriue
weathering elsewhere, Furthermore, the view that
luterie is ussociated with tropical conditions snd
silerete with aridiry was not supported because both
fortned ia similar climatic and biotic regimes; anly the
base levels. and groundwater conditions yaned.

Using chemical (Auton 1977), palynological and
suauigraphic evidence to suppart their argunient
McGowran et a/, (978) disagreed with Alley (1977)
that laterite and silerete formed concurrently on similar
rocks and under broadly similar climatic conditions
from Recene to Miocene times, However, Alley (978)
couniered the arguments presented by McGowran ev
al, (1978) and made a valuable contribution to the study
of latente venesis by hay hlighitine the influence of local
topographic and groundwater conditions in ats
formation, a well as questioning Clioatic aiflucrices
on laterite and silcrete development,

Minerals as climatic indicators af lateritisation

As noted above lateriasution is cotamonly associated
with intensive weathering under tropical climatic
conditions und certain minerals are suggested 4s
indicators of climatic conditions. Kor cxaniple, Woptnes
(1972) sugeesicd (hat miaghemite in motiles is a climatic

indicator, oripinating by thermal dehydration of
lepidoerowite Jormed by oxidtion onder fluctuate
water table levels and warm climatic conditions, He
voneluded that lepidocroente and goethite myy have
formed as pels that were subsequently dehydrated and
erystallised as maghemite and hematite under
conditions of low telief, warm climate and heavy
seasonal yalnfall,

However, mayherte in laterite rootiles is very rare
in southern South Australia as they are clominantly
henatine (Bourman 19894). Moreover in lateritic
arcs of the Mt Lofty Ranges, potentially-weatherable
minerals mcluiding felspies, muscovite, vermiculite,
chlorite and smectite have heen identified (Bourman
19894). In some cases there may have been neo-
formation of these minerals bul itdoes seen anomalous
that they should be so widespread in areas Considered
to have been affected by lateritie weathering processes,
Previously, Crawford (1965) had noled Iresh felspir
gravel ip motded material at Ardrossanand sed this
to argue Against literitic weathering

Falaeoclimatic mdicators

Depending on the climatic conditions considered
essential for laterite formation, the presence of laterite
has palacoenvironmental implications. There is little
deubt that the operation of chemical processes ts
aceclerated under hot moist conditions bat there 4 i
growin body of eyidence suggesting that iron
mobilisation and kaolindsation can occur under various
climatic regimes - see Bourman (1993) for a summary
- so that there ave considerable uncertainties linking
Jaterite formation with a specilic climate. For example,
theye are many reports of modern iran mobility from
localities in the Mt Lotiy Ranges, Kangaroo Island and
Fisherman Bay (Ferguson er a/. 984) under curren
Mediterranean und semi-arid climatic conditions,
These observations may also cust doubt on the
rehability of correlating lerrestdal ferruginous crusts
with evidence ol warm, hum chiates derived from
marine climatic indivators (MeGowrin 19796),

Interpretation of lateritic hindscapes

Many different hypotheses have been presented to
explain the distribution and evolution of laterite. These
include the deVelopment of laterite on a surface of low
relief, clase to sca level, followed by differential
tectome uplift and dissection of thre lateritic surface,
developmentof multiple erosion surfaces affected by
episodic weathering anc Javerite formuttion, differential
weatherme und laterite formation on a landscape
formed by uplift and dissection of a surface orginally
of low relief, and the weathering, erosion and
scdimentation of a landscage before, durin: antl afer
uplift.

Lo ROBERT TP BOURMAN

Reconsiruetion of lateritised landscapes

‘The models of landscape evolution presented to
expliin the development of latente depend ou
precomceptions of haw laterite forms. For example, il
is often assumed that isolated occurrences of taterite
represent dissection of a former continuous faterite
surivce and that the differemt horizens of laterite
profiles formed contemporaneously. In the past. many
workers have tacitly assumed that the present day
isolated and sporadic occurrences of baterite represent
the erosional dissection of a former contiguous and
unilonm Jaterite-mantled planation surface and that
these remnants are excellent and reliable morpho-
sauigraphic markers (eg. Twidale 1983). However,
discontinuous distributions tay reflect only localised
formation in favourable localines (Bourn |993a)
where optimum topographic and climatic conditions
did not penerally prevail, Henec, the oveurrenee of
faler{le need hor necessarily Tndicate uv tormer extensive
erosion Surhtee,

Preservation af uplifted peneplains

Interpretations of landscape evolution have
commonly depended upan recognition of uplifted ane
dissected former peneplains, and the preservation of
parts of the onginal weathered surface, which can be
used Wy reconstruc! the former surface. A review of
the churacter and age of the sanumit high plain of the
Mt Lofly Ranges was presented by Twidale (1976),
who argued thal (he summit surface as of Mesozoic
ue and has been preserved for some 200 Ma (Twidale,
97h), Recurrent uplift of the Mr Lofty Ranges, it was
argued. postponed the ultimate degradation of the
ranges by exposing new land to the area undergoing
reduction, However, other workers have suggested that
the best preservalinn of laterile is in relatively low
lying points and least in arews of greatest uplift (Milnes
eral. 1985). The development of the Mt Lofty Ranges
on an unlicline, the Manaks of which are faulted, was
one factor used by Twidale (19769) to explain the
preservation of the Taterite-capped phiteau, {t was
maintained that the bulk of the plateau is centrally
located close to the resistant compressional zone of Ihe
anoelime and remnants near the western margin are
huttressed by sandstone and limestone outcrops,
However, the folding. which occurred in the Cambrian.
was very complex and did not result in the formation
al a simple anhcline. Moreover, erosion of tis complex
structure bas been so pronounced thal yertical and near-
vertical rock structures are exposed. Furthermore,
subsequent tensional faulting bas occurred within the
runyes (Glaessner 19534), so that the core of the anges
should not be considered to be in Compresston_ The
preservation of the Mesozoic sandy A-horizon of the
Jalgrite profile was thought iq haye assisted
pilueosurface preservation by providing an absorbent

cushion to protect the underlying lerruginous harizan
from) rainfall (Twidale 19764). However, no evidence
has been found by the present wuthor of 200 Ma old
sundy A-horizons in the ranges. Conversely, sundy soils
ure common, especially on Pertman glaevigene
seditnents and oceur in landsurfaces. demonstrably ol
posi-Mesozuic ages. A petmeable and porous
ferruginous erust of the lalerile profile was also thought
fo render this zone resistant to erosion, Howeyer,
ferruginous crusts. are relauvely rare and discantinuous
with the thickest crusts Ocedrring in positions well
below the level of the postulated ancient surtace

Gully gravure. involving the alteration of the locus
of intense erosion through the protective influence ot
vourse debris, was implied io reduce the rate of scarp
retreat (Twidale \976a), However, no specific sites
were disetissed and the present author has not observed
the extensive operation of this process in the Mt Lofty
Ranges. The unequal activity of rivers, whieh incise
more rapidly than they erode laterally. was also
suggested as a Contributury factor in summit surface
preservation (Twidale |976a), While river incision may
operune more rapidly than valley-side processes in some
situations. che operation of the processes.of weatheriny,
surface wash and pullyiny on valley slopes and bill
summits for 200 Ma has Jed to considerable
modihieation of the landscape (Milnes et al, 1985;
Bourman 199%a),

A wodel of landsvape evolution involving increasing:
relief ammpliude inorder to account for the preservalion
of these presumed ancient palaeoforms was presented.
and evidence supporting this model for other areas,
was discussed, However, itis unlikely thal the summit
surface of the Mt Lofty Ranges has survived essentially
unehunged for this enormous period of tine,

Dixsection model

The dissection model assumes ree mduralion at the
top of former complete and continuous profiles and
the Lack of preservation of complete profiles is often
taken to imply dissection (Stephens 1946: Thomson
& Horwitz 1962; Johns (96la, b; Maud 1972;
Robertson 1974; Daily er af 1974; Twidale |983),
Johns (196la) considered that much of the Lincaln
Uplands of southem Eyre Peninsula is obscured by
fossil laterite and lateritte gravels and conglomerates,
a formerly continuous mantle now partly seripped
following regwinal uplift, drainage rejuvenation and
erosion. Johns (196/b) also interpreted the accurdanee
of summit levely in the eastern Mt Lofty Ranges as
a base-levelled lerrain of Phovene age, carrying
sporadic cecurrences of terruginous grils and tatertles.
He believed that onve-coutinuous ironstone cappings
of Pliocene or post-Plioeene age have heen largely
removed by crasion. The hest exposares of ironstones
were reported from “Lucernbrae” where depustis abelut

LATERITE [8 SOUTHERN SOUTH AUSTRALIA "

Im thick were noted to mame Kanmantoo Group
metasedimentary rocks.

Mid (1972) now! thal although laterite profiles im
the southery Mr Lofty Ranges are typically thick, with
well developed ujottled and pallid zones, laterite
horizons are rare. He ateributed this to erosional
truncation of the profile, Robertson (1974) reported
ironstone fragments and deeply weathered and
kaolimised rocks in che central seetionof the Mc Lofty
Ranges at about 450 m above sea level. He alse
interpreted the weathered material as a remnant of a
Tertiary laterite profile,

Geologists and pgeomorphologists have been
particularly interested in laterite, primanily to establish
denudation chronologies. to estiblish the apes of
particular landforms, to correlate widely spaced
planation surlaces and to throw lighton the tetonic
behaviour of upland areas. Examples of the use of
latenue Weathering in interpreting landscape evolution
are provided by the work of Sprigg (1945), Brock
(19648), and Twidale & Bourne (1975a). Sprige (1946)
considered Ialerrte formation in the Mt Lotly Ranges
to be short-lived, correlated it with mottled Pleistocene
sediments and believed that faulting and uplifr of a
peneplain occurred after laterite formation, indicating
land movernents of between 180 an ane 300 in during
the Pleistocene Kosciusko epoch of block laulbing, This
interpretation provides a very young age for
lateritisation and faulting. whereas Brock (19645)
interpreted the summit surface as a peneplain formed
after prolonged subaerial weathering and erosion in
the Palaeozoic, Culnmatag in a phase of crustal
stability in the Mesozoic and Barly Tertiary, when
latentisation occurred prior toupliit and dissection of
the surlace,

Even ereater antiquity of the Mt Lofty Ranges was
proposed by ‘Twidale & Bourne (19754) who
investigated the geomorphic evolution of the eastern
Mc Lolly Ranges. A summit high plain (Tungkille
Surfage) at 200) 300 im above sea level, ain etch
surface, occasionally surmounted by scattered lateritic
residuals up to 10 tm high (Whalley Surlace), was
identified. The scattered Jateritic remnants were
interpreted as remnants of a once-contiguous weathered
surface of low reliet. The Whalley Surface and its
agsiciated deep weathering were considered is be of
Mesoroic age by extrapolation from Kangaroo Island
(Daily er af, 1974), They also argued that it developed
under a humid, tropical climate. Dislocation of the
Whalley Surface by faulting was propesed although
there was fy evidence of tured laterite on the
downthrown side of the Milendella Fault. Tts absence,
if i} ever existed, was explained by sub surilice
dissolution of the iton oxides.

Kennedy siodel of development of lmerttised surfer
Twidale (INO8) accounted for jhe absenoc of

dowataulted remnants of the lateritised erosion surface
hy proposing an alternative to the traditional
explanation of the suminit surface of the ML Lofty
Ranges that it is an extensive literitised surface of
erosion, develuped close bo regional base level in ihe
Late Tertiary, and subsequently upthrust along ancient
fault lines, after which it suffered dissection, While
conceding that the ferruginous crusts of the postulated
laterite profile might have been removed by sub-surface
solution, the possible development of an extensive
landsurface in relationship to loval base levels in the
upper reaches, one of the possibilities suggested by (he
work of Kennetly (962), was proposed, However, ou
critical evidence was presented to show that the sumoniit
surface of the Mt Lofty Ranges developed in thos
fashion

Alternatives te trancated larerite profiles

While carrying oul regional geologeeal investigators
on Yorke Peninsula. Crawford (1965) deseribed
Pleistocene deposits, exposed in the sea-cliffs. at
Ardrossan, as mottled dark red oi) olive green
wrgillaccous sediments - The ‘Ardrossan Clays and
Sandrock’ of Tepper (1879). He suggested that the
mottling could be due to latentisation, with the upper
indurated zone having heen removed by erosion and
the pallid zone Oecurring sub-surface, However, fresh
felspar gravel (i the mottled material argued against
laterilic wealbering, Consequently, an ullermative non
lateritic explanation of mottling produved by alternate
wetting and drying in an environment of low relicf was
also Suggested, Crawford (1965) obviously considered
Iuterite within the framework of the standard laterite
protile and attempted to fit his observations into it by
postulating the erosional removal of an-upper indurated
zone. He did, however, also consider an alternative
non-la#teritic explanation for his observations,

The validity of accounting for meomplete profiles
by erosional truncation in landscape interpretation wits
questioned (Bourman er ui, 1987; Bournan 1993a) by
demonstrating great lateral variability in the spatial
distribution of bleached, mottled and lerricreted zones,
the development of which depended closely on local
tmicne-environments (Bourman, 199338), Presurued
remnants of lalerile crusts have heen shown lo be lags
of ferruginous motiles waccumalating at the surface
during landscape downwasting (Bourman 1894) and
thus litterile crusts, as such, tay never have existed.

Double planation theary af Fenner

Fenner (1930, 1931) presented a double peneplanution
hypothesis to account fof the evolution of the Mt Lofty
Ranges, providing # geomorphic and (eectome
(mevewurk lor the use of subsequent authors. The
renter part of the Mi Lofty Ranges wis thought i

18 ROBERT 2 BOTIRMAN

have been stripped of an easily eroded Miocene marine
vovermass. He saw the double planation theory as
necessury lo explain transverse drainage and exhumed
surfaces in the ranges. He postulated chat a pre-
Miocene peneplain blanketed with a Miocene marine
covermass had been affected by block-fiulling. Wilting
and ditlerential uplift in the Late Mincene or Barly
Pliocene, Subsequently, this irregular surface was
thought to have been peneplined, resurrecuing the oldec
sueface ir places and developing a new peneplain on
both Precambrian and Miocene rocks. Following this,
Pieismacene (Koseiuska Epoch) teclomism renewed
erosion. Some lault blocks retnained buried by Tertiary
sediments and others, exhumed from beneuth the
covermaiss, Were subjected to renewed weathering and
erosion,

Today it is generally agreed thal the Mt Lofiy Ranges
Were not tulally immersed by the Miavene seas. so that
the double planation theary cannot be decepted in its
entirely, However, lirge reas of the Mt Lofty Ranges
and Kangaroo Island (Milnes eral, 883) were covered!
by Mincene seas ut heights in exvess of 200 m above
sea level and there és evidence in the Bremer, Myponge
und Upper Hindmarsh valleys that the shorelines were
even higher than this (Bourman 19894). Moreover,
ever hough the covermass of marine deposits may not
hove totally covered the ranges, Tertiary lerrestritl
sediments oceur eyrensively and at higher levels thar
do the murme sediments. Consequently, the double
plunation theory has considerable merit but it is still
inadequate wo account lor all Of the geomorphic
complexities of the ranges. which haye been variably
exposed lo processes of weuthering, erosion and
sedimentation for immense periods of ume,

According to Sprigg (1945) laterite in the MI Lolly
Runes formed on both a Precambrian an Cambrian
hedrock undermass and a covermass ol Tertiary
Nimestones and lacustrine sediments. Initially critical
ol the double peneplanation theory, Sprigy (945)
subsequently made the observation that the widespread
ocenrrence Of laterite over the Mt Lofty Rangcs
presented a polent argument in favour of this theory

Lanwlscipes with multiple surfaces

Landscapes with mulliple erosional surfaces have
frequently been deseribed in South Austealia, with the
suriaves beige marked by different weathering
responses. Sonte examples follow, which illustrate
varying interpretations of multicyelic landseapes.

BuLkMAN, R. P. 11973) Georurpiic evelutin ol
southeastern Fleurieu Peninsula, MA thesis. The
Univernty of Adekide (unpubl. >

Arey, NS, BE, (1960) The Camozote History of the Mid
Nort of South Australia. MA, thesis, The Lmversiiy ol
Adelaivee (uiypubh 4

Bourtman (1469). 19737). identibed a multicvelic
landscape marked hy (wo major erosion surfaces, the
Spring Mount Plateau. developed during the time from
the Mesozoic tothe Rovene and the Green Hills Surtice
of Pliocene age on Pleuneu Peninsula. The former.
underlain by a literitic weathering profile consistine
of pallicl. mottled and ferrugious-rich zones, oecurred
at ahout 400 my above sea level. The surtace was
considered to have heen tilled to the southeas|. The
second erosional surface 170-L00 m above sea level,
was capped in places by ferricrete, 4 term used ty
desenbe iron-cemerted crusts nor underlain by deep
weathering profiles. The Green Hills fernereted surface
Was thought to have formed fram reworking of laleniie
material [rum the summil surface, Lismg stranded river
enivels and civer profiles, Bourman (19737) sugyested
thal base level durjag erosion of the Green Hills Surface
in the Pliocene wats approximately 60 7 above sea level
when fluvial action modified a resurrected pre
Mineene erosion surlice,

Forrest (1969) exarhined the geomorphic evolution
of the Bremer Valley in the easlern ML Lofly Ranges
and wWentilied two erosional surfaces of low relieh
Which he cémsidered) had formed prior to a mayce
murine trinseression in the Miscene Consequently.
both the surfaces and their ussociated cappings of
lateriie material were interpreted ws pre-Miacene im
ave. The Miocene sea was presumed to five
transgressed anurea with relict similar to thal of boday
and lateritisation of the bedrock was presumed to have
followed the development of the Whalley Hill and
Lucembrie erosion surfaces prior to the Miocene,
Another surfuce, an exhumed one wilh a remnant of
derived ferricrete, and thought to have formed by
stripping of the Mincene limestone cover was
considered fu be al Phiovene age.

In a study of landsurface development in lhe Mid
North af South Australia, Alley (1969*. 1973, 1977)
identified reinnants of u laterite surface, occurring high
in the landscape but below resistant quartzite ridges,
Remnants of the laterite surface were noted Lo he most
cammon at stream-heads hut also to pecur an
prominent hills that sland newrly 100m abeve modern
valley Moors. ‘Pho luterite capping of angular quariz
jraginents set in u matrix of iron oxides was observed
lo overlie severely Weathered ind locally kaolinitic
bedrock and (o be consistently thicker ory lower slopes
Seeuons of the laterile surface were lhought \o have
been dowe-lautled inthe Ear Tertiary and laler burial
by Middle “Ternary sediments. A consistently lower
silerete-capped landsurface was considered to be
younger than the laterite surtice.

1) interpretations of mulficyelic landscapes and (he
TecoysniGoan of the ages of difterent landsurluces (here
ure inevitahly many disagreements and warkers in
South Australid have not escaped these. An oxaniple
follows, King (976) reenenised several of hes world

LATERITE IN SOUTIIERN SOUTIT AUSTRALIA 1)

wide erosional surfaces in the Mt Lofty Ranges of
South Australia, He considered that south of the
Willunya Fault, ‘laterite-enerasted tablelands’
represented the Moorlands planation (‘great Australian
denudation cycle’) of Late Cretaceous to middle
Cainozoic age, whereas north of the fault. the Mc Letty
Ranges were thought to be surmounted by bis Rolling,
landsurface of Miocene age that Jacked a true saterite,
The Widespread landscape of Pliocene ape wis
recoymised in broad valleys and basins accordant with
4 Pliocene coastal plain at about [80am nbove sea level
and the Youngest Cycle was related lo deep valleys and
gorges in the ranges,

On the other hand, Twidale (1978) considered that
Ihe summut surfaces both neh and south of the
Willunga Fault were contiguous and of the same earls
Mesozoic age, However, areas of laterite mapped by
Twaidale (978) north and west ot the Willunga Fault
onthe Eden and Clarendon Blocks are variably covered
with weathered and ferruginised Eocene ty Pliocene
sediments (Sprigg 1942, (940; Ward 1966), In places

these have been eroded to expose an underlying:

weathered pre Tertiary surface, eroded and
tewedthered since exhumation (Sprige 1945).
Consequently. the summit surface here cannot be ot
carly Mesozoic age, Furthermore, there may be some
support for King’s generalised scheme, as weathered
zones have been stripped (rom large areas of Lhe
summit sunface north of the Willanga Fault, espectally
in the eastern Mr Lofty Ranges (Twidale & Bourne
19754), allowing further erosion and the potestial
development of a younger surhice, possibly equivalent
to King’s Rolling. surface.

Age of laterite

[lis very difficull to aseribe ages to laterine materials
because they may have developed over logy Orne
periods, samc inay have several possible modes of
genesis, others. aré polygenetic having been
considerably reworked and reweathered, and there are
severe limitations on daling techniques applied to
weathered materials (Bourman 19939), Furthermore,
there ure fiely constraining sediments, These
difficulties are apparent in South Australia, where
Jaterttsation has been ascribed to periods fram the
early Mesozoic to the present, There have been many
assertions about the age of laterite in South Australia,
offen withoul presentation ot convincing evidence.
There has also heen a tendency to prescribe a single
une of lateritisaon, When evidence of the timing has
commnly been derived from limited study areas, front
where there has often been widespread extrapolation.
The following discussion of evidence presented by
different workers in South Australia highlights the great
yurtability | the ages attributed to lateritisation

Early-Middle Tertiary

Woolnough (1927) considered that faterttisarion
eccurred during one period, in the Miocene, and many
subsequent workers have generally supported the view
of a ‘Yeruary age for lateritisation (e.g. Preseort &
Peridleton 1952) but mot necessarily inthe Miocene.
Aitchison et al (1953) reported Early Tertiary
lacustrine motued sands, argillaccous sandstone and
clays occurring sub-shorontally on a pre-Tertiary
erosion surface in the Adelaide area, implying
lateritisubon in the Teruary. und Campana (1955)
favoured widespread lateritisation between the Iurly
Bovene and the Miocene.

Sections of a hiterite surface in the Mid North ot
South Austalia were thought to have been down liulled
in the Furly ‘Tertiary und later buried by Middle
Tertiary sediments, so that Alley (1973) regarded the
Jaterite surface to be of (?)Early to pre-Tertiary age
and considered that it persisted until the Middle
‘Tertiary in the Barossa area.

Lron-stained rounded quartz gratos and ferruginous
pellets were reported from within a fossiliferous marine
limestone of probable Upper Eocene age (Bourman
& Lindsay 1973). intersected in a drill hole att 36 i
underlying part of the Waitpinga Creek drainage basin,
anid aban elevation of about 60 m above sea level This
observution was. interpreted as indicauny the
deyelopaient of fateriisation, or at least ferruginisation,
prior te the Bocene,

Phocene

Many concurred with Prescott (193]) and Whitehouse
(1940) that there had been widespread laterite formation
throughout Australia in the Pliogene (e.g. Stephens
1946 Crocker 1946; Northoote 1946; Rix & Hotton
1953; Johns W6la, b). Fenner (1930, 1931) also implied
a post-Miocene or Phincene age for laterite in the Mt
Lofty Ranges.

Working on Fleuneu Peninsula, Crawford (1959)
identified laterite capping Wilson Hill at 320 m, around
which af area Was mapped as the lower part of a katerite
profile developed on Kanmantoo Groyp metisediment
tary rocks. The occurrence of areas of hard laterite at
lower elevations (100 m above sea level), on quartzose
sediments. was interpreted as indigaunme a yery
irregular original lateritised surface of Lale Tertiary
ave. Subsequently, Bourman (1973) demonstrated thot
the two decurrences were distinetive and probably of
(wo difterent ages, with Miocene limestone separating
the two types,

The evidence presented by Horwilz (1960) for mayor
lateritisation in the Pliocene is also equivocal, In the
iritramontane Upper Hindmarsh Valley of Pleuricu
Peninsula over 150 m of cross-bedded and mottled
brown ferruginous sands, capped by crust of limonite-

ROBERT P. BOURMAN

ms cemented gravels, were reported, By extrapolation
oe 8 8 8 these were considered to overlie fossiliferous Early
1 ee el Miocene limestones. The sands were thus tentatively

assigned to the Pliocene. Horwitz also considered that
by these lateritised Pliocene sands were continuous with

ws limonite-cemented gravels on the high plateau (Fig. 9).
ae z, Thus he assigned them to the same Pliocene age. Brock
E © (1964°, 1971) questioned the contemporaneity of the
Roe high-level and low-level crusts and Bourman (1969! ,
a o£ 19737) highlighted their different characters and
m % & suggested that the higher crust was of pre-Miocene age
a 2 E und the lower one of Pliocene age.

G o 2 Harris & Olliver (1964) reported on palynologicul
E 8 @ analysis of organic material preserved in “coal balls”
cS @ exposed in Tertiary sands in the Barossa Valley. The

basal Tertiary unit was described as a lJateritic sand
and gravel overlain by laminated silty and sandy clays
and was considered to be of Early Tertiary age. The
clays were capped by an upper laterite. Previously the
sands had been assigned to the Eocene (Glaessner
1955) or Pliocene (Hossfeld 1949) but Harris & Oliver
(1964) suggested that the microfloras indicated a
Miocene or possibly an Early Pliocene age for the
sediments,

Twidale (1968) concluded that the deep weathering
in the Mt Lofty Ranges oceurred late in the Tertiary
(Pliocene) and may have even conunued into the early
part of the Pleistocene. Major & Vitols (1973) suggested
that ferruginous pisolites on the western end of
Kangaroo Island were of Late Pliocene or Early
Pleistocene age as an aeolian calcarenite (Middle
Pleistovene Bridgewater Formation) was thought to
overlie pisolites, and elsewhere blocks of ferruginous
" pisolite were noted to overlie marine limestone of
probable Late Pliocene age.

—<F
aa
Fntery

Lignitifetous clays

? Pliccene sano
Woigts
= Oligo-Miacene limestones

Etech surlace

Pleistocene

~ Sprigg (1946) noted similarities between inottled
zones of laterite of the Mt Lofty Ranges and mottled
Pleistocene clays of the nearby gulf lowland and
correlated the two disparate occurrences assigning
lateritisation ta a humid pluvial period in the
Pleistocene. Bauer (19595) also favoured a Pleistocene
age for lateritisation when he addressed problems

Upper Hindmarsh Vatey

£ zs 4 associated with lateritic soils on Kangaroo Island. He
a 28 noted that the Eleanor Sand, regarded by Northcote
2 Bes (1946) as a Pliocene fossil Jateritic soil, occurs both
= ns 2 on a Pliocene plain of marine abrasion at 50 m to 100
s Se ¢ m above sea level (Mt Taylor Plain) and on the highest
~ ses
: ays]

Fig. 9 Spur-line cross section through the Spring Mount-Upper
Hindmarsh Valley-Mount Cone area showing the

- Ferruginised sands .
Ste) | and calluvium akm

J vs relationships of ferricretes to Tertiary limestone. Note
s S a a 5 2 particularly the distribution of nodular to vermiform
2 * = ay | ferricrete and that consisting of ferruginised sands, which

Horwitz (1960) regarded as of the same Pliocene age.

LATERITE IN SOUTHERN SOUTH AUSTRALIA 21

poruions of an undissected Tertiary plateau surface at
heights up to 300 m abave sev level Repardless of
whether the Eleanor Sand lornmied om two separate
periods or one, Bauer (959°) considered (hat both
were no older than the Barly Pleistocene.

Rauer (1959°) also challenged the view ol Prescott
& Pendleton (852) that laterite is an exposed podzotic
Uluvial herdon of Terbary wae, as he noted the
oceurrence of Jaterite on presumed Pleistocene
surfaces. Several anomalous laverite oecurrences were
examined by Bauer (19597). who interpreted them to
favour im sire laterite formation although Northcote:
(pers. Comm. lo Bauer) suggested tit the pronstones
had heen derived by transport and could not have
formed i yitaw because of the stuall umounty of
associated clays. However, Bauer (19599) doubted this
explamition on uge and topographic grounds.
Subsequent work (Milnes etal, 1983) revealed that the
critical limestones used by Bauer (1959°) (o date the
laterifes are older than he thought, Consequently, many
of lis objections to transported origins for the laterites
may. be remuved,

Lalentsavon hus also been ascribed to the
Pleistocene by Horwitz & Daily (958), Crawtord
(1965) and Wopfaer (1972) wha deseribed mottled
Pleistocene sediments in various locations.

Multiple periods of literitisation

A detailed Stratigraphic study by Giliessner (1953b)
and Glaessner & Wade (1958) on the western margin
of the Mt Lofty Ranges allowed them to sugpest several
periods of lateritisution. provide information on the
character and timiny of leetonic activity and ky elucidate
aspects OF lundseape evolution. Rocks and sediments
of Precambrian, Cambrian, Permian and ‘Tertiary ages
awere noted to be variably lateritised of to contauy blocks.
of lakerile, However, many of the iron oxides within
these sediments attributed to Jateritisation may have
formed since exposure, in recent times, by oxidation
of primary iron minerals such as glauconite and
sitlerite.

Several groups of workers have consiclered clrat
luteritic weathering has proceeded over lang periods
of time, For example, Campana & Wilson (1954)
attributed Jateritisation to Pliocene ta Recent
weathering, Brock (1964) fron) Mesozvic to the
Early Tertiary and Firman (1981) observed weathering
affecting matenals varying m aye from the Proterozoic
i) the Pleistocene, Milnes eval, (1985) considered that
weathering has been ongoing since the Permian,

Aller tavesiyeuons on Fleuneu Peninsala and in ihe
Mid North, Horwite (1960, 196)) considered evidence
relating (© the nature and age of laterite, which
suggested two major periods of lateritisation, in the
pre-Eocene and Pliocene, Boutmnan (979!) alse

presented evidence for Wwo ferruginous duricrusts of
diflerent apes, one on the summit surtice and the other
ow the sands that overlie Miocene Jimestoae in the
Upper Hindmarsh Valley Previously, Horwile (1960)
had regarded these crusts as contiguous and of the sume
Pliocene age Bourntan ((973’) also favodred the view
thar deep weathering procecded ufter summit surface
uphitt,

Ward (1966) believed that fhe peneplain of the
western Me Lolly Ranges is nor of the same uge
everyWhere and was ni younger than the Barly
Phocenc. Relationships between soil morphologies and
degrees of laleritisulion of mialerials were noted, as
were well-developed laterilic mottled zones. formed
beneath surfaces attributed to the Lute Pliovene. Barly
Pleistocene and Late Pleistocene. bn contrast. Maud
(1972) noted scattered erosional remnants. ol laterite
Surviving above the level of Miacene limestones
deposited in partly exhumed glacial valleys. This
Suggested that the laterite surface pre-dated the
Miocene, by which time i was being destroyed.
Consequently, Maud (1972) believed that faulting, and
Uluing Of the lateritised surface had occurred earlier
than the Pleistocene upe favoured by Sprigs, 1942). wath
the lateritised surface antedating the major perind of
diastrophism, Me equated the surtaee with the
Australian Surtace of King (962), Furthermore, Mate
(1972) interpreted outcrops of faterive afonstone at
various levels in the landscape ws retigs OF episoutic
Juteritisation, affecting alluvial sediments, inclucting:
reworked crusts, on former broad valley floors. He
suggested that the ironstone terrace remnants varied
ul age Hon Phocene for the highest to Recent for the
lowest, These valley ironstones were desembed as
locimnag parts of typical laterite profiles, with bleached,
though rarely kuolinisect, pallid: zones

Mexozpic

Daily eral. (1974) argued that evidence an Kangaroo
Ssland enabled direet and precise dating of the laterite
developed on the uplifted planate summit surface ol
the Mt Lofty Ranges. They described Kangaroo Island
us 4 dissected, tilted and block-faulled platewu wath a
caprock of laterite, in places. breached by faults.
Adjacent lowlands were noted to be essentiatly
coincident with Permian glacigene sediments Unal were
also lateritised and overlain by basalt of Jurassic uge.

The lateritie capping of the summit platesu surface
of Kangaroo [sland was described as part of a laterite
profile and they explained the lack of a complete laterite
profile in the Late Palaeozoic sediments beneath the
busalt hy erosion of the ferruginons horizons prior to
hasalt extrusion, No evidence of deep weathering on
the basalt was observed during their investiyations
Consequently, they ruled out the possibility of the
sarface on Ube basalt being an etch surhies

22 ROBERT P. BOLIRMAN

They aigued (hat as the basalt is of Midille Jurassic
age. both the laterite and fhe summit sudfiace rust be
older. ‘The hutwrite was regarded as an inelioalor oF a
hamid tropical climate and us a reliable morphostrat—
graphic marker, Using stratigraphic and palaenelinvwatie
evidence [ey suppested that the sumajt surface Was
eroded and fateriised during the Late Triassic, Barly
Jurassi¢, or both, Suppert for this consluston was
derived from evidence of warm. humid conditions
ussochited with the Triassic flora of Leh Creck. in
the Flinders Ranges, and evidence of tevtonism and
uplif) of a deeply weathered kaolinised zone during
the Mid-Jurassic. which had led to the development
ot the Polda Basin on Eyre Peninsula and the extrusion
of the Kangaroo Island basalt.

A Middle to Late Tertiary age fur the literitiscd
surluce was preferred by Northcote (979) whe
considered that the correlation of the summit surbice
weathering with that beneath the Jurassic basalt was
Unresolved,

Sclinudt e al. (1976) presented pilactmsgnetic
evidence (hal required sub-basallic weathering duriiy
a Late Oligocene ta Early Miocene period of dominant
latenue Weathering. tdi & Sentor 1978) favoured
acsynchronous Australians wide laterite remagnedsation
over this period diring a major weathering event, ‘The
superimposibon of a Mid-Tertiary weathenig event
on the earlier weathering profile was accepted by Ditily
eral, (1979) fut they also presented further evidence
for deep lateriiic pre-Eocene weathering., Milnes et
al, (1982) also pomled out problems with the sub-
basaltic weathering hypothesis. including the
preservation of a sharp basali/weathered zone contact,
the ubsence of leaching or kvolinisarion of the basal
basilt, und the fact that the busall everywhere ww lane ly
unweathered.

Subsequent isolupic dating of kaalinite (Br 196s")
and alunite (Bird eral, 19901), collected by the presen!
author tram the sub-basaliie weathered zone at
Kingsgote, together with kavlinised bedrock from the
sucinit strkiee of Kangaroo Island and Fleurieu
Permysula sigwests thal the kaolinite weathering
benvath the basalt 1s of Barly Mesozoic age, but that
the summit surface kaolinite samples are of Middle
Tertiary age. Furthermore, the alunile is not synch-
ronous with the pre-Jurassic weathering but possibly
relates to the posulated Middle Tertiary tron
Mobilisation of Schmidt er al. (1976). This illustrates
the complexities involved in some weathering matenals
and highlights potential dangers in extrapolating even
over guile short distances, and especially inter-
regiovally (e.g. Bourne 1974; Twidale & Bourne
75a, b: Twidale er al. 1976: Twidale 1983).

v Bird. M. J. (1988). AL oxygen and hydrogen isotope stil:
of Igteriles and deep weathering. Ph Di thesis, Austral rin
National University, Canberra (unpubl, }.

The many confliciitig views on the age and
development of lateriic materuils, largely arise from
investigations in wolaled localities ahd extrapalation
from them over sub .continental areas, These apparenl
vonilicis may be resulved by the application of the
ongamng wedthering hypothesis

Ongoing weathering

The evidence presented for a wide vanely of possible
ages for luteritisation and reworking of ferruginous
materials in southern South Australia. ranging
throughout the Mesazme and Cainozore including the
present (see Fig. 2 in Bourman 1993b), prompted
Bourmun (19894, 1993b) t) propose continual Jareritic
development interrupted by geological events urel
ongoing transformations of femeretes ovet long perioes
ot time. There may have been some tones wher
weutheting was More extreme but there 1s no relivble
evidence ol discrete and episndic periods pl
luleriusdoon,

Lalerite as a morphosttatigraphie marker

Durtecusts including iaterite have been widely used
US Miorphostratigeaphic markers for dating and
correfating land surfaces, in some cases of continental
exten|, Some workers, such as Twidale (9831 regard
duncrusts as execllent morphostratigraphic rourkers
and Firman (98) considered that original materials,
now ferricreted, have separate lithostratrpraphie status
und that continuous shect ferrierete hus both rock anit
soil Stratigraphic staluy, However, there are difficulties
in-using duricrusts as morphosiratigraphic markers,
For example, they may uke loag periods of time in
form so that any correlation would be extremely course.
Furthermore, as noted if the section on the ages ul
lateriles, even Short distance correlation ol apparently
similar materials cai be unreliable

Horwitz (1960) used lateritic materials in
morphostraligraphy when he associated pwazed
pisolines, pebbles and fimonite pisolites wath a pee-
Tertiary surfave on Fleurieu Peninsula. aller observ iy
similar ferruginous. materials elsewhere beneath
Tertiary limestone, However, the correlation of
ferruginous materials, which superficially uppear
Similac, may not be reliable. For example. a surface
in the lower Hindmarsh Valley, carrying ulleged Early
Tertiary pisoliths.could not have developed until post-
Miocene dimes (Bourman (19737) as it had been
covered by the Mioverie seas. Moreover, the occurrence
of pisaliths in reworked Early Tertiary sediments 1s
nota erifeal indicator of their maximum possible ape,
They may he of variable ages. or be older clasts
reincorporated inte younger sediments: In addition, it
appears that some of these pre-Eocene ferruginous
nuttenals represent the (runsgressive marine Compton

LATERITE IN SOUTHERN SOUTH AUSTRALEA y

Conglomeraie (Oligecene) of the Murray Basin
(Ludbrook 196), Lindsay & Wiliams 1977), and de
AOL relate to exposure ina terrestrial environment

A useful and innowelive approach to
Mmorphostrutipraphy was reported by Woptrer (1872)
who recorded identical mouled profiles From the Mid
North and the South East regions of South Austral,
where maphemiie motiled profiles im Camozare
sediments were capped by brown ferraginous: jer
Miaghenine crusts. Maghennte was reyarded as cat
climatic indicator and as presenting Opportunities for
correlation at the Cainoxoie sediments,

‘Twidule and co-workers (ee. Pwidale eral, 176;
Twidale & Bourne 1975a, b. Bourne 1974: Twidale
1983) have used duricrusts extensively as morphostrat-
rraplue dukers ji southern Australia, For example,
Twidale er af, (1976) desertbedt eight palaedsurfiaces
on kyre Peninsula. Among these was an epipene
surhice of Jow relied (Lineoln Surface) proteeted by
a lateritie durigrast and tormed under humid tropical
climatic conditions during the carly Mesavoig, A
younger surhice characterised by a lerrugiious
duricrust was aseribed ra the Late Tertiary. These
surkices Were used as evideree for the progressive
exposure of Inselbergs on Eyre Peninsula, The Linealn
Surlace was regardesl ay a dateate-capped dissected
pepeplain formerly contiguons with summil surkices
inthe Mt Lofty Ranges and on Kangarou Island and
disrupted by faulting.

However, the summit surface of Blue Range
described by Bourne (19744) and Twidile ef al (976)
us the Most portherly occurrence oF a postulated
Mesozoi true laterite surtice, and equivalent to
suininit surfaces on the Lincoln Uplands, Kangaroo
Island and the Mt Lofty Ranges was tought to hive
no comtinnous laterite profile beneath i (Bourmian
1k94, Neur-horizontal Precambrian melasediments
ure bleached and mottled but the mottles were
interpreted as Superficial stains of tron oxides
(Bourrnan 9894), Tabular blocks of iron-stained and
Wom impregnated sandstome litter the surface und
superticully resemble a crust but they were considered
to be reninants of Matlytug steata within the
Precambrian bedrock, Purthermore, tringing the
highest sections of Blue Range, are bleached

Precambrian metascdiments overlain by up to 2 im of

calearcous fine carth, out of syinpathy with a leached
laterite environment, and capped by a sandy grey soil
containing fragments of ferruginised sandstone bedrock
and glazed magnetic pasoliths, Thus the surface is a
comples feature, much younger thun the suggested
Mesozoje (yee

Pedopenic accumulalions of iron oxides lacking
motticd and pallid zones were reported by Twidale er
al, (1976) al lower elevations below relics of silerew
darterust assigned to the Middle Tertutry and thus were
attributed wi the Late Terthry and correlated with

similur ferricreles on Yorke Peniasula amd an the
southert Mt Lofty Ranges. The Glenville Surfiwe was
also Thappedl iq the area and was regarded as a eteh
pho cquivalept of the laterite surtice

A summary of views concerning duricrusts has been
presented by ‘Pwidule (9X3). Laterites and bauxites
were regarded as ferruginois anc dluminous menihers
of comparable origins willl siiitlar physiogmaphie add
chimatic implications, developed on contiguous land
surtices and of lhe same age ranges in piven reports,
A cmap of Australia was compiled, reaffirming the
general peripheral distributien Of laterite and an mteriot
preservation of Silcrele in arid Australia (e.g. Stephens
1971), Both promary bilecte and sulcrete were regarded
as relable strarigriphic markers, uselul in dating
fandiorms and fundseapes.

Twidale (1983) considered (hit the dating of laterite,
sulerete and ther assorted surfaces fas beer confused
by the assumption thal all rebe Gaterles are oF the same
uge und that primary and secondary literites have been
eontused, This mity have been the cause in some areds,
but in the Mt Lolty Ranges, Brock (964°), Bourmin
(969!) and Forest (969%) clearly dishayuished
faterttes formed in place and those developed hy
transport. Although some workers have stressed the
influence of geomorphic progesses affecting Miterite
development during deep weathering (eu. Alley 1973,
977), Twidale (98%) consmlered (ial (hese processes
had not been given sufficiewt appreciation, Twidale
(983) alsa thooght thal silorete developed mainly
during the Early and Middle Tertiary, Jorminy, under
warn-humid te sub-humid gonditions, but ts today
preserved im andily, ‘This ts i contrast to the views
expressed in MeGowran er al, (1978),

Twidale (983) believed that curing the Late
Mesozoic and “Tertiary mueh of Australia was base-
Jevelled and this surface of low rehef was deeply
weathered under burmid. warm conditions; laterite and
bauxite formed in the marginal areas with external
drainage, while silerete developed an uitertot
cathments. The fornjution of the duricrust was
Tnlerpreted as having been interrupted by peologic wid
geomorphic events so that the duration and timing: ol
events were not everywhere the same, Climatic
cunditions sutable for dunerusting were thought ro
have lasted for at least 60 Ma and possibly for 200
Ma and ferruginus and siliecous crusts were related
to the same extended period of warm, Humid climate
but were separtled from analowous Canmore
development by tectonie rather than by climatic events
(see alsu Alley 1977),

Kourman (1993b) noted thar the relinbility of dari
crusts (lerri¢rete) and weathered mottled and bleached
zones us murphostrapzraphie markers depends or
whether literitisalion has been ongoing: ae discrete,
episodic and related 10 periods of intense tropical
weuthering. Evidenee of continual wealhering in

24 ROBERT P BOURMAN

southern Soutdt Ausualn throughout the Mesozaic and
Connemoic favours the former view. Even Where lateriic
materials are Stratigraphically constrained there js no
evidence that they relate to humid (roped cynaligyris
or Unt their cessation depended on climatic change
rather than burial by sediments. Most commonly, dere
arc no constraining sediments und some laterite
materials have been affeeted by ongoing
transformations over long time periods.

Conclusions

‘This review of the latente litemture of southern South
Australia reveals many fundamental contliels
concerning the nature of laterite, its classiticwion, the
processes of laterite formation, the relationships of
horizons within laterite profiles, the topographic and
climatic requirements for laterite formation, the
interpretation of lateritic landscapes. the age of lateri|e
and is viability as a morphostratigraphic and
pulacoclimatic marker.

i particular, there is considerable confusion and
lack of consistency about lhe nature of materials called
laterite, these varying from superficially tron-stiined
sediments. without associated profile differentiation,
to iron-mouled and kaodlinised hedrock tornting, part
of a weathering profile. Different types of laterite
fabrics have long been recognised but unril recent work
{ee Milnes ey al, 987; Bourman ef al. 1987, Bourman
1993a) there bas been na recognition nor discussion
of their significance, Resulung trom these factors there
is nO previse Uefinition of the term by many workers.
Distinetively. different materials have often been
regarded as equivalents leading to the allocation of
spurious ages for the laterites, Qn the basis ol much
equivocal evidence, the tectonic behaviour of parts of
the Mt Lofty Ranges and apes of lateritisalion have heen.
implied.

Many studies have been merely comeidental to olher
geological investigations and others have been very
broad scale geomorphic reports that have involved
inter-regional correlations based on the use of laterite
as morphological und palaeosol-stratigraphiv markers:
(Intil quite recently there has been 4 dearth of studies
involving detailed chemical, mineralogical and micro-
morphological analyses (Milnes ef af, 1987; Bourn
eval. 1987; Bourman 1993a)

There bas been a shackling effect on landscupe
interpretaion by the model of the normal laterite profile,
which implies the original occurrence of a complete
profile including ferruginous, mottled and pallid zones
having,developed by the in site Weathering of regolith
materials, Evidence of former lateritisalion has been
attnbuted to the occurrence of weathered, bleached and
mottled bedrock as well as to ferruginous crusts, Thus,
often, the present distribution of Taleritic mutlerials on
upland areas hay been explained by the dissection of

formerly continuous latertte afer disnuption and uphitl
by faulung. 'The absence of fetruginous and/or mottled
yones fas heen explained by various degrees ol
Truncation ob af original and complete profile, rather
than considering differential development wt, and
lateral variability in. the distributian of terricretes,
mottled and bleaehed zones, depending ot local
environments of formation. Surprisingly, often only the
laterite crust has been reported missing, even though
this as likely to be the most cesistant part of the profile.
Where crusts are present they are all younger than the
unoedetely uidenying materials.

The common association of laterite development
with hunid, wopieal conditions on peneplains close
ly base level (sea level) has fed lo the development of
circular arguments relating climale, lopography and
laterite and there have been implied or specified
associating of Laverne with deep weathering by most
previous workers, whereas hypotheses offering
allermutive explanations to the view ol laterite bey
a fossil soil profile formed on peneplains under tropical
climatic conditions have fitiled to find yveneral
acceptance.

There have been suggestions of the age of faterine
formation yarying from the Mesozoic to the present.
The views have been promulgated that laterite of grew
amiquily persists in pristine form in the comemporary
regolith environment and that it is an excellent
THorphostraligeaphic marker, thereby facilitating inter
regional extrapolations. This interpretation 7s at odus
with the view that Jateritic materials are demonstrably
complex, polygencne features. having been weathered
and modified over Jong periods of ume and are
notoriously difficult (0 date. There have also been some
questionable correlations of luteritic materials berween
remote locations, based on relatively superficial
ubservations such as the shape, size and colour of iron
oxide niotiles.

As laterite formation has often been equated with
humid, torrid conditions jt has also been Used asa
Palacoclimatic indicator although some workers have
considered that current chrnaue conditions may be
suitable for its formation, There isa paucity Of dsolupic
and palacomagnetie data und other age dating
associations, such as palynology, in demonstrating the
tuning of lateritisation. Clearly there is a need for tur
more analytical work to be cartied oot in the
investigation of laterite in South Australis.

Acknowledginents

“The author would like to thenk the following peuple
for commenting on the manuseript at various times
during, its nictamorphosis — Neville Alley, Rob
Fitzpatrick, Tony Milnes, Brian MeGowran, Coli
Murray-Wallace. Malcolm Oades and Maleulm Wright

LATERITE IN SOUTHERN SOUTH AUSTRALIS 25

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CLADOCERA RECORDED FROM AUSTRALIA

BY R. J. SHTIEL* & J. A. DICKSON*

Summary

Shiel, R. J. & Dickson, J. A. (1995) Cladocera recorded from Australia. Trans. R.
Soc. S. Aust. 119(1), 29-40, 31 May, 1995.

One hundred and sixty-five taxa of Cladocera, in 53 genera, are recognized from
Australia. Seventy-two of these are endemic, with another five also recorded from
New Zealand. Species names, with published synonymy, are listed systematically,
Distributions are given by State/Territory only.

Key Words: Crustacea, Branchiopoda, Anomopoda, Ctenopoda, cladocerans,
Australia, checklist, taxonomy, distribution.

Transactions of the Koval Sever of So ates W998), LY) 29-10

CLADOCERA RECORDED FROM AU

YPRALIA

by R. J. SHIEL® & J. A. DICKSON*

Summary

Suid
MM May, 1995,

Rod. & Dieksus, LA. (95) Chulocers recorded fram Australia, rans R. Som Sd, LIC, 29-40,

One hundred und sixty-five tisd of Clidoverdt. in 33 genera. are ceoognived from Austeajia. Seventy-two ot
these ure endemic. wilh another five also recorded from New Zealand, Species names. with published sy nonyrny,
ave listed systematically, Distributions are given by Shite! Terrilory only

KEY Worps: Crusticea, Braneliipodd., Anomopoda, Clenupoda, cladoverans. Australia, checklist, taxonomy,

Jistnhubon

Introduction

The small branchiopod crustaceans commonly called
cladoceruns are vital links im aquane food webs as
Intermediite grazers between alpac and bacteria und
higher order consumers, e.g. Macroinvertebrates und
lish, They generally have been peglected in Australian
coological studies. jo part because of the lack of
suitable local laxonumie references or expertise.

The first brief deseriptions of Australian cladocerans
appeared in the expedition reports of Dana (1852,
1853). with more detailed descriptions of peculiarly
Ausiralian cladocerans by King (1853, 1854. 1866),
from the neighbourhood of Sydney, Later. G.O, Sars
(cg. ISAS. ISR. R89, 1896, 1897), working in Norway,
described specimens raised from dried mud mailed ta
him from Australia, Subsequent incidental records, ¢.z.
Henry (1919, 1922), Gurney (1927), Serventy (1929)
Brehm (953, b). Petkovski (973a, b) brought to ca
60 the taxa of cladocerans known [rom Australia.

barly records were collated. und new taxa described.
hy Smirnov & Timms (983), in the Tist revision at
the Australian Cladoceru. They listed 125 taxu, and
provided keys and figures for most of them.
Subsequently, significant taxonomic changes were
made (eg. Benzie 1988; Korovchinsky (992) and a
further 45 cladocerans were deseribed (e.g, Frey
199]a, b: Sergeev 1990u, b; Sergeev & Williams 1985,
Smirnov 89a, hb, 1992). Pive additional indizenous
chydorid taxicare deseribed in manuscripts only parily
completed by the fare D. G, Frey (indian University)
Frey was working on Gondwanan chydorids, with
ginphasts on the nadtiation of the farnly in Australia,
when he died in 1992,

Particularly as.a result of Grey's highly detuled work,
i has become apparent thal many of the cladocerans
revorded trom Australia but described (ron elsewhere,

* Murnw-Durling Breshwiler Research Centre. PO. Bow O21,
Albury, NSW 2640),

‘ire not conspecific with the nominate species. The
wide dissemination of authoritative (albeit northern
hemisphere) taxonomic references ty partly to blaine,
compounded by lack al careful discrimination, Also.
possibly as a consequence of the widespread
aeceptance ul cosmopolitaninm, some eurlier authors
did not figure their finds, but merely listed them. It
is thus impossible to determine the true identity ot an
animal if the description rs minimal, there are no
Ogures, or the original nmaterial has beeo lost.

Qur intention in proviuling a checklist of the
cladocerans recorded from Australia is to bring
together a disparate and olten indecessible titerauire.
The listing below is a starting point and includes all
the tax and references which have appeared since the
revision by Smirnov d& ‘Timms (1983). We stress thut
17S or opinidn that at least some oF the taxa named
are not conspecific with the poaminate species,
particularly those-desertbed initially from (he northern
hemisphere. Ir will clearly nike considerable effort to
resolve the systemuric uncertainties,

Systematics

We continue to use “Cladocera” because iis familiar.
although the tero) no longer has taxonomic
significanee. The chissilicuuion of the Crusticea:
Branchiopoda is oullined by Dodson & Frey (199),
Most Australian cladoceran families are placed in the
Anamopoda (families Duphnidue (5 genera/2l
species). Moinidase (2/7), Bosminidae (2/2), Ilyo-
eryptidaé (1/4), Macrothvicidae (5/24) and Chydoridae
(29/01). Only one of the two families in Ctenopoda
occurs here - Sididue (3/11), the Holopedidae do not
Similarly, only one of the three families in Onychopoda
accurs here - Podonidue (murine. 4/5), The
Polyphenndae and Cereopagidae are nar recorded here.
Neither are the Haplopoda: Leptodoridue known from
Australi This zooveographiec disparity 1s sigaibcunt

M ROI SHIELD & J A, DICKSON

ecologically; these absent tamihes contain the larger
carnivorous cladocerans,

Cladocera recarded from Australia

In the following checklist, all genera and species. of
Cludoceruns recorded front Australia are fisted
alphabetically m their respective farnilies. Original
authors are cited fully in the reference lst to fucilate
location of original descriptions, Sec also references
cited by Hawking (1994) and Shiel (1995) The first
recorded Australian locality follows the describing
author, wilh subsequent finds outside the
State/Territory of Sirst cecord also given with citing
author(s). To minimize repetitive citation, exhaustive
listings of later finds. in the same State/Territory are
not given, Published synonymies or reassignments
relevant to the Australian fauna also are given. as are
auihors of synonymy. In the followings List, # =
endemic to Australia, ** = Australia and New
Zealand. A>? with the locality record indicates thar
the taxon was listed with a “ef” and is pot positively
identified Irom that State/Territory, Unless specifically
noted. all records are from Australian inland waters,
both fresh wod athailasie saline. Families are treated
systematically in the sequence as given by Smirnoy &
Tinims (1983), However, the /yoeryplus species are
separated into the family Myveryplidae as proposed by
Smirnoy (1992). For convenience genera and species
within cach family are listed alphabetically. In the
author citations, two authors who are sometimes
confused are separated as follaws: (O,F.) Miiller, with
urmlaui, who published in the late 1770s-80s and (P.E.)
Mueller. with ve, who published in the 1840s.

Kamily Sididae Baird, 1850

Diaphdnosoma Fischer, 1850

“PD. australiensis Korovehinsky, 1981; Qud
(Korovehinsky 1981). Later finds all in Qld

DPD excisumn Sars, 1885; Old (Sars 1885); NSW (Jolly
19646): SA (Shiel er al 1982): NT (Paice 1982'):
WA (‘Timms 1988)

D. sarsi Richard, 1894; Qld (Korovehinsky [981);
NT (Tait eral. (984), WA (Timins 1988)

“LE unguiculaium Gurney, 1927; Qld (Gurney 19271;
Vic,, NSW (Walker & Hillman 1977); SA (Shiel
et al, 1982): NT (Tait 1982!) WA (Brock &
Shiel 1983); ?Papua-New Guinea (Korovehinsky
1992)

D. vulzi Stingelin, 1905, NSW (Korovehinsky 1981)

Tati, R, D. (1982) Plankton of Magela billabongs, No
M)Sc, thesis, Macquarie University, unpubl,

“Ds. BV. (1973) A comparilive study of the linmalogy
ol three maar lakes m western Victorat. Ph.D. Thesis,
Monash University, unpubl.

Latonopsix Sars, 1888
L, australis Sars, (8&8: Old (Sars (888); Vic., NSW
(Shel 1978); NT (fulli 986); WA (Timms 1988)
°L. brehmi Petkoyski, 1973; WA, NSW (Petkovshi
1973b); NT (Juli (986); Qld (Timms 1986)
Penilia Dana, 1852
P. avirostris Dana, \852; NSW (murine, coastal)
(Dakin & Colefax 1940)
Preudosida Herrick, 1884
*P susirafiensiy Smirnov & Timms. 1983: NSW
(Korovehinsky, in Smirnov & Timms 1983)
FP. szalauyt Daday 1898; Old, N'T, WA (‘Timmins 1988)
Sarsilatena Koroyehinsky, 1985
3. papuana (Duday, O01); Pseudasida papuana
Daday. 1901; Sursi/atona papuana: Korovehinsky
(1985); NT (Korovchinsky 1985); Qld, WA
(Timms 1988)

Family Podonidae Mordukhai-Boltovskoi, 1968

Pleopsis Dana, 1852
P. polyphemoides (Leuckart, 1859): E. Australia
(marine) (Dakin & Colefax 1940)
Psendevadne (Claas, 1877)
& tergestina (Claus, 1877); E, Australia (marine)
(Dakin & Colefax 1940)
Podon Lilljeborgz, 1853
P. intermedius Lilljeborg, 18532 View (estuarine)
(Neale & Bayly 1974)
Fvwidne Loven, 1836
&. spinifera Mueller, 867: E. Australia (murine)
(Dakin & Colefax 1940)
&. nordmannl Loyen. 1836, BE. Auswalia (marine)
(Dakin & Colelax 1940)

Family Chydoridae Stebbing, [802

Acroperts Baird, 1843

A. alanvides Hudendorff, 1876: NSW (Smirnow
971); Qld (Timms J988)

A. Hlarpae (Baird, 1834); Lynceus harpae Baird,
S342 deraperus herpee: Baird (1843). NSW
(Smirnov 1971)

AL neglects Liljeborg, 900; Acroperus avirestris
Henry, 1919: Smirnoy & Timms (1983); NSW
(Henry 1919)

*4_ sinvarus Henry, 919: NSW (Henry 1919)

Alon Baird. 1849

A archer’ Sars. 1888; Qld (Sars 1888)

*4_ beverlevae Smirnov, 1989; Qld (Smirnov 1989n)

A, cambiuei Guerne & Richard, 1893; NSW (Henry
1919), Vie (Shiel 1976); Qld, NT, WA (Tims
1988)

%4, clathrata Sars. |888: Qld (Sars I8&88), NSW
(Henry 1922)

A, caslate Sats, 1862; NSW (Smirnov 1971); Vie
(Vimar 19732): NT (Tait 1982!)

CLADOCERA RECORDED FROM AUSTRALIA i

A. crassicaudait Sars, (96; Qld, NT. WA (Timms
1958)

A, diaphana kang, (853: Alonella diaphana (Kings:
Sars (888); Alona david? Richard, 8S: Frey
(199la); Alona david var thering? Richard, 97
Frey (19912), Alona punctaia Daday, 198, Prey
(199la), see Prey (1991a) for comments on
synonymy; NSW (King 1853); Qld (Sars 1888):
View (Shiel 1976); SA (Shiel 19814); NT (Tait e¢
al. 1984); WA (Timms 1988)

A. ynitand Sars, \R62; Alona mricrotata Henry, 1922:
Smirnov & Timms (1983); NSW (Henry 1922);
Vic. (Timms 19732); SA (Shiel et al, 1982); NT
(Tait er al. (984): Qld. WA (Timms 1988)

A. imreticalata Shen Chia-jui, Sung Ta-hsiang &
Chen Kuo-hsiao, 64: Vie. (Morton & Bayly
1977); Tas, (Sniimey & Timms 1983)

“ad, investix Smirnoy. & Timms, 1983; Vie. (Smirnov
& ‘VYimins 1983)

“4. laevivsima Sars, IS88; Qld (Sars 188K): NSW
(Henry 1922)

“4, tlaecracontia Synimnoy & Timms, 1983; NSW
(Smirnov & Timms, 1983)

AL mondedntia Sars. 1901; NY Guth t986), Old
(Timms 1988)

A, poppe? Richard, (897; Vie, (Shiel 19813)

A, pulehella King, 1853, NSW (King 1853); Vie,
(Shiel 19814), "Qld. NT, WA (Timmins 1988)
A, quedrangularis (Miler, 1785); Lyaicens
quadraneulars Miller, 78S: Alana
quadrangularix: Smirnov (97D. Vie. WA

(Smirnov & Timyns 1983)

A, rectangula Sars, 62: Old SA (Smirnov 1971);
Vic.. (Shiel 1976). NSW (Walker & Hillann
77). NT, WA (Timms 1988)

“4. serdloides Snvirnov & Timms, 183; WA
(Smirnov & ‘Tins 1983)

A, sirtolater Sars, 1916; “tropical Australia” (Srnimov
1989a)

‘4, truncaia Smirnov, 1989; Old (Smirnov 1989)

“A, unguiculaia Smimov, 1989; Old (Smirnov |98%a)

Aleomella Sars, 1862

Al elathranda Sars, 1896; NSW (Sars 1896), SA
(Shiel J981%): NT (Julli 1986); Qld (Timms
980): WA CLinms 1988)

4 excise (Fischer, 1854): Lynceus excisus Fischer.
1854: Alonella exeiva: Sars W62b; NSW (Henry
1922), Vie. (Shicl 1976); SA (Shiel 19817); Old
(Hawkins I9KK) YWA (Bayly 1992)

‘Su, Ro (98t) Plankton of the Murray-Darling over
system, with particular reference to the zooplanktor. Ph.D.
Thesis, University of Adelaide, unpubl.

‘Mortons, 1D, W. (1973) Studies an some temporary Victorian
witlers with special reference 10 the Microcrustaven, B.Sc,
(Hons) Thesis, Monasl University, unpubl.

A. exfeva (Lilljehorg, [853); Lyneeny exiguns
Lilkjeborg, 1853; Alonella exigua: Mueller, 1867;
NSW (Smirnov I97l): NT (Tait er al. 1984)

tArchepleuroxus Smirnov & Timms, 1983
“4. beryl Smirnov & Tintms, 983; Vic., Tas.. WA
(Sturmoy & Timms 1983)
Adustralochvdorus Smirnov & Tininis, 83
44. gporus Smirnov & Timms, (983; Old, NSW
(Smirnov & Timms (983), NT (Tit er al, 1984);
WA (Timms 1988)
Biapertura Smimov, 1971
*B, abbreviate (Sars, 1896), Alona abreviata (sc)
Sars, 1896: Biapernira abreviare (sic): Smirnov &
Timms 1983, NSW (Sars 1896)
Comment! The spelling of the spectes name with d
sige b as abreyiata in the original description (Sars
1896; 40) appears to be 4 typographical error, as 1 is
later spelt (p.43 text: p. 79 fig. caption) as abbreviara.

B. affinis (Leydig, 1860), Lynceus affinis Leydig.
1860: Alona whirelevgii Sars, 1896: Henry 1922.
Alona affinis: Sars 1901, Alona longirasirts Henry,
1919: Smirnov 1971; Biapertira affinis: Smirnov
& Timms 1983; NSW (Surs 1896), Vic. (Timms
19732), WA (Williamns 1979); SA (Shiti 1981");
NT (Tail 1982'); Old (Tinuns (986)

*B. duoodonta (Heary, 1922); Alonella ducodonra
Henry, 1922; Biapertura duoddoata: Smurnoy &
Timms 1983; NSW (Henry 1922) INT (Tait ep
al, 1984)

*R imitateria Smirnov, 1989: WA (Smirnoy [98&9a)

B. intermedia (Sars, \462), Alone intermedi Sars.
862: Biapertura intermedia: Smirnov 971, Qld
(Gurney 1927); NSW (Smirnov 1971); Vic, (Shiet
1978); WA (Bayly 1962): NT (Timms 1938)

B. karwa (King, 1853); Alona karua King, (853:
Alonella kara. Sats, W888. Bapertara karna:
Smirnov & Timms 1983; NSW (King 1853); Old
(Sars, I88%); Wie. (Morton 19734), NT (Tait
1982'); WA (Timms 1988)

Comment: King’s description is inadequate hy modern

shindards. There are differences in the post-abdomen

morphology of his species and that later hatched frean
Qld mud by Sars (1888), although Sars considered the
taxa identical, There ms now good evidence that &
karua represents 2 species complex worldwide (Alonso
& Pretus 1989). In our opinion the O00) km separation
of the King and Sars taxa is sufficient to doubt
conspecificity, hence their respective identities are not
satisiactorily resolved at this time. The problent is
compounded by eyrors in Smirnov & Timms (1983)
(see Incertae sediy below).

*B. kendallensis (Henry, 1919); Alona keridallensis
Henry, 1919; Bianertira kendallensis! Smirnov
1971; NSW (Henry 1919), Qld (Smirnov 1971):
Vie. (Timms 19732), NT (ulli 1986); WA
(Grows ef al, W925

a2 RJ SMIEL & 1A

“R. lonsiqua Smirnov, W871, NSW, Old (Smirnov
97h; Vie. (Shiel 19815). WA (Timms 1988)

"B. macrecaper (Sirs, O94): Alona macrocoper Sars,
1894; Bianerrera macrecopa: Smirnov & Tinos
1983 (author dale given us 1895); Qld (Gurney
1927); Vic. (Morton & Bayly 1977); WA (Bayly
1982); NSW (Timms 1982)

*B. rivuticandix Smirnov, OT, Alena intermedia
Gumey 1927 (misidenuification): Biaperura
rividicaudis, Smirnov 1971: Qld (Gurney 1927):
Vie (Shiel (976); NSW (Timms. 976); SA (Shiel
1981), WA (Bayly 1982), NT (ull 1986)

"Bo rusticaidey Smirnoy & Timms, 1983; Tas,
(Smuirnoy & Timins 1983)

RB. setigera (Brehm, 1941); Alona vattani seligore
Brehin, 1931, Alana setizeni: Petkovski |97Au:
Biaperiura setigera. Smarnoy & Timms 983;
NSW (Bayly 1970): Vie, (Shiel 1976), SA (Shiel
ORE): Old (Timms 1986) ?WA (Storey er al
1993)

B. verracoase (Saes, WO, Alona verrucosa Sars.
901: Alone reclanwule pulehre Hellich. 1874
Smiron IW7l and Smirnov & Tirnms 1983;
Biupernira verrnceusas Smupnoy 9895 Old
(Smirnov 197: NT Gull 986): WA (Timms
1988)

*B. willisi Smumdy, (989; Old (Smirnov 1989)

Camptacercus Baird, 1843

*#C> australis Sars, 1896, NSW (Sirs 1896), Vic.
(Shephard eral '918): Qld (Smirnov 17> NT
(Tuit 1942"); WA (Timmins 1988)

*Celsinonine Frey. 199]

*C. Aypsilopham Frey, 1991, NSW (Frey 19910)

°C. putraneusis Brey. 9) NSW (Frey 19914)

*C. plataumentes Frey, 991; NSW (Frey, 1991)

Chydorus Leach, |16

CC. eurvantus Sars. Ol: Qld (Timms 67); Vie
(Walker & Hillraan 1977)

C. herrmanni Bret, 1933; Qld (Timms 1967); Vie.
(Shicl 19814)

C. kallipyeos Brehm, 1933, NSW (Petkowskr 19738):
Old (Hani (975%)

Comment. Smienoy & Tints (983) regarded Perkoy -
skhS record as a mistdentification of C. enryneuty,
However Hann (1975°) independently recorded C. kal
fipvges from NSW and Qld. Petkovskrs record should
stand unlil.a thorough revision of the genus is mude-

“CL ohscuiirosteis Brey, RT, NTL WA (Prey 1987)

*C. opachy Frey, W87: NT. Qld, WA (Frey, 1987)

©. parvuy Daday, (898; “tropical Australia” (Smirnoy
O89)

C. rencufarns Daday, WWY8; “tropical Australia”
(Smirnoy lk9a)

“lass. BLD. 1975) Taxonomy ot Chydoridae m Ontara and
genus Chvderis worldwide MS Thesis, Unversity af
Watertoo Ontario, ripubl.

DICKSON

Comment: This specres was listed without comment
by Smirnov (989), Ibis given as a synonym of ©
sphaericuy (Miller) in Flassner (1972), The relitton-
ship of this taxon to the other ‘feavifarnislike relicu-
jated taxa deseribed by Prey (1987) remains unresolved.
We consider it unlikely to be Daday’s species.

C. sphaeriens (Miller, 1783), Lynceny sptheertony
Muller, 1785; Chyvdorius sphaeriens; Baird 1843,
Chydarus cleland; Henry, 1919 was synonynnzed
with Chydorus leonard? by Henry (i922),
Chydarus leonard King, 1853: Smirnoy (1971) C,
leonard’ was attributed to Sats. [896 by Smirnov
& Timms (1983); NSW (King 1853); Vie. (Morton
1967), Qld (Tims 1967): NT (Tait 9829) SA
(Shiel e7 a, 82), "WA (Bayly 1992)

Comment: In view of the restricted distribution of
Chydorus sphaericus s.sue Brey 1980), it rs likely that
a complex of species accurs in Australia, none of whieh
is Ihe nominate taxon (D.G. Frey pers, comm.)
Dadava Sars, 1901

Do meacrops (Daday, 1898): Alans macropy Daduy,
1898; Daedeya macrops: Sars OL, Qld (Srainey
OTD, NT (Tait-ez al 1984); WA (Timms 1988)

Dispuralina Pryer, 65
D. weurvesrris (Birge. IN79); Plenroxus acalirostriy
Birge, IK79. Akmelle acutirastris; Prey W59
Disparalona acutirestris, Fryer J971, “tropwal
Australia” (Stairnov 1989a)

Duntievedia King, 1853
D, erasyu King. 1853: NSW (King 1833); Qld (Sars
la88); SA (Henry 1922); Tas. (Brehm IWS3a),
Vic. (Marton 19734), NT (Tait eral, 1984): WA
(Timms [9&8)

Ephemeroparus Frey. (982
2. tridentatus (Bergamin, 193912 Chydonets
tridemarus Bergamin, 1939: Chydorny barreiy
Richard, (894, Fig, 329 jn Smirnov [97],
Ephemeroporus trideniatus; Frey W8la; Qld
(Smmienoy 1971): Vie. (Shicl 19814): NT (Tait ef
al, 198A. WA (Timms 1988)
Comment: see /neertue sediy tor other taxa referred
to this genus in Australia.

Eurvalona Sars, (901
F arientalis (Daday, (898); Alonapsis avientelis
Daday, I8Y8. Evryalona acctdenralis Sars, OO:
Smirnov I971; Auryelena erientalrs: Daday 05,
Old (Sryirnew de Timms 19K3)) NT (Tait eral
(984); WA (Timms 1988)

Grapteleherts Sars, (62
GG lesindinaria Crhiseher, W48)2 Lincens
texindinariuy Fischer, [848 (cited as IRSL in
Smirnw & Timms |[1983]); Graploleberis
lestucinarie, Kure. 1874; NSW (Henry 1919): Vie
Shuel W76) NT (Tait 19824. Old WA Timmins
aS)

CLADOCERA RECORDED FROM AUSTRALIA 4

Kura Dybowski & Grochowski, 1894

A. latissime (Kurz, 1874), dlonopsis latissima Kurz,
1874; Kursia latixste Dybowski-& Grochowski
R94, Vic. (Shiel 1976)

RK. lungirostris (Daduy, 1898); Alona longirostrix
Daday, 1898; Kursia longirostrix: Harding 1957;
NSW (Timms 1972), NT (Tail ef al. 1984), Old,
WA (Timms 1988)

*Leberis Smirnov, 1989
“L. aenigmataya Survey, 1989; WA (Smirnov

hb)

Leydigia Kurz, 1874
L. acanthocercoides (Fischer, 1854), Lynteciey

acaunthaeercoides Fischer, 1854; Levydleia
acanthovercuides, Kurz, 174. NSW (Tins
1970), NT (ull 1986); Qla, WA (Timms 1988)

{., australis Sars, BRS; Qld (Sars 1885); NSW (Shiel
1978). View, SA (Shicl 19813)

L. ciliata Gauthier, 1939; NSW, Qld (Smirnow 1971);
Vie, (Shiel 19814)

*L. laevis Gurney, 1927; Qld (Gurney 1927): NSW
(Snel 1981°): WA (Growns er ai. 992)

L. fleydigi (Schoedler, 1863): Alena levdivit
Schoedler, 1863; Leydivia leyiligi: Daday 1902;
SA (Henry, 1922), Vie. (Shiel 1976); NT (Tait
19821), WA (Growns ¢f al. 1992), NSW
(Kobayashi 1992)

*Monope Smirnov & Timms, (983; Monoports
Smirnoy. 1977: Smitnoy & Timms (1983: 34)
*M. reticulata (Henry, 1922); Pleureaxus reticularis:

Henry 1922: Monoparcs henrvyae Smirnov, 1977;
Monope reticulata: Smirnov & Timms 1983; non
Plenroxus reticulatus. Henry, 1918; Frey 199 1b:
NSW (Henry 1922): WA (Bayly 1992)

Coynment: Henry's (1918) taxon as figured is, according

(o Frey (1991b). a species of Alonella, probably 4,

cluthraruld Sars, 1896.

Monospilus. Sars, 1862
“M, diporns Stainoy S ‘Turns, 983, SA (Shiel

1978) (as Monospilus sp. nov.), WA (Brock &
Shiel 1983) (as Monaspilis sp); NSW (Shiel
1981') (as Monaypilas no. sp. Dy Vie, (Shiel &
Croome, unpub), data)

*M. vlongains Smirnov & Timms, 1983; SA (Shiel
Wsl3) (as Monespiluy n. sp. 2)

Comment: Neither of these taxa is referrable te

Monaspilas s. str; indeed they are probably sor even

vongeneric (DG, Frey, pers. camm.).

Noralana Rajapaksa, 1986
N. vlobulasa (Daday, 1898); dlona globulosa Daduy,

1898: Notealona glebulosa: Rajapaksa &
Fernando 1987. The nominate species iy not
recorded from Australia, However a geographic
subspecies ts kruwn:

CW. globuloxa australiensis (Rajapaksa & Fernand,
1987); Indiqlona (Petkoveki, 966); Rajapaksa a
Fetnando IS&T) NT (Smurnoy & Timms 95 as

Indialona sp.); (ld (Rajapaksa & Fernando 1987;

WA (Timms 98%)
Comment: Indialona was reported {rom the NT by
Smirnov & Timnis (1983), species not piven. Julli
(L986) reported J. globulosa. also from the N.T As
Rajapaksa & Fernando obtained their material frorn
BY. Timms, who also collected the Smirnov & Timms
material and identified the Julli material, it ts probable
that all NT. records are N. globules austrelienscs.

Oxyurella Dybowski & Grochowski. 1804

Q, singalensis (Daday, 1898); dlonepsis singalensis
Daday, 1898; Oxyvurella singalensis: Smurnoy
1971, Old (Smirnov & Timms 1983). NT (Tolli
1986); WA (‘Timms 1988)

OL tertuiceicdis (Sars, \862); Alona tentuicatdis Sars,
1862) Alona wallaciana Heaty, 919, Oxyurelle
wallaciane: Sinimoy WT); Oxyurella tenuicauelis:
Smirnoy & Timms i983; NSW (Henry 1919):
Vic. (Timms 1973?) (as Oxyurella sp.

*Planicirelas Frey. 1991
*P) alticarinatus Prey, 991; WA (Prey 199\b)

*Plurisping Frey, 199]
*P. chuulivdiy Frey. 991; WA (Brey 1991b)
#2, multituberculata Frey, 991. WA (Prey 1991b)

Pleuroxus Baird, 1843

*P foveatuy Frey. 1991: WA (Prey 1991)

*P inermis Sars, 1896; Chydonis denticularus Henry,
1919; Frey (1991b); NSW (Sars: 1896). Vic. (Haase
1903); Qld (Gurney 1927): SA (Shiel 1981;
WA (Bayly 1992)

*P jugosuy (Henry, 1922); Chydorny jugosus Henry.
1922: Smirnov & Timms (1983); NSW (Henry
1922)

Comment; Frey (J991h) states thal close study of the
type specimen did not reveal cnough positive characters
to towke 4 firm decision (regarding Pleureauy cf.
jugosus), and hence this taxon, al least forthe present,
must be regarded as a nomen dubiumn. Pleuroxus
Jugesus io Smirnov & Timms (1983) ws not Chyderus
Jugoses Henry, 1922: rather most of the description
und all of the illustrations in this paper are lor
Plarispina chauliodis Frey, 1991.

*P. kakadidensis Smimay, QR NT (Smirmov 1989b)
P laevis Sars, 1862; NT, Qld, WA (Timms 198%)
PF similis Vavea, O00: Po ausralis Henry, 1922:
Smimoy & Thrinis 1983: NSW (Henry 1922)
Comment: Probably absent from Australia (Frey !991b)
*P mocellarus Smunnov, 989. WA (Sutirtioy 1989b)

Pseudochiydarus Fryer, 6%

P. globosus (Raird, 1844); Chydoruy globosus Baird,
1843; °Chvdoerns aveustis King, 1853: Sars
(888); Pseqdoclivdarus elobosus: Fryer 1968;
NSW (King (854): Vie. (Shephard er af. 1918);
Qid (Timms & Midgley 1969); SA (Smimoy
71, NT, WA (Timms (988)

Ja R, J. SHIRL & J A. DICKSON

Rak Smirnoy & Timms, 1983
+R. labrasus Smimov & Timms, 1983: SA, Tus. Vic
WA. (Sotirnoy & Timms (983)
*R. obtusus Smirnov & Timms. 1983, NSW, WA
(Smirnoy & Thrans 1983); Qld (Timms 1948)
Comment; Several new species of Kak from W.A, are
included in an incomplete MS by the late D.G_ Frey.
He also found Rak in South Africa
The Rak MS will be completed by RIS.
*Khynchochyderus Smirnoy & Tirniis, 983
*R, australiensis, Smirnov & Timms, 19823,
Amblyaryachis: Bayly 1992 (nemen nude),
NSW (Smirtiov & Timms 1983); WA (Bayly 1992)
eSayera Sars, 04
“5. cnoki (King, 1866); Euryeereus cookl King,
1866; Saycia erbicularty Sars, 904: Smirnoy
1966; Suyela coekt: Smirnoy 966: NSW. Qld
(King 1866); Vie. (Sars 1904)
Comment: Alter examining a N.Z. population, Frey
(1971) coneluded that it represented a new geographic
subspecies, Saveia cook? novaezealandiue Prey, 1971,
The Australian subspecies is designated Saycla ceokr
cookt (King, 1866): Smirnov & Timo 1983

Family Tlyocryptidae Smirnayv, 1992

Hyveryptus Sars. \862

1. hrevidentainy Ekman, 905; Vie. (Shiel 19813),
NT (Tait et al. J984)

“1 raridemaris Smirnoy, 1989; WA (Smirnov !989b)

1, sordidus (Lievin, WAR), deanhocereus sondicus
Lievin. I848: Mywerypiis serdidus: Sars. (896;
NSW (Sars 1896): Vie. (Henry 1922): SA (Shiel
OBIS). NT (Tart et uf, 1984)

1. spinifer Herrick, 1882) 1 dongtremis Sars, 1888"
Smirnoy & Timing (983), 7) ahi Brady, (886
in Gurney (1927); Smirnov & Timms (1983): Old
(Sars, 1888): Vie. [Timms 1973): WA (Walliams
1979), SA, NSW (Shiel 1981); NT (full 1986)

Farnily Mactrothri¢idae Baird, 1843

Grimaldina Richard. 1892
G. braszat Richard, 1892: Qld. WT (Timms (988)
Macrethrix Baird, 1843; Eehinisca Lievin, 48:
Smirnov 1992

tM. breviseia Sinirnov, 1976; Qld (Smirnov 1976);
Vie. (Shiel 19814); WA (Growns. er ai. 1992);
INSW (Timms 1993)

M. capensis (Sars, 1916); Behinisea capenstry Sars.
16; Smimov 1992; Vic (Smimney 1974). NSW
(Shiel 19812), Tas., WA (Smirnov & Timms
W&3). Old. NT (Timms 1988)

SMITCHELL, B.D. 980) The coology of waste stubrliziation
ponds. Phy, D. thesis, Liniversiry of Adelaide, unpubl

~M. carinata (Smirtiov, 1976); Echinisea carineara
Smirnoy, 1976> Smirnov (1992), Qld (Smirnoy
1976), NSW, Tas., WA (Smirnov & Tintms 1983)

*M. flabelligera Smirnov. 1992; Qld (Smirnov W992)

+M_ flagellate (Smirnov & Timms. 1983): Eehinisea
Hagellata Sinirnoy &¢ Timms, 1985; Sininev
1992; Tas: (Smirnov & Timms, 1983)

4M. harding? Petkovski. 1973: Echinisea hardingy-
Smirnov 1976; WA (Petkoyskt 19736); NSW (Shiel
19814)

M. hirsuticorniy Nortivan & Brady, 1867) Vie
(Smirnow 1976); SA (Milehell 19804)

4M hystrix Gumey, 1927: Old (Gumey 1927): NT
(Juli 1986)

+M. indisrinceta Smirmev, 1992, NSW, WA (Smirnov
1992)

*M. longixeta Smirnoy, 1976; Vic. (Smimov 1976);
“tropreal Australia” “Tas, (Smirnoy 1992)

M. mafavensis Idris & Fernando, 1981, Old (Tinwes,
OBR)

+M. pectinata (Smirnov. 1976); Eehinisca pectin
Smirnov: 1976; Smirnoyw 1992; Old, Vie, (Smirnoy
1976); NSW (Smirnoy & Timms 1983)

M. rosea (Laevin, IB48); Echinisea rosea Lacwit.
i848: Smirnov 1992; Old (Smirnov & Timms
1983)

M. schaunislandi Sars, 19035, Macrothriv bursialix
Smith. 1909: Smirnoy & Timms (I9B3)2 Tits
(Smith 1909), Vic. “tropical Australia” (Somrnoy
(992)

M. spinesa Ring, 1$S3; NSW (King, 1853); Vie,
SA (Shiel l981*)

AM. rimnist (Smirnov, 1976); Behinisea tinmnsi
Sournoy, 1976: Smirnay 1992; NSW (Sunrney
1976); Qld (Tinwns 1986)

M_ trisertalis Brady, 1886, Fohinisea triserialis
Smirnov 1976; ?NSW (Henry 1922), Old, Vic,,
SA (Smirnoy & Timms, 1983); NT Chulli 1986);
WA (‘Tinims 1988)

*M. williami (Smirnov & Timms, 1983); Behinise
williams: Smirnov & Timms. 1983) Snvirnov
1992; Old (Smirnov & Timms. 1983); NT (oll
1986)

Neothrix Gurney, 1927

WEN, armata Gurney, 1927; Old (Gurney 1997); Vie.
(Morton 19734); WA (Bayly (982); NSW
(Kobayashi 1992)

+N paucisetosa Smirnov, W89hy Marrorhpiy
paucisetosa Smirnov Y89b: WA (Smirnov 1289b)

+N. superarmat Sinimov, 1989b, Qld (Sinimoy
(989b)

“*’Pyendemoina Sars, 1912
+P lemnad’ (King, 1853), Moina lemnae Katy.
1853; Psevdomoina lemneae Sars 19125 NSW
(King 1853); Vie. (Shephard er al, 918); Tas.
Smirnov & Tins. 1983); SA (Shiel & Koste
1985)

CLADOCERA RECORDED FROM AUSTRALIA 34

Streblocerus Sars, (62
S. serricanmedatuy (Pischer 1849); Daphnia laticarnis
Fischer, 1849: Srreblecerus Yerricaudurus
Lilljeborg, 1900: Smirnov (976; Vie. (Smirnov
1976); Qld, Tas. (Smirnoy & Timms 1983)

Family Moinidae Goulden, 1968

Moina Baird, 1850

“KML anstraliensis Sars. (896: NSW (Sars 1846);
Vic. (Shiel W814); WA (Smirnov & Timis
1983), NT (Tait eral. J984)

*M. baylyt Porro, 18S; Moina mongelica Daday,
9QL in Bayly (1976), Smirnoy (1976). Smirnov &
‘Yinuns (984) (misidenritied): SA (Bayly 1976);
NSW (Williants 1986); Qld (Timms 1987)

“M1 fletuosa Sars, 1897, WA (Sars 1897)

M. micrura Kury, 174: Moina propingua Sars,
1845: Goulden (1968); Moina dubia Richard in
Gurney (1927) (misidentified): Goulden 1965);
Qlu (Sars 1885): NSW (Timms 1970): Vic.
(Timms 1973°): SA (Shiel 1978); NT (Smirnov
& Timms 1983)

“M. lenuicorniy Sars, 1896, NSW (Sars. 896); Vic.
(Henry 1922).

Cominent. Possthly ulsa from South Africa
(Unverified); Goulden (1968)
Moinodaphnia Herrick, 1887

M inacleayi (King, 1853); Moina macleayi King,
1853; Moinodaphnia macteayr: Sars 1888; NSW
(King 1853); Qld (Smirnov & 'Timms 1983); NT
(lull) 1986); WA [Timms 1988])

Family Bosminidae Sars, 1865

Bosmina Baird, 1845

B. meridionaliy Sars, 903 (not 1904 as in Smimoy
& Timms [1983]).

For extensive synonymy. see Smirnoy & Tinuns
(1983). See also Jncertae sediy below; Tas. (Smith
1909, as 8B rotundant); NSW (Jolly 1964); Qld
(Timms & Midgley 1969); Vie, (Timms (973%);
SA (Shiel et al. 1982); NT (Tait ev al, 1984); WA
(Timuns 1988)

Bosnunapsix Richard, 1895

B. dietersi Richard, 1897, NSW Uolly 1966); NT

(‘Tait 1981); Old (Timms 1986); WA (Timms 198%)

Family Daphniidae Straus, 1820

Ceriadaphnia Dana, 1852
C. coriuta Sars, 1885; Qld (Sars 1885); NSW
(Henry 1922), Vic. (Shie! 1978); NT (Tait 1981);
SA (Shiel ef.a/. 1982); Tas. (Koste & Shiel 1987):
‘A (Berner 1987)
Comment: Evidently more than one small species of
Ceriodaphnia with ai acute “beak” occurs in tropical

Australia (cl Berner 1987), Unt a thorough revision
of the genus has been made, these laxa should be
referred w C. vortur 5.1

C. dubia Richard, 1894: Qld (Gurney 1927): Vie.
(Shicl 1976); °NSW (Timms 1989)

C. latieaudauis Mueller, 1867, ‘Vic, (Shiel 1978);
2QId (Timms. $988)

C. quadrangila (Mover. 785; Cerindaphnia hakea
Smith, 1909; Brehm (1953a)2 ?Ceriodaphnia
planifrons Smith) Brehm (1953a); Tas. (Smith
1909); NSW (Jolly 1966): Vic. (Timms. (973+);
SA (Shiel 1978)

C. rotunda Sars. 1862, Vie (Shephard er al, 1918)

Daphnia Miller. 1785

D. carinara King, 1853.5,/. For extensive synonymy.
see Benzieé (J988: 136-139), NSW (King 1853):
Vic. (Shephard 1898); Tes, (Shephard 1917), WA
(Serventy 1929); Qld (Tinims 1968), SA (Mitchell
1978): NT (‘Timms & Morton 1988)

DP, vephalata King, 1853; For synonymy, see Benzie
(98S: 129); NSW (King 1853); Vic. (Sars 1914)

*D, jolly’ Petkayski, 1973: WA (Petkavski 1973)

D. lumhaltsi Sars, 1885: For synonymy, see Benzic
(ORK: 113-14): Old (Surs 1885); NSW. Vic. SA
(Shiel 19814); WA (‘Timms & Morton 1988)

*2 nivalis Hebert, 1978; For synonymy, see Benzte
(1988: 122); NSW (Hebert 1977)

*D. aceidentalis Benzie, 1986; WA (Benzie &ha)

Daphniopsis Sars, 1903

*P australis Sergeey & Williams. |S; Tas,
(Sergeev & Williams 1985); SA, Vic. (Williams
1986)

*D. pusilla Serventy. 1929; WA (Serventy 1929):
Vic., SA (Bayly & Edward 1969), Tas, (Sergeev
& Williams 1983)

*D. quadranguluy Sergeev, 1990: Vic. (Sergeev
19904)

*D. queenslandensis Sergeev. \990; Qld (Sergeev
1990h)

Scapholeberis Schoedlen 1858

S. kingi Sars, 1903; Daplmia mucronata Miller,
1785: King 1853 (misidentification);
Scapholeberis kingit: Sars, 1388 (namen nudem):
Smirnay & Timms 1983; Scapholeberts kingt
Sars, 1903; NSW (King 1853); Vic. (Henry 1922);
NT (Sulli 1986); Qld (Timms 1988); WA (Halse
el al. 1993)

Simecephalus Schoedler, 1858

5. acutirasrramus (King, 1853); Daphnia elisabethae
acutirastrata King, 1853; Simecephatus
acutirostratus: Sars IRR; Stmoecephalus
dilvertonensts Smith, 1909; Dumont in Smirnov
& Tiinms 1983; NSW (King 1853); Vie, (Haase
1903}; Tas. (Smith 1909); NT (Tait et ad. 884);
Qld (Timms 198%)

46 RO oT SHIEL & J A, DICKSON

S exypunesus oustradienyis (Dana. (852); Daphnia

anstrativeasis Dana, (52. Simwvephalas
ausrraliensis! Sars, WRB, SS. exspelnasis

austratiensis; Durnont in Snyrmov & Tipps
(1983); Qld (Sars 1888), NSW (Surs. 1896); ?Tas,
(Stith 1909) SA (Henry 1922): WA (Servenry
1929): Vie. (Morton 19734)

S. Jatirastris Stingelin, 1906; ?S. theringi Richard,
1897; Dumont in Smirnov & Timms (984):
NSW (Henry 1922), NT (Tait ef af 1984); Olid.
WA (Tins 1988)

S. serrulatus (Roch, Wal, NT ull x6): Qld
(Timms (88s)

Si vetulies (Miller, 1776), 2 ssp. recognized Fray
Australia (see copiments by Dumont, in Smirnoy
& Timms [I9834> 98-102])

S verlus elisabethue Using. B53), Daplinie
elisahetliae King; Simmacephalus elixabethae. Sars
(SSS; NSW (King 1853); Vic. (Shiel 197K) SA
(Shiel 9815), Qld, NT (Tiims Tks)

S. vetdlus gibbosus (Sars, 1896); Sirecephealis
gihbosus Sars. 1896; Simecephalus ventas
glhbusus! Dumont in Smirnov & Tinims 1983:
NSW (Sars i896). Vic, (Shephard era/, 1918),
SA (Shiel 1981+)

*S. vicroriensis Dumont, 983: Vie (Dumont in
Sroirooy & ‘Viontas 84)

Ineertiae sedis

Alona hairdi King, 1853, NSW (King
Desenpuon inadequate.

Alona karte King, 1853; NSW: Biapertura karva in
Srirnoy & Tires (983) is erroneously referred to
King, They list King’s dlona kare as incertae sedis.
The species they have mistabelled is dlonella karua
in Sars (1888), which is dpparently a
misidentification of another species, not the
nominate A, keruva, As tygured by Sars, itis clearly
nol the taxon figured by King, und should be
relocaled if King’s species is rediscovered.

Alona mascufa King, (853. NSW: Inadequately
deseribed,

Rosina maritima Mueller, 1867; ~ off the Abrolhos,
300 miles north of Fremantle, Western Australia,
nt November, 1910" (Searle 1936; 172). Not recorded
again. or mentioned by Korinck in Smirnov &
‘Yirnms (1983). 8 maritimes is recorded as a synonym
ol B longixping Leydig, 1860 in Plossner (1972)
This is the only record of a murine bosminid from
Australia and its identity is uncertain.

Chydorus barroist (Richard, 1894): Qld (Smimov
OT): = Ephemereporus burroisi, nomen dubium
(See Frey 1982a), Frey noted (p, 234) that the
figured specimens from Prospect Reservoir, NSW,
in Srairnoy (1971 Figs 328, 330, 331, 332) are nat

ISSA:

conspecific with &. rrideemaiis (Fig. 329 in the same
series). nor are they conspeciic with B harraist
5, Str.

Chydonis livbridus Daday, 1905; Old, NSW (Sntnov
& Timms 1983): Frey (92a) relocated ©, hybrids
5. str. tow new genus, Evhemeroporny, and the xen
hecame &. dvbviday (Daday). The limited features
of the Australian taxon assigned to °C. Ayhridus® as
figured in Smirnov & Timms 983) are neither
Chydorus nor Ephemeruporus, bul more correctly
those of Rak (Frey, in MS).

Clivdoras ovals Kurz, 1874: NSW (Henry 1922): No
fipures or material are uvailuble of the tuxon
identified by Henry and it has not been recorded
again, Itis a Holarctic species and regarded as absent
Irom Australia by Smirnov & Timms (1983).

Daphnia honerata King, 153; NSW (King 1833): A
speeles of Ceriadaphina, insdequately described anu
figured. Sars (I888) vonsidered if close to the
European CC. reffenlata (Jurine, 1820), but
specifically distinct

Dunhevedia podaera King, 1853; NSW: Not seen sinee
Original deseription, which is inadequate.

Euryeercus chnninghami King, t853: NSW (King,
1854). A chydorid, but not referable to Aunecercay,

Auryeercns spinosus King, 1853, NSW (Ring. 1853)
A chydorid, buat not relerable to Banwerciy.

Plearoxas aduncus (urine, 1820) in Smimoy & Tins
(1983), Aloriella nasuta Smith, W909) Chydorus
denticulaius Henry, 1919, Chyderus anispinus
Henry, 1922: southern Australia: Smirnov & Times
(1983); These tixe were synooymized with the
northern hemisphere Po aduncus by Smirnov &
Timms (1983: 24). Frey (1991b), after examination
of these and other extensive materials, considered
that P udurcis does not occur in Australia, The
entities of these various axa have yet to he
resulved.

P. denticulatus Birge, 1879) non Chydoris denticudanes
Henry, 1919, NSW (Smirnov 1971); Vie. (Timms
19737); 7Old, ?NT, 7WA (Timms 1988).

Comment: After examination of the available material

in the Australian Museum labelled as P denciculatus,

Frey (1991b) concluded that none of the specimens was

of the nominated taxon and they were Certainly not

conspecific. He considered that the species probably
was absent from Australia.

Zoopgeography

Qur comments here must be considered preliminary.
given the rapid changes in cladoceran taxonomy in
reeent years. Widespread recognition of non:
cosmupolitinism has provided a significant impetus
to a more critical approach (cf. Frey 1982b), Ibis clear,
particularly from some of Lhe last works of Frey (1991,
b),. that a considerable degree of endemism is obscures!

CLADOCERA RECORDED FROM AUSTRALIA 37

by cosmopolignn Warnes in the Austratian fauna [avr
opiion, any cladoceran in Australia relerted to a
species described from the ortiem hemisphere shoul
be viewed with suspicion until critics! reviews of all
fymilies, to the standard of Frey (991b), are achieved,

On present evidence, Australia hay more cladoceran
species: 165 vs ca. 120 (Frope) and 10 (U0S.A.) than
are found inother Comparable areas Overall the level
of endemicity Stands. al 43%, with five additional taxa
also known trom New Zealund. ie Ausiralisjan
endemicity is ev. 46%. To the endemic genera
Neotlriv. Pseudamdina (Macrothricidae),
Archepleurosus, Australochyderus, Monape, Rak,
Rhyachovhydorns and Saveiae (Chydoridae) listed by
Smirnov & Tinims 0983), Celsinonan, Leberis,
Planicivclus snd Plurispina (Chydoridae) are added.

Most radiation appears to have wecurred in the
Chydoridae; 45 of the 94 recognized species (48%)
are endemic. Australia may differ from other cegions
in the Selvctive pressures which cause genetc
divergence (4 Prey 19916). In any event there has been
poarked specmbon iqarcas Where water 1 limited, c.g.
southwestern WA, where the habitats are not those
‘normally Muicative or supportive of a diverse aguatic
mivrolauna, @y:. rock pools. salinized wetlands. The

aquatic microfauna of these habitats, iy cemimon wath
those of billabongs and wetlands of the opposite side
ol the Continent. have generally been jenored. We
suspect (hat a diverse array of indigenons clacocerans
is yet lo be discovered.

Acknowledgments.

Our eladoceran work continues to be wded by the
umcomparable- observations dnd literature collection of
(he late David G, Frey, Without his expertise and keen
eye and support during 1982-42 [he study of Australian
chydorids would have been overwhelming. Our gratetul
thanks wy Libby Frey, who supported RJS jo travel to
Bloomingjon in Dec, 1992 to help sort the incomplete
manuscripts and recover Australian material from the
Frey collection, For their willing assistance in his
neophyte ‘cladocerai’ days (before being subverted to
rovers), RJS thanks Nikolai Smirnov (Moscow) and
Brian Tinims (Cooranbongy. Hendrik, Segers (Gent)
kindly sought obscure references unavailable in
Australia. Our thanks also lo Alice Wells (Canberra:
taxononuc advice), Jan Bayly (Melbourme; distaibution
records) and Jean Just and the Australian Biological
Resources Study (Canberra: grant support)

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SKUSEMYIA ALLOCASUARINAE, A NEW GENUS AND SPECIES
OF CECIDOMYIIDAE (DIPTERA) DAMAGING LATERAL
BRANCH BUDS OF DROOPING SHEOAK,
ALLOCASUARINA VERTICILLATA IN AUSTRALIA

By P. KOLESIK*

Summary

Kolesik, P. (1995) Skusemyia allocasuarinae, a new genus and species of
Cecidomylidae (Diptera) damaging lateral branch buds of drooping sheoak,
Allocasuarina verticillata in Australia. Trans. R. Soc. S. Aust. 119(1), 41-46, 31 May,
1995.

A new gall midge genus Skusemyia and a new species S. allocasuarinae are described
from South Australia. Detailed descriptions of the larva, pupa, male and female of the
new species as well as its gall on drooping sheoak, Allocasuarina verticillata, are
given. The new genus is placed in the subtribe Schizomyiina within the tribe
Asphondyliini.

Key Words: Cecidomyiidae, Cecidomyiinae, Cecidomyiidi, Asphondyliini,
Schizomyiina, Skusemyia gen. nov., Skusemyia allocasuarinae sp. nov.,
Allocasuarina verticillata, South Australia.

Transactions af the Royal Saeiery ofS. Aust. W998), TA}, Alaé

SKUSEMYIA ALLOCASUARINAE, A NEW GENUS AND SPECLES OF CECIDOMYHDAE

(DIPTERA) DAMAGING LATERAL

BRANCH BUDS OF DROOPING

SHEOAR,

ALLOCASUARINA VERTICILLATA IN AUSTRALIA

by P. KOLESIK*

Summary

Kovesi€, P1995). Skasernvin allecasuerinae, anew genus und species of Cecwdomyiidae (Diptera) damaging
lareral branch buds of droopme sheoak, Allocasuarina vertieillake in Austealia. Trot. Bo Soe: 5. Aust. U9¢1).

4\-46, 41 May, 1995.

A new gall midge genus Skusemyia and anew species & alfpeesmarinae are described fram South Austrulia.
Detwiled descriptions ofthe lurve, pupa. mate and female oF the new species ds well as Tis pall en deoeping sheowk,
Allocasuarina vertictlum, are given, The new genus is placed in the subtibe Schizomylina within the tribe

Asphondy liar,

Kiey Words: Cecidomyiidae, Ceedomylinac, Cevidomytidi, Asphondylini, Schizomy ina, Skavenoia gen, nov,
Skuseniva altocusuarinae sp. now. Allocnsaurnny verticillata, South Austratia

Introduction

“Vhis paper is the second part of a study Gn the South
Australian Cecidomytidae. Kolesik (in press) described
the first South Ausiralian gall midge species,
Becinetivarnia matarskii, trom Encalypies fasoiculosa.
A new species is deseribed here that was found
damaging the lateral branch buds of drooping sheoak .
Altlocasuarina vertieillate (Lam.) L. Johnson
(Casuarmaceae) in South Australia. The large nurubers
of infested and ultimately killed branch buds al one
site indicate that this species could become a serious
pest (Pig. 31). The new gull midge has one generation
per year in Adelaide, South Australia.

A new genus is erected for the new species. It
belongs to the subfamily Cecidomytinae and superinbe
Ceeidomyiidi. tt is compared to other known genera
of the subtribe Schizomyiina of the tribe Asphondytiinr.

Materials and Methods

A survey of the Cevidomyiidae associated With galls
on plants in nature conservation parks around Adelaide
was carried out beeween November 1992 and May
1993. All galls sampled were dissected and examined.
Those which contained larvae of Cecidomyiidae were
described, photographed and conserved for luter
authoritative identification of the host plant species.
The larvae from the gall kind described here were
processed in lwo ways, A small number was preserved
in 70% ethanol after notes were made on their colour
The larger number was brought to the laboratory to
rear to adults. Here the galls. were carefully dissected

* Department of Horticulture, Viticulture and Oenology,
Faculty of Agricultural and Natural Resource Scienves, The
University of Adelaide, PMB L, Glen Osimond, South
Austealta S064, Austeatia-

and the larvae transferred with entornolagical tweezers
into rearing pots containing sterilised, wet sand
(Skubravil & Skuhravy 1960). Pots were examined daily
and emerged adults preserved together with their papal
skins in 70% ethanol after their colour had been noted.
Microscope mounts of a series were prepared by
miuverauion in 20% KOH. followed by processing
through distilled water, 70 and 99% cthanol and xylene
(o Canada balsam mountant for examination by
interference-contras| and bright-light microscopy.
Larvae, entire or dissected into two pieves, and entire
pupae, were mounied doarso-venirally or laterally,
Adults were dissected into tour (female) or five (male)
pieces und the particular parts were mounted
separately. Wing and head were mounted frontally.
thoras laterally, abdomen dorso-ventrally or laterally
and male genitalia dorso-yentrally, Measurements were
made with an eyepiece reticule. Drawings were done
with the uid of a camera lucida. The type series and
other material retained in 70% ethanol are deposited
in (he South Australian Museum, Adelaide [SAM] and
United States National Museum. Washingtoo |USNM],

Genus Skusemyia ven. noy.

Adult

Wings with R. joining C al wing apex, Rs weak, R,
joining C near wing mid-length. M, 5 absent. M,
weakly developed, Cu forked. Maxillary palpus with
4 sepments. Male antenna with 12 Nagellomeres.
Female antenna with I! flapellomeres, the last three
successively and progressively shorter, the last apparent
Magellomere evidently a combination of the eleventh
and twelfth, Flagellomeres cylindrical with necks, first
and second not fused, with Jong and stout setae in two
whorls, bearing closely appressed circumnftla. Tarsi
with first segment substantially shorter than the secand.

43 rl KOLESIK

first Lersomeres lacking yventrodistal spine, tarsal claws
siinple. empodia mach shorter thun claws. Male
jerminaha: gonovoxites Free ventrally, produced to form
a roundly tiangular apie! process; gonosty)us situated
dorsally on gonacoxtie, short and wide, with truncated
apex bearing teerh of umtorm length: cerci simple,
founded upically. hypopreet divided into two apical
lnbes; claspettes large; aedeagus long. srout. tapering
distally, Female abdominal sternite 7 longer than
Stvrpile 6, Ovipositor: protructile, elongate, sclerolized,
without basal lobes: verct fused. divided at apex,
SEHOSL,

Larva

Head capsule with short posterolateral apodemes:
Ameona short. Sternal spatula bilobate. Anus ventral.
Thorucie and first through seventh abdominal segments
with 6 dorsal, 2 pleuraland 4 ventral papillae. Biphth
abdominal segment with 2 dorsal, 2 pleural and 2
ventral papillae. Terminal segment wilh § dorsal umd
Aanal papillae. AlL collar Unoracie and abdominal
papillae Usetose with exception of the dorsal pupillie
onthe terminal seyment that are bearing very. short
sche,

Pupur

Frans without projections, One of three lower fayial
papillae with a seta, Lateral facil papillae absent,
Cephalic selerite with two strongly chitinized swellings
und two papillae wath long sete. First through seveoth
abdominal seyments with 6 dorsal asctose pupillac and
one pleural setose papilla. Sewend through eighth
abdominal segments dorsally with 3 indistines
(ramaverse rows of spines On anterior hall, Last
abdominal seament with large pouch. emarginate
medially but nat completely divided in io parts

Whe species: Skusenvia allocasiarinad sp. wv.
Hiyinology

The genus 6 named after FAA. Skuse, author in
I888/(890 of the first taxonomic studies on Australian
Cee domy ndite,

Remarks

Skasemyiee lits in the tribe Asphondyliini of the
supertribe Cecidomyiid) because the female seventh
abdominal stermle 15 1.5 lines as long as the sixuh
sternite and the eqhth tergite i wider than the seventh
tergite, combined with the male genitalin having a
ventroapical gonacoxal lobe and a dorsally situated
fonostylus that is about as broad as long: it belongs
lo tle subtribe Sehizamyiina becuse the first
tarsomeres lack a ventrodistal spine. the male genitalia
have claspettes, and the female lacks cere\-like lobes
immediately posterior to the eighth tergue (Gagne
994), Within the subiribe Schizamyiina, Skusemyia
resembles most closely Placerhela Rubsaamen. known
from three European and one El Salvadorean species

(Molin 1960, Skulirava 1986). The male aacennae of
the iwo genera are the only ones in the subtribe with
relatively simple cireumfila and with fagellomeres
made ap ofa bulbous basal nodé and long neck an!
resembling these of Desineure wid relalives
(Oliotrophini: Lasiapteridi). The female antennae of
Skasennvie und Placochela ure also similar except that
the eleventh and twellth layellomeres of the new gerius
are apparently amalgamated, which is unique mi
Asphondyliini. The genitalia of both sexes are generally
similar cso (Mohn 196))_ except in details of the
oyipositer whieh differs in Skuse beeause the vere
are diserete, at least attheir apices. and the distal setae
are longer. The immature stages of Skusemyia are
unique in Schizomyina, The pupal cephalic selente
has two swellings thal are longer than the antermal
homs. The larva has very reduced papillar sere. and
the papillae of the terminal seyment are all situated
at the end al separate lobes.

Skusemyia allacasuarinae sp. nov-
FIGS 13

Halorype: Oo, Black Hill Conservation Park, Sauth
Australia [34°53'S., I38°44°E.|. 15.11.1993, P
Kolesik. reared from larva tron lateral branch bud yall
of Allpeasuarina vertiettaia (ham) L. Johnsen
salnpled 25.1, 1993, 121270. [SAM].

Allvlype: 2, same dala bub emerged 16.11, 1894,
(2127) [SAM].

Pararypes (all sampled with holotype): 4cr.cr and
$09, emerged 13-19.111-1993; 7 pupae, emerged
13-18 71-1993; § larvae [SAM].

Other maternal 4 F 39 9 |SAM[|. 300,390 9
[USNM|..4 pupae [SAM] and 3 pupae |USNM], same
dats us holorype but emerged 16-22-1717. 1993. 6 [SAM]
and 3 larvae |USNM], same data bat sampled
25111993, 3 larvae, Wistow, South Auestratia
[35°07 'S_. 158°53’E_], 23.41.1993, P. Kolesik, on A.
vertierdlate.

Description
Male (Fig W)

Colour: antenna prey, head black. thorax. brown.
abdomen with sclerotized parts black and non
sulerotized red, legs yellow, all setae black, Toul legeui
of the body 3.24 mm (range 315 9 3.33-mm), Wing
length 2,44 mom (2.34 - 2,52), width 0.98 mm (0,93
- 1,00). Wing membrane und especially veins densely
vovered with selac, 50 - 60 xm. Antenna total lengeh
1.72 min (1.57 - 1.83); Mavellomeres with stout sete
54 - 160 am, longer at the distal whorl than at the basal
whorl; closely appressed circumfila consisting of one
transyetse and one longitudinal bands. Eye bridge +
Ww 4 facets medially, eye facets rounded. Claws curves
beyond the second third. 39 wm (37-41). Empodiuny
I pm (8 — 14).

A NEW GENUS AND SPECIES OF CECIDOMYIDAE 43

Fig. 1, Male of Skusemyia allocasuarinae gen. et sp. noy.: A. genitalia in dorsal view; B. wing, C. last tarsal segment with
claw and empodium; D. head in frontal view; E. sixth flagellomere; F, last three flagellomeres. Scale bars = 100 pm.

dd P. KOLESIK

ra
| r
| 2
ry ( )
} 7 la B} in

c-— D-——_ E
Fig. 2. Female al Skusennia allacasuarinae gen, et sp. nov.
A. ovipositer in ventral view, B. end of ovipositor im ven-
tral View; C. first four antennal segments; D, fast five Mag-
cllomeres, E. sixth (itgellornere, Seale bars = 100 wn,

Terminalia: gonocoxite setose and setulose: zonostylus
selose and setulose, bearing 17 - 20 sclerotized tapering
teeth that are narrow and about 15 pm in length:
hypoproct bearing one setaon each lobe. setulose; cerci
deeply divided medially, setose and setulose; claspettes
setulose

Female (Fig, 2)

‘Total length of the body 3.63 mm (3.41 - 3,80). Wing
length 2.65 mim (2.32 - 2.80), width 0.89 min (0,74 -
1.00). Antenna total length 112 mm (0,95 - 1.20),
flagellomeres with setae, 5] - 15 jan, Last Mayellomere
with a shallow constriction medially, Seventh
abdominal sternite about 1.5 times longer than sternite
6. Ovipositor with one ventral sclerotized longitudinal
band forked distally. distal hall of ovipositor with §
-9 pairs of 6-9 jm Jong setae, cercus with 8 pairs
of 5 ~ 48 pm tony setae. Colour.and other characters
as an male.

Mature larva (Fig. 3A-D)

Colour red. Total length 3.70 mm (3,24 - 4.44).
Integument smooth, ventrally with several transverse
rows of spiculae on antenor half uF each segment. Head
capsule width 61 jam (53-77). length 63 wm (S51 - 74),
length of posterolateral apodemes 61 jum (51 - 64),
Antenna 17 pn (15 - 20). Sternal spatula 268 yim (230
- 320) in length, with apical enlargement 86 am (77
- 105) in width and 42 yom (38-51) in length. Terminal
segment dorsally with 8 lobes bearing papillae: @ with
very short setue and 2 with corniform sete:

Pupa (Pig. 3F-H)

Colour, prothoracie spiracle, cephalic swellings and
untennal horns dark brown, remaining: parts pale
brown. Total length 3.03 mm (2.46 - 3.56). Integument
covered with spiculae, ventrally 2-4 mand dorsally
4-6 um long. Antennal horns 33 ym (28 - 38) long.
Cephalic swellings 77 yn) (74 - 80) in length. Cephalic
pupilla with seta 76 aim (58 - 90), One of three lower
facial papillae with seta 45 pm (38-51). Prothoracie
spiracle [46 pm (140 - 151) long and 23 ym (20-28)
wide across the base, with trachea ending at apex. Seta
on pleural papilla 9 am (8 - 10), Dorsal spines of the
first row [3 . 20 in pumber, 5 - 30 pm, spines of the
second row 13 - 20 in pumber, 25 - 45 yam: spines of
the (hird row 9 - 12 tn number, 35 - 65 jum.

Gall (Fig. 31)

Swollen lateral branch bud, forming spherical to
spindleform rosette, 7 - 12> mm in diameter,
polythalamous, pale brown in colour. One larva in cach
of the 2 - 3 cells. Gulls appear in January - March
Larvae eave galls to pupate in the soil.

Eryinalogy
Derived from the generic name of the host plant

Acknowledgments

I thank the South Australian Museum, Division of
Natural Science, for providing the facilities that assisted
this Work, Lam grateful to the museum collaborators
Ms J, A. Forrest, Dr E. G. Matthews and Mr DB.
Hirsi for their support. The Ministry of Environment
and Planning, South Australia, kindly gave permission
to collect in the nature conservation parks of Black Hill,
Morialta and Cleland. I wish to thank Mr M, C.
O'Leary, State Herbarium of South Australia, Adelaide,
for the identification of the host plant species, | an
grateful to Dr Raymond J. Gagné, Systematic
Entomology Laboratory, U.S. National Museum,
Washington, tor carelul review of the manuscript

A NEW GENUS AND SPECIES OF CECIDOMYIIDAE

viel, 4 f it

TaN

tem

Fig. 3. Skusemyia allocasuarinae gen, et sp. nov.: A. - D. larva; E. - H. pupa; I. gall. A. stigma; B. head capsule in dorsal
view; C. anal segment in dorsal view; D. sternal spatula; E. sixth abdominal segment in dorsal view; F. last abdominal
segment in dorsal view: G. anterior end in ventral view; H. prothoracic spiracle, I. Allocasuarina verticillata - lateral
branch bud galls caused by Skusentyia allocasuarinae gen, et sp. noy. Scale bars = 100 wm. A. - H.; 2 em I.

46 P. KOLESIK

References

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

KOLESIK, P. (in press) A new species of Eocincticornia
(Diptera: Cecidomyiidae) on Eucalyptus fasciculosa in
South Australia. J. Aust. ent. Soc.

Moun, E. (1960) Gallmiicken (Diptera, Itonididae) aus El
Salvador, 2. Teil. Senckenbergiana Biol. 41, 197-240.

(1961) Gallmiicken (Diptera, Itonididae) aus El

Salvador. 4. Zur Phylogenie der Asphondyliidi der
neotropischen und holarktischen Region. /bid. 42,
131-330.

SKUHRAVA, M. (1986) Family Cecidomytidae pp. 72-297 In
Sods, A. (Ed.) “Catalogue of Palaearctic Diptera.” Vol.
4. Sciaridae: Anisopodidae (Akadémiai Kiad6, Budapest).

& SKUHRAVY, V. (1960) “Bejlomorky.” (Statni

zemedelské nakladatelstvi, Praha).

TRANSACTIONS OF THE

ROYAL SOCIETY
OF SOUTH AUSTRALIA

INCORPORATED

VOL. 119, PART 2

NEW GENERA, SPECIES AND A NEW SUBFAMILY OF
XYALIDAE (NEMATODA: MONHYSTERIDA) FROM
OCEAN BEACHES IN AUSTRALIA AND THAILAND

By WaRWICK L. NICHOLAS* & AIMORN C. STEWART*

Summary

Nicholas, W. L. & Stewart, A. C. (1995) New genera, species and a new subfamily of
Xyalidae (Nematoda: Monhysterida) from ocean beaches in Australia and Thailand.
Trans. R. Soc. S. Aust. 119(2), 47-66, 31 May, 1995.

Five new species of Xyalidae are described from Australian ocean beaches and one
from Thailand. Gullanema fragilis gen. nov., sp. nov., possesses many long cervical
and somatic setae, but is distinguishable from other similarly hirsute Cobbiinae by a
distinctive narrow peri-buccal region. Rhynchonema tomakinese sp. nov., R. collare,
sp. noy. and Prorhynchonema gourbaultae sp. nov., in the Rhynchonematinae, differ
from congeneric species by their spicules and amphidial regions. R. chiloense
Lorenzen, 1975 and R. gerlachi Vitiello, 1967, and P. warwicki, Gourbault, 1982 are
commented upon. Two new species with characters intermediate between the
Cobbiinae and Rhynchonematinae are placed in a new _ subfamily, the
Corononeminae. The cervical region, enclosing a cylindrical buccal cavity, is shorter
and less attenuated than in Rhynchonematinae, longer and narrower than Cobbiinae.
The labial region its set off from the buccal region by a narrow indented ring.
Corononema parvum, gen. noy., sp. nov., is described from Australia and C. thai sp.
noy., from Thailand, the two differing in the shape of the head.

Key Words: Taxonomy, nematodes, Xyalidae, beaches, Gullanema, gen. noy.,
Rhynchonema, Prorhynchonema, Corononema, gen. nov.

Transactions of the Royal Somety of S. Aust (1995). 1t9(2),47-06

NEW GENERA, SPECIES AND A NEW SUBFAMILY OF XYALIDAE
(NEMATODA; MONHYSTERIDA) FROM OCEAN BEACHES IN
AUSTRALIA AND THAILAND

by WARWICK L. NICHOLAS* & AIMORN C, STEWART*

Summary

NicHouss. W, bo & Stewart, AC. (1995) New genera, species and u new subhimily of Syalidae (Nematoda,
Monhysterida) [ror ocean beaches in Australia ancl Thailand, fins. Ro Soc. 8. dst 192), 47-66, 31 May, 1995,

five new species of Xyalidac are described from Australian ocean beaches and one from Thailand, Gullaneme
fragthy gen, low. sp, nov, possesses many long cervical and soralic sefae, but is distinguishable from other
similarly hirsute Cobbiinae by a distinctive narrow peri-buceal region. Rivachanema tomakinese spo noy.. KR.
collate, Sp. nov. and Proriyichoneme gourhantiae sp, nev.. in the Rhynchonematinay, ifler from congeneric
species by their sprules and amphidial regions, R, chileense Lorenzen, 1975 and RK. gerfarhr Vitiello, 967. and
P. warwteki, Gourbault, 982 are commented upon Two new species with characters intermediate between the
Cobbiinae and Rhynehonematinae are placed ina new sublamily, the Corononeminae. The cervical rexion, enclosing
a cylindrical buceal cavity. is xhorter and less attenuated than in Rhynehonematinae, longer and narrower than
Cobbiinac, The labial region is set oll [rom the buceal repion by a narrow indented ring. Corwioiema parvnin,
pen nov., ap nov, is described from Australia and. thai sp. nov., from Thailand, the twe differing in the shape

of the head.

Key Worns: Taxonomy, nematodes, Xyulidae, beaches, Gullanema. gen,

Prorkvachonend, Caronanema, gen. nov.

Introduction

Neratodes, have been collected from sandy beaches
on the southern, castern and northern coasts of
Australia and southern Thailand. Previously we
(Stewart and Nicholas 1994), described eight new
species of Xyalidae Chitwood, 1951 belonging to well-
known veners of Cobbiinae de Coninck, 1965, In this
paper we describe 4 new species and genus of
Cobbiinae, numely Gullanema fragilis gen. anv. . sp.
nov., and threé new species from the other subfamily
of Xyalidae, the Rhynchonematinae de Coninck, 1965,
namely Rhyachonemu temukinense, sp, nov,, R.
collare, sp. nov. and Prorhynchanema gourbanttae sp.
nov. Species of Rhynchonente Cobb, 1920 are common
in Australian sandy beaches, as in other parts of the
world, and we cotiment on Australian specimens
belonging to two previously deseribed species, The
genus was comprehensively reviewed by Lorenzen
(1975). We also erect a new subtamily, Corononeminuae,
w hold Corononema parvum gen, nay:. sp. nov. and
©. thai sp. nov., with characters intermediate between
Cobbtinae and Rhynchonematinae. The last named
species is from Thailand.

* Division of Botany and Zoology, Australian National
University, 0200 Canberra. Australia,

voy, Riivnelaneni,

Materials and Methods

Collections were made on the 90 Mile Beach ut
Seuxpray, Vicloria (147° 23'E, 38 46'S); South
Moruya, Broulec, Rosedale, Tomakin and Kioloa
beaches, New South Wales (berween 150° 9'E, 35
55'S and 150 20'E,*35 32'S): Southport beach,
Quéensland (153° 25'B, 27° 58'S) and Rapid Creek
beach, a suburb of Darwin, Northern Territory: (30
50'F, 12° 23'S). Some specimens were also collected
trom Pathaya beach. Chonburii. Thailand (100° 53’E,
12) 45'N). Specimens were collected in samples of
sand dug up at low tide between the tidemarks to a
depth of 40 em. Exceptions were samples of sub-tittoral
sand wken from a boat in shallow water off Cronulla,
New South Wales (151° 10'E; 34° 05'S),

Nematedes were extracted from the sand by re-
suspension in tap water, allowing the sand to seule and
collecting the nematodes ona 50 jum. nylon mesh sieve.
The nematodes. were washed olf the sieve imto sea
water, then fixed in 5% tormalin in sea water.

Specimens were picked up under the microscope
with a fine pipette and transferred to 5% aqueous
glycerol which was slowly dehydraled to anhydrous
glycerol at 40°C. Permanent tnounts were made in
anhydrous glycerol, and the cover slips ringed wiih
Glyceel (Gurr). Glass beads (ballatini), selected under
the microscope to approximate the diameter of the
nematodes, were used to support the cover slips.

ak WARWICK L, NICHOLAS & AIMORN C. STEWART

Drawings and measurements were made using a
caverta lucida, When mounted the nemilodes Jie ou
their sides presenting a lateral view (except in Pig, 19)
Our draWings show setae on one side only, that lying
uppermost as mounted -

Measurements are in pau front specimens fixed and
mounted in this way, De Man's ratios are piven, i.e,
a = body length divided hy greatest body width, b
= length divided by length of pharynx, ¢ = length
divided by tai) length, c = tail length divided by width
at anus. V = anterior end to vulva as @ percentage
of body Jength, and spicule measurements are arc
length, For Rhynchonematinae, Lorenzen's (1975)
formula has been used to sumumarise the characteristic
features of each species described. In this formula a
Jeter code, referring to drawings of characteristic
structural leatares, is used to describe successively the
form of body annulation, symmetry of the spicules,
form ot the spicules, annulation surrounding the
amptids, relative size of amphids in both. sexes,
position of amphids relative to end of buccal tube, form
of buccal cavity, and form of the vulva. An important
character is the direction, either towards the anterior
or towards the posterior, of the saw-tooih edge of
culiculur annulation (reiferartie dick nit sageatiger
Aussenkortur). In some species there is an abrupt mid-
body change im direction. Where this is so, the distance
from the head end to the inversion is expressed as a
percentage of body length. In other species only the
cervical and tail regions: have saw-tooth amulation,
the inid region of the body having rounded annule
profiles,

Scanning electron micrographs (SEM) were made
from specimens that had been postfixed in 14% aqueous
osmium tetroxide, freeze-dried and coated with gold
palladium. Type material is deposited im the South
Australian Museum, Adelaide, and the museum's
numbers are given im the text, Holotype numbers (all
males) follow SAMA and the prefix V. Some paratypes
are On the same slide, to give both a male and a female.
Other paratypes are on slides labelled AHC followed
by Oo nuniber.

Taxonomic descriptions

Tamily Nyalidac Chitwood, 1951,

Annulated cuticle, ciccular eryptospiral amphids,
female with single prodelphic ovary situated to left of
gul. male with one or two testes, anterior situated to
lefi GF gut, posterior if present to right of gut.

Subfamily Cobbiinae de Coninck, 1965

With characters of family. Head and cervical region
only slightly attenuated. base of buccal cavity conical,
enclosed by pharyngeal musculature, 10 sensillu in
second ring of cephalic sensilla, usually segmented.

Genus Guilanema goa. nov.

Type species: Gullanema frayiliy
Diagnostic definiuten

With characters of Cobbrinae, Cephalie region
surrounding conival buccal! cavity eylindrteal. sec ol
from wider cervical region. Strong mouth ring
supporting six labial setae and six thin flexible lips.
Very long angulated double and iriple sets of cervical
setae and numerous long single and double sematir
selae extending as far as the anus.

Etymalogy

Named after Dr Penelope Guillen im the School of
Lite Sciences, Australian National University

Gullanema fragilis sp nov
(FIGS 1-12)

Holotype; Male, SAMA V4259, Kioloa beach. NSW.
27. x1.J986.

Pararypes; 4 males, AHC 24808 and 24813, Kiolog
beach, NSW, 27.21.1986, 15.x7.1987 respectively: AHIC
24809 und 24812, Broulee beach, NSW, 3viii 1980. 8
females. AHC 24810 and 24813-15. Kioloa beach.
NSW. 27.15.1986, 3.viii 1980, 26.yii.1978 and 15.01 1987
respectively; AHC 24812, Browlee beach, NSW,
3viii. 980; AHC 24816, Moruya beach, NSW,
T.xuU.t98%, AHC 24817, Scaspray beach, Vic.
3.411 1988,

Meayurementy: ‘Table |

Deseriprion of Holarype male,

Body cylindrical, cuticle finely annulaled, tail w long
narrow tapering cylinder, Head widens sharply at buse
of narrow peri-buceal ‘iret-lke’ region enclosing
buceal cavity, Almost cylindrical buteal cavity, taperest
al base, six longitudinal bars reinforce buccal cavity,
strong circam-oral mouth ring, surmounted by six
fexible very thin lips, about 6 gm high. and six 4m
long inner labial setae. Six 18 wm outer labial, four
12 pin cephalic setae, in one ring, insert near hase of
peri-buccul ‘turret. Complex hexaradiate array at
cervical setae insert between base of buccal region and
amphids: most anterior, six IL jan short setae; next,
six longer unequal pairs of cervical setae, 36 and 14
pm; then six longer triplets of unequal length, 166, 102
and 50 am. The successive sels of cervical setae nut
inserted in rings around the hody at the same level,
but cach set slightly staggered relative to the long axis
of body. Labial, cephalic and cervical setae segmented.
Amphids, almost circular, 9 xin diameter, 31 yan behind
mouth. Long unagual pairs and single setae inser
irregularly between amphids and anus. longest. 209
pun, iN pharyngeal region. Also a few shart setae along
hody continue posterior to anus. Two caudal setae, 168
and 145 wm long, at tip of 230 um tong tail, Pharynx
cylindrical, cardia conical, rectum short, Two

NEW XYALIDAE

TABLE 1. Measurements of Gullanema fragilis gen. nov., sp. nov.

Type

Leneth

Width

Lips

Inner labial setae
Outer labial setae
Cephalic setae
Amphid diameter
Mouth to amphid
Buceal cavity
Nerve ring
Pharynx

Mouth to vulva
Mouth to anus
Tail

Anal Width
Spicule
De Man’s 4

" b

" c

ul V%

nn ttt tttEIttESESS

* Excluding female with truncated tail (n=7)

Figs 1 and 2. Head of male Gullanema fragilis sp. noy. Setae, which are annulated, are illustrated only on one side of the body.

Holo
Male

1443
36

=o w
, oH HKHeraeam

Male Paratypes n = 4

Range
1368 - 1482
24 - 36
3-8
8-10
22 - 26
13-19
7-8
27-35
9-18
68 - 90
262 - 330

1015 - 1293
17] - 225
21 - 33
37 --46
41-55
44 -5.4
65-9
6.8. -9.1

**Truncated tailed female

Mean + SD
1444 258
29 4.85
a: 2.10
9 1,16
24 2,07
16 2.50
78 0,50
30 3.78
13 3.78
77 9.27
290 30
1200 126
199 24
26 5.25
42 4.43
SI 6.74
5 0,43
73 0,70
1.01

7.83

Female Paratypes n =8

Runge
125] - 2025
35 - 62
5-9
7-9
22-29
15 - 20
71-9
22-45
10-23
82 - 130
275 - 365
751 - 1279
JOL9 - 1645
30** - 380
21-48

26-43
3.9-S,5
41,7** - 8.6
L1**-11
74-81

Mean + SD
1580 261
47 8.82
6.9 0.58
8 0.93
25 2,06
17 0.53
7.8 0.70
32 7,28
1S 4.30
100 17
328 28
981 160
1332 213
279+ +.
347.91
34 7.36
AS 0.50
6t ¢
&.2* *
76 2,58

3) WARWICK 1

a :

ranger wr

Figs 3 and 4. Gullanema /rayilis sp. nov. 3, entive male, 4.
spicules.

outstretched lestes, antertor to left of gut, posterior to
right, gonoduct filled with spermatozoa, spicules weak,
hardly cephalated, uniformly curved, blunt bilid tips,
48 yum long.

Puratypes, The numbers and location of somatic
setae are rather variable, especially the oumber of
caudal setae, which vary from one to three of unequal
length, possibly due to breakage during preparation.
The number dnd location of short cervical setae are
varluble, 9-id ym long, Two sets of longer cervical
setae are consistently present, the first set of six
doublets of unequal length, the second of six triplets
also of unequal length, but with (he points of insertion
of each group staggered with respect to the longitudinal
body axis. As an example of setal Jength in one female
paratype doublet 58, 42, triplet 99, 62, 45, cervical
200), 55 pm. The post-amphidial setae apparently not
inserted in regular rows or circles, the most anterior
the Jongest, decreasing to short scattered setae behind
the anus.. The distance the amphid lies behind the
mouth is rather variable both in absolute terms and
relative to pharyngeal length. Female paratypes differ
significantly from males only in reproductive organs.
Females are monodelphic and prodelphic.

Most specimens of both sexes have long tails, but
occasional specimens with much shorter tails haye been
found: an extreme case is illustrated in Fig. 5 (on slide

NICHOLAS & AIMORN ©. STEWART

AHC 24810, which also contains a second female with
a slightly longer tail), Because the degree of truncation
is variable and found on infrequent individuals amongst
numerous long-tailed individuals, the shert-tailed farm
is not considered a separate taxon but an infrequent
aberration, Because of the variation in tail length, the
position of the vulva is more usefully related fo the
percentage distance from head to anus rather than the
more usual head to tip of tail,
Differential diagnosis

The new genus resembles Stemeria Mivoletzky,
1922. and less closely 7richorheristus Wieser. 1956,
in possessing many very Jong setae. In Sremneria, as
in Gullanema, but unlike Trichotheristus, groups of
long cervical setae are inserted between the mouth and
the amphid. Gullanema is quite unlike either genius
in the formof the head and shape of the buccal cavity,
In these two genera the head tapers gradually to the
base of the large lips, whereas in Gullanema gen. nov.
the head narrows sharply to a cylindrical ‘wrret-like’
region surrounding ihe buccal cavity, The outer labial
und cephalic setae are inserted near the base of the
turret, the inner labials at the mouth ring. The lips are
smaller, supported by 4 very strong mouth rim. At the
specific level, the weakly cephalated spicules with
blunt bifid tips are an important character.
Habitat

Sandy ocean beaches.
Dispributton
New South Wales, Victoria,

Subfamily Rhynehonematinae de Coninek. 1965,

After Lorenzen, 1975: head und mouth region very
attenusted, buccal cavity long and tubular, cuticle
strongly annulated, first cephalic annule wider.
amphids circular. tail conical. Uncertuinty regarding
cephalic sensilla because of difficulty in resolving tiny
head with hight microscope, inner Jabial sensilla
undescribed, single ring of 6 or \O cephalic setae. Less
than 900 um tong.

Genus Rhynchenenia Cobb, 1920

Buccal cuvity in two parts; short anterior chamber,
at level of cephalic setae, narrow tubular part at least
25 ym Jong., Amphids placed over or very close to
end of buccal tube. Male with two testes.

Rhynchonema collare sp. nov.
(FIGS. 13-18, 25-30)

Holotype: Male. SAMA. V4260, Rosedale beach, NSW.
5.11. 1986.

Pararypés: 2 males, AHC 24818, Rosedale beuch,
NSW, 5.111.1986 and AHC 24819a Broulee, NSW,
3yiii, 1980; 7 females. AHC 24819b-c, Broulee beach,
NSW. 3.vili J986,

NEW XYALIDAE

Figs 5 and 6. Gullanema fragilis sp. nov. 5. female, infrequent short tail form. 6. common long tailed female.

52 WARWICK L. NICHOLAS & AIMORN C. STEWART

_- OLS

Figs 7-12. Gullanema fragilis sp. nov. 7. head to show buccal region. 8. showing amphid. 9. spicules. 10. SEM of buccal
cavity. Il. entire nematode. 12. SEM cephalic and sub-cephalic setae. SCS sub-cephalic setae, OLS outer labial setae,
AM amphid, SP, spicule, BC, buccal rim, LF, lip flaps, ILS inner labial setae, CeS, cephalic setae.

NEW XYALIDAE 53

13,14,16,17,18 ———__________2's im

15 ———————. 50 pm

Figs 13-18. Rhynchonema collare sp. nov. 13. female head. 14. male head. 15. entire female. 16. inversion of annules. 17.
male tail and spicules. 18. female vulva and operculum.

54 WARWICK L. NICHOLAS & AIMORN C. STEWART

TABLE 2, Measurements of Rhynchonema collare yp. mer.

‘Type Holo
Male Male
Length 476 412
Width 20 Y
Body setae - 3
Ruveal eavity 53 ay
Amphid. length 7 1k
Amphid, width 10 10
Mouth to amphid 5? 40)
Mouth to nerve ring -
Pharyns 125 14
Mouth to vulva
Mouth to inversion’? 36 55
Mouth to anus 399 428
Anal width 17 1k
Tail 77 05
Spicule. long 16.8 18.7
Spicule, short 13.7 13,2
De Man's a 24 pad
w bh ams) an
Z e 6.2 63
£ c! 15 3.6
" VGE =

Mate Paratypes

Female Paratypes n=

Male Range Mein + Sb
579 311-552 333 19
7 20.23 218 110
17-17 7 0.00
52 47-34 3] 2,63
19 6-95 8 (.32
12 6-7 67 0.45
a2 44-55 Ay 4.00
AS R499 90 AR?
162 124 - 152 138 |3
316 -39] 3S 35
38 56-73 60 5.07
515 439 - 483 465 iv
16 15-17 15.6 ORY
75 34-74 65 8.72
Ih2 - - -
12.8 -
34 22-26 24 1.52
3.6 36-41 34 0.27
a7 71-99 8,3 142
47 3.0 -4.9 4.2 O48
- 61-73 69 5.41

Measurements: Table 2.

Description of Holotype mate

Very small, with long attenuated cervical region.
plump post-cervical region, rather long broad curved
conical tail, recurved at hp. Cuticle strongly annulated,
annule profiles sharply angled forward on Front half
of body, backward on posterior, abruptly changed
(inverted) at 56% body length; very thin somatic setae
spaced uniformly between amphid and anus, Buccal
cavity a shallow cup leading into long narrow parallel-
sided tube with strongly cuticularised walls extending
Jength of narrow cervical region to level of middle of
amphid. Six cephalic setae at base of buccal cup,
extremely large elongated amphids enclosing posterior
40% of narrow cervical region: no annulation between
amphids. Pharynx cylindrical, cardia heart-shaped.
Spicules unequal, weakly cephalated, lacking
rectangular bend, tips turned up, Large gubernaculum
encloses mid region of spicules, strong dorso-caudal
apophysis.

Paratypes: Amphids strongly dimorphic, in females
relalively large, but much smaller than in males,
separated by strong non-annulated cuticle. Vulva with
operculum, on which annulation greatly reduced or
absent. Terminal vaginal canal cuticular. Lorenzen’s
formula:- bwaokg,+.2,0.w: “new letter because
spicules do not correspond to any of those figured by
Lorenzen (1975),

Differential diagnosis

The new species can be distinguished from some
other species by possessing sharply angled annulation
along the whole body and by the possession of sexually
dimorphic amphids. It lacks the pre-anal supplements
and strong somatic setae and equal spicules found in
R. hirsutum Hopper, 1961, It more closely resembles
R. chiloense Lorenzen, 1975, and R. scutatum
Lorenzen, 197! but has quite differently shaped
spicules. The male amphids are larger than either of
these species, It differs from R. tomakinense sp. nov.
in the larger male amphid, absence of annulation
between amphids, and in possessing a cuticular vaginal
canal.
Habitat

Sandy ocean beaches.
Distribution

New South Wales.

Rhynchonema tomakinense sp. nov.
(FIGS 19-24)

Holotype: Male, SAMA V4261, Tomakin beach, NSW,
1. xi1986.

Puratypes: 3 males and 5 females, additional male and
3 temales on holotype slide: male and female. AHC
24820a, Rosedale beach, NSW. 22.xi.1986 and female,
AHC 24820b, Rosedale beach, 5.11.86; male, AAC
24821. Rapid Creek beach, NT. 31vii. 1986,

NEW XYALIDAE 55

19,20,21, 22,23
24

€ a,
oy,

Figs 19-24. Rhynchonema tomakinense sp. nov. 19. male head, dorsal view. 20. male head, lateral view. 21. female head.
22. vulva and operculum. 23. spicules. 24. entire female.

56 WARWICK L. NICHOLAS & AIMORN C. STEWART
TABLE 3. Measurements of Rhynchonema tomakinense sp. nev.
Type Holo Male Paratypes m= 3 Fermale Paratypes n ='5
Male Range Mean + SD Range Mean + SD
Length 331 438 - 495 ARS a0 4Xy - 502 545 ay
Width 22 19 25 AW 3.12 22-34 24 15
Body setac tl 9-18 12.6 - 13-16 [4.6 -
Ruceal cavity 44 Ax 57 49 473 j4- 49 47 1,52
Amphid, length 4 9-12 2 L7s 6-9 72 04,10
Amphid. width iT) K-12 8 2A 6-38 6.4 O89
Mouth to amphid as AO - 43 3 158 48 - 44 420 20s
Pharynx 121 15-110 10 LOS 120 - [ay 134 1)
Mouth to vulva - - - - - 345-440 ARG 46
Mouth to inversion 54 S266 57 7457 5) - 58 55 2.86
Mouth to anus 474 382 - 434 413 29 424-324 476 18
Anal width Is 17-18 17) O.SR IS 18 16 ),52
Tail 77 36 GK 60 6.93 64-73 68 4.05
Spicule is I) 19 I4.00 0 4.04 - - - -
Gubemaculurn 1] - - - - -
De Man's a 25 22-24 23 1.00 20 26 a4 245
" bh 4.6 41 47 44 O30 35-45 41 O44
We c 7.2 TL RS 7.9 ORS 74 87 8.0 O63
a e 43 3.) - 4.0 Ad 0.52 42-47 4.2 O40)
I Vo - - 66-734 71 285

Measurements: Table 3.

Description of Holotype male

Very small, cervical region long and attenuated, post-

vervical region plump, tail curved, conical, rather long
and broad with recurved tip. Strongly annulated, sharp
border angled forward, especially in amphidial region.
direction sharply inverted 54% from anterior end. Six
very small cephalic setae inserted at base of buccal cup,
from which narrow, parallel-sided cuticular buccal tube
extends length of cervical region. Large circular
umphids located over end of buccal tube, annulation
continues between amphids. Cylindrical pharynx,
heart-shaped cardia. Thin setae spaced along body
from cervical region to anus. Spicules cephalated,
asymmetric, slightly unequal size, without strong
rectangular Curvature, tips turned up. Gubernaculum
encloses middle of spicules, strong dorso-caudal
apophysis.
Paratypes: Amphids dimorphic, smaller in fenrale,
vulva with operculum, vaginal canal not cuticularised.
Lorenzen’s formula:- b,a,p*%r,+ .2.0,u. *new letter
because spicules do not correspond to any of those
figured by Lorenzen (1975).

Differential diagnosis

This species is very close to R. collare sp. nov. The
two are sibling species from the same beaches bur can
be clearly distinguished by several features. Annulation

continues between the amphids in both sexes, whereas
there is smooth cuticle in RK. collare sp, noy, The male
amphids are not as large. The vaginal canal is not
cuticular and the spicules show greater asymmetry, KR.
tomakinense resembles R. chiloense Lorenzen, 1975,
and R. scutatum Lorenzen, 1971, but has quite
differently-shaped spicules.

Habitat

Sandy ocean beaches.

Distriburion

New South Wales, Northern Territory.

Rhynchanema chiloense Lorenzen, 1975

Material examined

One male and one female and juvenile, off-shore,
Cronulla, NSW, one temale, Rapid Creek beach, NT,

Description

Lorenzen’s formula:- b.s,ijr,+,2,0,u, De Man's
ratios:- a = 18-23, b = 3,2-3.6,¢ = 8-10, V = 73%;
inversion of annulation 52%. Spicule are length 26,
more than twice length given in Lorenzen’s paper, but
that seems by comparison with the drawing to be chord
length, and by comparing drawings our male's spicules
are only slightly longer in a larger male. The cuticle
and body setae are stronger than in the other species
ol Rhynchonena we have found

NEW XYALIDAE 4)
26

20m . "eg / } >Oum
—_-__—_—_

: _ —_

Rigs 25-30, Rhynchonema collure sp. nov. by SEM. 25, male head, and cervienl region. 26. female head and cervical region.
27. annulation pharyngeal region: AAyichonema tomakinense sp. nov, 28, female head and cervival region 29. tale cervical
region. 30. female posterior, AM amphid. OP operculum, AN anus.

Habitat

Sandy ocean beach and shallow sub-littoral sand.
Distribution

New South Wales and Northern Territory.

Rhynchonema gerlachi Vitiella, 1967

Material examined

Three males and one female, off-shore Cronulla,
NSW,
Description

Lorenzen’s formula‘- i,s,z,r,=.2,u. Agrees with
Vitiello's (1967) description except that he did not
illustrate the vulva. Our specimens possess an
operculum but no strongly cuticular terminal duct
corresponding tou in Lorenzen’s formula. According
to Viticllo the spicules have bifid tips, but his igure
shows sharply pointed tips with one spicule rotated on
its long axis, The spicales in our specimens are just
like his illustration. According to Vitiello there are
three pre-anal papillae in males but these are not shown
in his illustration. We observe one 1o three minute pre-
anal bumps, but cannot determine whether they contain
papillae because of the strong annulation, Our
specimens are smaller than Vitiello’s adult specimens,
L=390-632 compured with 742-793 but De Man’ rauos
in our specimens:- a=20-26, b=3.5-4.8, c=6.340.2,
c =2,8-3.9. V=75% are in agreement with Vitiello’s,

Habitat

Shallow sub-littoral and intertidal beach sand.
Distribution

New South Wales

WARWICK L. NICHOLAS & AIMORN C. STEWART

Prorhynchonema Gourbaull, 1982

Buccal cavity in two parts; anterior short, at level
of cephalic setae, posterior tubular, not more than 15
um, Amphids circular, placed well posterior 10 end
of buccal tube,

Prorhynchonema gourbaultae sp. nov.
(FIGS 31-35)

Holotype; Male, SAMA V4262, Kivloa beach, NSW,
31.vii 1986,

Paratypes: 3 males and 3 females, AHC 24822, Kioloa
beach, NSW, 31.11.1976.

Measurements: Table 4.
Descriptian af Helarype male

Body cylindrical, anterior atretiuated from level of
amphids to smal! head, about 10% of body length: tail
conical, curved ventrally, tip slightly reflexed. Cuticle
weakly annulated, annules about 1 ym wide, first
annule wider, inversion of annule direction 50%: sparse
thin somatic setae. 11 ym Jong. Inner labial setae not
visible. six short cephalic setae in one ring, less than
1 ym long, amphids circular, cryptospiral, 30% body
width, situated much farther posterior, 9.5% of length,
about 55 annules from mouth, well beyond end of
buccal tube. Buceal cavity long narrow parallel-sided
tube, 3.8% of body length, slightly expanded at level
of cephulic setae; pharynx cylindrical, widens
gradually as body widens behind level of amphid,
cardia rounded. Spicules cephalated, smoothly curved,
slightly attenuated narrow spoon-shaped rounded tip,
gubernaculum simple plate with small caudo-dorsal
apophysis, three past-anal caudal glands,

Taare 4. Measurements of Prorhynchonema gourbaultae sp. nov,

Type Holo Male Paratypes n = 3 Female Paratypes n = 3
Male Range Mean Range Meun

Length 452 460 ~ 49| 476 439 - 453 445

Width 15 16-18 (7 19 - 20 19

Ruccal cavity 17 15-16 16 15-16 13

Amphid 2.6 25-3 28 3-35 3.3

Mouth to amphi 43 40 - 47 43 39 - 42 4\

Pharynx 144 1Q2 - [22 7 J 28 - 138 (32

Mouth to vulva - ---= - 30% - 322 414

Mauth to anus aK4 39) 6 420 4dOR 3K0 - 396 AMT.

Anal breadth \4 l4 4 \4 14-14 i4

Tail 66 62-7] 68 53-62 57

Spieule 22 23-25 24.5 - -

De Man's a ay 27-3 29 22-24 23
J b 3.1 3.8 - 4.4 4.) 4,.2-3.5 a4
us c 648 66-77 TA 71-84 AB
” a Aq 44-51 48 38-44 41
" V% - - - Ju-7) 7!

—_

NEW XYALIDAE 59

32 33

\ ae

_—

, 4

x ae

ee 31,32 25 um
a 33,35 25 um

34 50 nm

Figs 31-35. Prorhynchonema gourbaultae sp. nov, 31. male head. 32. female head. 33. inversion of annules. 34. entire female.

35. male tail and spicules.

60 WARWICK L. NICHOLAS & AIMGRN C, STEWART

Parutypes: Female paratypes resemble male holotype.
apart from reproductive organs, but amphids slightly
larger, 2.7-3.5 ym cf 2.6-3 yim. Single anterior ovary,
vulva simple not cuticularised, no operculum.

Habitat

Intertidal sandy beaches.
Distribution

New South Wales.
Differential diagnosis

The new species differs from the other described
species, 2 warwicki Gourbault, 1982, by having very
different spicules and a longer cervical region and
buccal tube. The relative length of the buccal tube is
similarto R. brevituba Gerlach, 1953 but the spicules
ure different and in R. gerlachi annulation does not
show. inversion,

Prorhynchonema warwicki Gourbault, 1982.
Material examined

Three males and 3 females from Rosedale and Tomakin
beaches, New South Wales.

New subfamily Corononeminae

With characters of tamily, Buccal cavily cylindrical,
deep and wide, not enclosed by pharyngeal
musculature; amphid situated above base of buccul
cavity, Cervical region only slightly attenuated. Six
inner labial setae in one ring. six outer labial and four
cephalic setae in second ring. Strong cuticular
annulation begins at base of buccal region.

TABLE 5. Measurements ef Corononema parvum sp. nov.

Genus Carononema gen. nov
Type species: Corononema parvunt
Diagnostic definition

Cuticle strongly annulated with cight longitudinal
ridges. Lips high, incised. Deep groove around the
head just posterior to the insertion of six inner labral
setae, in [ront of outer labial and cephalic setae, sets
off the lip region from the slightly tapered buccal
region.

Frymoalogy

Corona, Latin for crown, because the head appears
to be crowned by the incised lips, set off from the
cylindrical buecal region by a deep groove,

Coranonema parvum sp. nov.
(FIGS 36-40, 50-53)

flolotype. Male, SAMA V4263, Rapid Creek beach.
NT, 31vii. 1986,

Faratypes, 6 males, 5 AHC 24823a-c, Rapid Creck
beach. NT, 31.vii.J986; | male, AHC 25824, Southport
beach, Qld; 7 females, 6, AHC 24823, Rapid Creck
beach, NT, 31.vii, 1986; 1, AHC 24824, Southpert
beach, Qld, 31.vii.1986,

Measurements: Table 5
Description of Haletype male.

Cuticle strongly annulated, annules LS jm, with
eight equidistant longitudinal ridges, weakly developed
in cervical region, pronounced mid-body, which is
polygonal in cross section, extending almost to blunt

Type Holo
Male Range
Length RSS BON = [045
Width 27 22-26
Annulition 1.6 |.5-2
Lip height 3,5 45.6
Outer labial setac 5 4-6
Sub-eephalie setae \2 \}- 19
Body setae if 13-19
Amphid 5 4.48
Buecal cavity, length {3 Hh 46
Bovedl cavily, width 9 R10
Mouth t nerve ring 97 90 - 120
Pharynx 322 285- 360
Mouth to vulva - . 2.
Mouth to anus FRI 714.473
Width at anus 2) 19.23
Tail y4 72-87
Spicule, are 2 25-31
De Man's a 32 32-44
" h 7 2.6 - 3.6
ml c LLG O34 - 14a
by c a5 33-45
M V% - :

Male Paratypes n= 6

Female Paratypes n = 6

Mean + SD Range Mean + SD
9 i) 840 - 1084 977 71
24 «1.46 22 - 38 28 4.3
18 O19 1.7-2 19 (12
34 059 4-55 5 050
46 (89 6-9 6.9 1.0L
IS.4 0 2.97 {- 12 ia 1.00
16.2 2.39 95-2) a5 499
46 0.40 44-6 3.20 O82
I3.2 0 1.97 13-18 5.7 1.80
9 0.63 8-11 9.9 95
1o0 \] LNQ~ | 32 107 13
4 30 310-380 452 26
> ° ON7 ~ ¥70 799 63
843 = 107 TH - 975 900 74
2t 133 20-25 22 (A,27
79 5.92 68 - 105 80 12
278 2.56 - - -
7 SOI 26,0 - 40.0 33.0 At
29 0.36 2.4-3.1 24 0.27
1G 204 OB - 15.2 |2-4 191
AR 0.47 320 - 5,30 3,80 O7t

= : 80-84 $I

NEW XYALIDAE 6]

36 ——_———_______ 25 um Pe
3g: —_——_—_———__. 25 1m hy

37 —&- cm — 100 um

38 ——————————_ 50 um

40 ————————- 100 um aye

\ \
Trt
prvi nitl

yy vi

pysa prey

VUVHPVIV PET py yyy

Vii

Figs 36-40, Corononema parvum sp. nov, 36, male head. 37. head and pharyngeal region. 38. male tail. 39. spicules. 40.
entire female.

62 WARWICK L, NICHOLAS & ALMORN C, STEWART

cylindrical tip of tail, Cervical region enclosing buccal
cavity nol annulated or ridged. slightly amenuated
rowurds mouth. Deep groove, with less strong cuticle,
surrounds buccal region just below the lips, Uwe
scalloped cuticular rings: lie just within mouth. Six

incised leal-like lips, six inner labial setae al base of

hips. six outer labial and four cephalic setae insert
posterior torhe groove: four strong cervical setae, |2
pen insert at base of nen-annulated buccal vegion on
first annule, numerous body seiae, amphid fovea
situated over base of buccal cavity, 5 wm diameter, 25%
head width. Wide, deep buccal cavity with strong
almost parallel-sided walls. Cylindrical pharynx, Two
outstretched, inactive testes: spicules cephalated,
rectangular curvature, simple pointed tips;
gubernaculum surrounds spicule tips.

Paratypes: The spicules ditter somewhat in the degree
of curvature and may appear different because of partial
rotation about their axes and the pointed lips may be
turned ouiwards. Females. apart from the reproductive
organs. closely resemble males, Females possess a
single anterior gonad; the vulva has no operculum. The
labial and cephalic setae are difficult ro measure by
light microscopy because of their small size, but from
scanning electron microscopy the inner labials are
about 1.2 pm tong, the outer labials about 0.5 um and
the cephalic about 2 wm long.

Differential diagnosis

The deep parallel-sided buccal cavity, without teeth,
with lour strong setae at the base is distinctive, The
indented weakly cuticular groove below the insertion
of the lips is anlike that found in other Xyalidae.

TABLE 6. Measurements af Corononema thai sp. nev.

Habitat
Sandy ocean beaches,

Distribution
Tropical and sub-tropical beaches in the Northern
Territory and Queensland. Australia.

Corononema thai sp. noy,
(FIGS 41-49, 54-56)

Holotype: Male. SAMA YV4264. Pathaya. beach,
Chonburii, Thailand, 30.ix.1985.

Pararypes: 4 males, AHC 24825, und 7 females, ABC
24826 Pathaya beach, Chonburii. Thailand, 30.ix 1985.

Measurements; Table 6
Description of Holotype mate

Strongly annulated cuticle, anmules. 1.6 jan wide, 8
longitudinal ridges from buccal region almost to (ip
of tal, Buccal region not attenuated, only slightly
widening [rom mouth to base, enclosing deep, wide
unarmed buceal cavity. Lips high, leaf-like, fold over
wide mouth, Lip region sepanuted from buccal region
by deeply indented weakly cuticular circum-oral ring
sandwiched between two circum-oral erenellated
cuticular rings. Six inner labial setae inserted at base
of lips. $1X outer labial and four cephalic setae inserted
below indented ring. Long thin body setae spaced along
bady, six cervieal setae inserted on first anoule at base
ol buceal region, Cylindrical pharynx. Two
outstretched testes. anterior to left of gut, posterior lo
right. spicules cephalated, rectangular curvature,
simple pointed tips; gubernaculum surrounds spicule
Ups,

Type Hole

Male Runge
Lenuth 9900) 642 - 95%
Width 27 25-24
Annulation 1.6 16-17
Lip height 44 4-5
Outer labial sete 6 5-8
Sub-cephalle setae 4 yold
Body setie 3 l0-16

|
|

Amphid + 7 -5.3
ie

Buceul cavity. length 2 D168
Buceal eavity. width 2 K-11
Mouth to umphid 7 6-15
Mouth to nerve ring 76 8a - 118
Pharyns 264 24) - JKR
Mouth to vulva - - "=
Mouth to urus 19 AR - RIS
Width at anus 2a 20 «24
Tail 7\ Hl -6%
Spicule, ure 32 2h - 32
De Man's a an 24.0 - 384
" b An 2.5 -F.5
ls c 149 Ws IS2
4 c 0.4 2.6 3.)
fi Vy

Mule Parutypes ne 4

Female Purdivpes p= 6

Mean + Sd Runge Meun + SD
74 138 TO LOK 946 106
21 1,29 a) - 35 32 1st
a) 0.05 14-14 1 OS
ti DAS 5-75 7 At
64 144 h- 10 94 |.o3
\2 2.50 1O- 12 " 1
has «2.50 Ihe 14 12.4 152
46 O75 4-6 340 OH
125 2,89 1-13 1.7 (2
Ya 800 W- 12 \! ttl
1 3.74 3-74 58 LAT
O5 16 7-42 h2 OMI
262 21 2K5 - 424 Alb SQ
- m0- 871 TOR Kou
OM] 13h 722 - O8k 858 1M)
i V7 22 - 26 24.2 143
th 2.99 on = TOT BO \3

2s 263 isp

28.00 6.46 23,0 - 34,0 293 393
25 (AS 20-34 3.0) O34
HY 228 YO - 13.3 1.2 1.74
aN 24 1,10 ~ 4.00 60 Oda
- nO - B+ at 1.60

NEW XYALIDAE

feames varia n
“e fy Sie 2,8 ‘ »

p~ C—O

43, ——_—————_ 100 pm

42 ———————. 50 jim

Figs 41-44, Corononema thai sp. nov. 41. male head. 42. male tail. 43. head and pharyngeal region. 44. spicules.

64 WARWICK L. NICHOLAS & AIMORN C, STEWART

Figs 45-49. Corononema thai sp. noy. 45. cross section drawn from fractured specimen viewed by SEM. 46. spicules in
ventral view. 47. female posterior end. 48, annulation by SEM successively near head, mid-body and near tail. 49. entire
female.

NEW XYALIDAE 65

30 pim

Figs 50-56. Corononema yen. noy. by SEM, 50-53. C. parvum sp. nov. 54-56. C. thai sp. nov. AM amphid, LP lips, ILS
inner labial setae, OLS/CeS outer labial and cephalic setae, SCS sub-cephalic setae, SP spicules, VL vulva, AN anus

66 WARWICK L_ NICAOLAS & AIMORN C, STEWART

Paratype. Female paratypes possess a single anterior
ovary, vulva without operculum, prominent post-vulyal
hand. otherwise females closely resemble males.

Differential diaynesis

© thai sp. nov. diflers From C. parva sp.nov: by
possessing a shorter broader head, Asan index of this
ditlerence the ratio of length to breadth of the pre-
annulated buccal region has been measured. In C.
parviin the ratio ranged from 0.87 te 104 (=30), in
C. that trom 0.64 to O75 (n=10), The annulation is
shallower in C. thai. Both these properties are most
vlearly seen in scanning electron micrographs of the
hesel (Fig. 52).

Habvitert
Tropical sandy beaches.
Distribution

Vhuiland.

Discussion

The taxonomic position and rack of the
Corononeminac presents difficulues. The circular
eryptospiral amphids and single prodelphic female
gonad are characteristic of Nyalidae. The strongly
annulated cuticle and buceal cavity lacking teeth are
also-shared with most Xyalidae, although some possess
toott-like ridges in the buecal cavily, The deep
unarmoured buccal cavity with strong cylindrical walls
is found in other Xyalidae. such as Omicranemtee and
all Rhynechonemutinae. but Whereas in
Rhynchonemutinde the buccal cavity terminates as a
narrow tube of varying length, the buecal cavity of
Crrenoneme is relalively wide and short. The cephalic
region isattenuated.. but not to the marked degree so
chargoteristic of Rhynchonematinae. For these reasons
the new genus, Caroyionemer, 18 best placed within the

Xyulidae, but would stretch the definition of the nwa
previously recozmsed subfamilies too far so that the
erection of d new subtamily seems warranted,
Lorenzen (1978) in tis review of the supertiumily
Monhysteroiea does not recognise any subfamilies
within the Xyalidae. noting, in this regard, that essential
aspects of the phylogenetic relationships within the
Monhystepojdea have not been cleared up. We do not
find this sufficient reason to abandon the previously
recognised division of the Xyalidae into two clearly
separable subfamilies, namely the Cobbinue and
Rhynchonematinae. Whether we are justified in
creating another subfamily for two species with
intermediate characters must be a matter of personal
judgement. In Coronanenia the form of the cervical
region and buccal cavity are intermediate between that
tound Rhynchonematinae from Cobbiinue, but there
is.no overlap.and clear diflerences remain between the
three taxa. The deeply incised labial region and a
flexible indented ring separating the cephalic region
from the buceal region ure mm our view significint
distinguishing attributes of the Corononeminae. Tt may
well be when the very poorly known nematode faunae
of Australia and South East Asia ure betier known more
species will be found assignable to (he subfamily.
The indented cireum-oral ring is probably flexible
und may facilitate the mgestion of larger paruicles than
would otherwise be possible. A flexible oral region
has been Observed to lacilitate the ingestion of relatively
large diatoms by other Xyalidae such as Dapronema-

Acknowledgments

We are grateful for a grant from the Australian
Biological Survey which made this work possible. We
thank Dr Russell Hanley, Northern Territory Museum
of Arts and Sciences for making it possible lor us po
collect on Darwin beaches,

References

Geriack. S. A. (1953) Die Nematodenbesiedlung des
Sandsttandes an des Kiistengrundwassers an der
iluhenischen Kliste. J-Systematischer Teil, arehe. Zo,
fal 37, 517-640,

GOURMALILT, §. (1982) Nematodes marins de Guadeloupe.
I, Xyalidae nouveau des genres Rhynchonemea Cobb ol
Provunchneme nov. gen Bull, Muy natn. Hist Nat, Paris
HW ser, AL Sec, AL tas. 1-2 75-87.

Hopper, BE. 961) Marine nematodes trom the coastline
of the Gultal Mexico. Can. Zaol, 39, 359 365.

Lokisven. oS (1971) Die Nematodentauna im
Verklappungseebeit fir Industnieabwasser norehwestlieh von

Heigoland. |. Araeolainida und Monhysterida. Zod/, Ane
187, 223-248,

(TY75) Rhynchynema -Arien (Nematodes,
Monhysterutae) aus Sidamerika und Europu. Mrcrafdiya
des Meereybodeny $5, 225-251.

_—____ W978) The system of the Monhysterdides (nerialodesy
- A New Approach. Zant, Jb. Svs. 105, 515-536.

STEWART, AC) & NieHolas, WL, (1994) New species of
Xvalidge | Nemutoda: Monhysteridy) from Ausiralian ocean
beaches, Invert, Texan, 8, 91-15,

Viigo, PL 1967) Deaux nouvelles espetes du venre
Rhynchonema (Nemittoda, Monhysteridae), Bull, Sac, cel,
Fe 92, Al3-121.

A NEW GENUS AND SPECIES OF CRANGONYCTOID AMPHIPOD
(CRUSTACEA) FROM WESTERN AUSTRALIAN FRESH
WATERS

By J. H. BRADBURY* & W. D. WILLIAMS*

Summary

Bradbury, J. H. & Williams, W. D. (1995) A new genus of crangonyctoid amphipod
(Crustacea) from Western Australian fresh waters. Trans. R. Soc. S. Aust. 119(2), 67-
74, 31 May, 1995.

A new genus of crangonyctoid amphipod (Crustacea) from Western Australian fresh
waters, Totgammarus, with a single species, T. eximius, is described. The species was
collected from roadside pools in the south-west of Western Australia.

Key Words: Amphipoda, Totgammarus, Western Australia, crangonyctoid,
Paramelitidae.

Transactions of the Reval Soerery of S. Ause (W9S), WH2). @T74

A NEW GENUS AND SPECIES OF CRANGONYCTOLD AMPHIPOD (CRUSTACEA)
FROM WESTERN AUSTRALIAN FRESH WATERS

by J. H. BRADRURY*® & W. D. WILLIAMS*

Summary

Reapnory. 1 H& Witaws, W. BD, (1995) A new genus of crangonycloid amphipod (Crustacea) from Western
Australian fresh waters, Tras Ro Sec. 8S. dust. 92), O774, SL May, 1995.

A new genus of crangonyetoid armphiphod (Crustacea) from Western Australian fresh waters. Jorguninaras ,
wilh a Siigle species, Fo cranius, 1s described) The species was onllected from roadside pools in the south-west

if Western Australia

Kiy WoRxps: Amphipoda. Jorgiminaris, Western Australia. cranganyeioid, Paramelilidae.

Introduction

All known crangonyctoid species of Australian fresh
witlers [9 IY87 were comprehensively reviewed by
Williams & Barnard (1988). In their review, knowh
species were re-examined and redescribed, and some
new species were described. They dealt with a total
of 12 genera and 33 species. They considered the
number of genera probably represented about half the
number expected to oecur in Australia and noted that
the number of species within genera was probably
small. Purther work (Barnard & Williams, in press)
supports this view; they deseribed two new genera.

euch monotypic, ay well as a further new species of

both Ansrogammaras and Uractena. This second
review by Barnard & Williams (in press) described,
ier alia, most material available to them from Western
Australia, They did not deseribe, however, a taxon from
thal State represented by only a single specimen,
pending the collection of further material,
Unfortunately, all attempts to obtain more specimens
hive proven unsuccessful; exhaustive exammation of
all the known collections from the area have yielded
no further specimens and nor did collections made in
1994 on our behalf by A. J. Boulton.

Since the single available specimen represents in our
view u new genus, and to Jacilitne further studies of
freshwater amphipods in Western Australia in
particular and Australia in general. we now consider
it appropriate to describe this single specimen.

Methods oF dissection, description and potation
follow those of Williams & Barnard (1988). To expedite
the use of figures in the present publication, the
abbreviutions are as follows; “Antennal sinus” refers
wnly to the cephalic sings receiving umtenna 2. A -
iumcnna Abd - abdomen; ace - acteessory; art -article;
C - coxa, cox - coxalsd - dorsal; dact - dactylus; @ -
eye; E- epimeron, fag - flagellum; g - gill; G -

gnathopod: Hd - head) i- inner, | - lefts lac - tacinia
miobilis; lat - lateral; LL - lower lip: MD - mandible;
micd — medial; mol - molar: MP -maxilhped; MX -
maxilla; o- outer; O- oostegie, p-palp: P -
percopod: PC - prebueeal complex; pl - plate: post -
posterior, Pp - pleopod: r - right; ret -retinaculum:
st - sternal; 'T - telson; U - uropod: UL -upper lip: v -
ventral; 1,2,3...7 - first, second, third. seventh,

Genus Jotgammarus gen, nov,
Eymtulogy

Named for the combination of features of several
gener,

Type species: Totgenemerks extanus

Diagnosis

Pleon with sparse dorsal setation, pastrum weak
Lateral cephalic lobes strongly. proyecting, antennal
sinus moderate. eye not discernible in preserved
specimen. Flagellum of first antenna lacking mayor
armainents, moderately long. about 05x body length,
iwive A2, Ratio of peduncular articles 2;2:1, Flagellum
of second untenna and peduncle of sub-equal length,
culecolt of type 9 present (Lincoln & Hurley 1981).

Ratio of mandibular pulp urticles abour 2:9:6, article
2 moderately Setose. arucle 3 faleute. setae = ABDE
(Barnard & Barnard 1983). Lubium lacking inner
lobes, Maxillaeé 1-2 medially setose, inner plate
wholly or marginally pubescent, Maxilla | outer plate
ovate. medially ind laterally setose, palps asymmetric:
left with thin apical spines. right wilh thick apical
spines. Maxilla 2 inner plate with row of apico-medial
weakly sub-muarginal setal spines. medial margin
heavily setose. Muxillipedal palp arucles 2 ~3 with
few lateral setae. article 3 with fine factal pubescence
dorsally and a ventre-facial row of moderately long.
curved setac.

O8 1H, BRABBURY & WD. WILLIAMS

Coxue 1- 4 willha cow of postenor spines. coxac
) 4-moderitely elongate. coxa (tapering below, cox
d deeply emurginate post-dorsally, coxue 5 - 6 shorter
thin 4, coxa 7 shorter than 5 - 6, Gnathopods unequal,
soathopod | O.5x gnathopod 2. Carpus ol gnathapod
| long, of gnathopod 2 short. Scythe spine absent from
artele + of both gnathopous. each with a weak lobe,
Propodus of both grathopeds rectangulir, palins
weakly to moderately corivex, palmer corners
prominent, lurned oul. First gaathopod lacking stron
spines at palmar corner, second with G strony spines.
Spines along palms of both gnathopods short, simple,
without triggers, numerous.

Perenpods 4 - 4owith posterior spifie sets on article
6 evenly spaced, PS - 7 moderately elynwute, urticle
2 broudly expanded, lobate Dueryls with | -3
sprites.

Coxac 2 - 7 with sav-like gills. Thoracic segments
2-7 with dateral steraal gilts.

Basumedial selae on inner rami al pleapods 1 - 3
plumuse, peduncles each with paired retingcula and
paired (Hirst and seeond pleopod) or single (hind
pleopod), plumose kecessory retinacula,

Pleomtes With few dorsal setae and/or spines,
Epimera with few ventral spines. posteriar qmargins
Weakly setulite. Rami of uropods | - 2 extending sub-
equally, each with 2 rowe of spines. Facial armament
of Uropod Tweak, largely absent on uropod 2 which
hears u strong, elongate upteo-medial spine Liropad
J extended, magninumous, peduncle shart, ouler ramus
2 aniculine. artiele 2 short,

Telson longer than broad, 100% eleft, not laverally
turin, apically and disto-laterully weakly selose,
bearing a single sub-apical spine on either lobe

Additional déseription

Flagellum of Al-2 lacking major armaments.
Apical thargin of labrum. extended. Accessory blades
(rakers) on mandibles with inter-raker plumase setae
interspersed among rakers und additional Short plumose
4elle Iving between rakers and molar Matar trilurative,
with plimose apical seta, Mandibular palp article 3
shorter than 2, palpurticle 2 Jacking baso-anterior setae
with few median and apico-anierior setae. Both plates
Of second maxillae with rows af long distal setae
Maxillipedal palp inoderately long. Article 3 weakly
produecd and finely pubescent at the upex which bears
lone terminal setae; baso-medially hearing a sirle sub-
marginal seta, medially Wilh a row of scythe selae
extending to the base of the duetyly selae of [he venirul
Jace constituting a comb row, aswell as u single long
mid-lacial sela and 4 naw of shert setae basal te the
comb row: jhe mid-distal dorsal face beanny fine
pubescence

DactyLof first gnathopod not reaching palmur comer,

bearing a small, bent tiner tuoth-spine. Bucry) ut
second ynuthopod reaching i end of palm, bearing
Jsmallinner spines, Peeeopod 7 shorter thin 6. Article
2 of pereapods 5 - 7 equally setose pasteriorly.

Sternal processes: fleshy sausage-shuped pills on
thoracic segments 27, aniehed to mid-lateral argeins
of segments,

Puslero-ventral apex of epimera | - 3 blunt. as in
Ausirogaminaras, Pleopods similar, except for numbers
of retinaculae, rami approximately equal. Outer cunius
of urupods |~ 2 slightly shorter than jnner ramus.
Apidolateral corner of peduncle of uropod UL wilt 2
Spines, ram of both first and second uropods with >
upical spines. Phe third weopod extending beyond the
first and second in the entire animal, pedunenlar spines.
apical. and Sub-apical, some medial sete of each taitius
Plumose. Ventradistal spine on urosumite 1 ur base pl
drypod | short, as in Auatragennaerin,

Relarlonship

This genus displays the eharicteristies ul
crangonyetoid amphipods in possessing (Ly) sternal gills
(2) un accessory Magellun of the first antenna with two
or more drticles (3) ealeeoli of type 9 or linear (4)
uropod | licking a basolacial spine on the pedunele
(5) a lower lip without ianer lobes (6) a frst ynathopod
thitt is not melita or mittenform in shape (7) 4 first
enathopod that does not dominate the second, and (4)
a inandibular palp of typical form (Williams & Barnard
JORK).

The new genus fits the essential criteria of the family
Paramelitidue in possessing sausage-like sternal pills,
dorsal setae on the telson, and linear or type-9 calceo|).
VW differs front the Neoniphargidae in the ubsence of
rugosities on the third article of the maxillipedal palp
and gnathopeds. the form of the gnathopods (not srl!
und mittentorm), the form of the carpi (not short and
lobate), and non-dendritie or luip bearing sternal gills.
It differs from the Perthiidae in thar the first antenna
is significantly longer than the second. the mandibular
molar is normally developed and triturative. rhe outer
plate oF the maxilliped is nor very small die
enathopods are not large. nor are the carpi short and
deeply lobate. the curpi and propodi are not eusirid,
dnd the sternal gills are pet dendritic,

Joteannarns. bears atures in Common with alter
paramelitid geuera, such as blindness, which occurs
ie several, and m possession of an eclangate spine on
fhe secand male uropod (as in-some Urvetend spp.)
The combination of characteristies however, is uniguc.
The genus varies from Austrogamtimarus, the most
primuuve Australian paramelitid genus, in-several ways,
In Jorgaminarns, dorsal setation of the pléonites: is
weak, the lateral cephalic lobes project strongly, the
antennal sims is Moderate, eyes ure absent, and the

A NEW GENUS AND SPECIES OF CRANGONYCTOID AMPHIPOD 69

Fig. 1. Totgummarus eximius, sp. nov. holotype, male 10.6 mm. Whole animal, antennae and mouthparts, Scale bars: adult
and antennae = | mm, mouthparts = 200 pam.

WW | H. BRADBURY & W, D. WILLIAMS

second urlicle af the mandibular pulp 1s relatively long
will lew wpieo-unterion and no baso-anterior sete,
Additionally. there is a extension of the apical margin
of the labrun, 5 rather than 3 apical spines occur on
the wher plate oF the maxilliped, cosae 1-4 are
mderdely long ratber than elongule, the apex of coxa
| Lapers, pereopods 5 - 7 are more even in length, and
a seythe spre iy hol present on article four of the
gnuthopods although a small lobe js present, The
daetyls of legs 3 - 7 are mulliespinose, The peduncles
(the pleopods are moderately setose. the apieo-lateral
and apico-medial spines of peduncles of the Sivst
uropod differ, usdo the relative lenpths of the urepod
vam which also bear busu-ficial armaments. The third
Uropod is miagniranois,

Toteainmaruy eximiuy sp. nov,
FIGS | - 3.

Pryvinelaes!
From eviniay. mieaning exceptional or alone.

dype hacetiry
lomporary roudside water in sands along the Scott
River Road. south-western Western Australpt

Dern
With the charaateristies ab the gens (only trate
Known,

Material Evanined Hatoiwpe

Western Australian Museum WAMI4-95. male 10.6
Min in rype series,
Nooather specunens available.

Beseripiton uf halvivpe (male)

Body (Fig. t), pleanites 3 = 6 with Sparse transverse
dorsal selalion and dorso-laieral spines on 3 - 6,

firstamtenna (Pig, 1) primary Oagetlim sparsely
s¢ioxe, Mugellunt of 35 articles. 18 x pedunele. No
culeeoli, Accessory flayellunm 7 - 8 articuliiie. reachine,
teanicle & of (he priury flagellum. Sccond antenna
(hig. Ves Jength 0.25 x beady length, pedunecular articles
4S subequal, Magellum of 19 articles, setae sparse.
Calecol) on urticles 1-13. Labrum (Fie 1: broadly
rounded with apex slightly extended. harerully and
apically pilose, Labium (Fig. 0: medially and laterally
pilose with (0 vurved upico-medial spines on either
lobe

heh mandible trig. Ue palp article 2 senjtien
1A 2B - 15D - AL. article 2 with 2 Inedial »etic and
Ob)QUe PAW EO apico-imedial setae, Incisor 6-tuelhed,
lheinia mobilis +-roothed, 9 setose accessory: blades.
3 short plinose sete und | short bluntspine taward
baseot molar, Anterior of molar densely pilise. Motar
with) shork plumose seta. Right mandible (Pu 4.

incisor 4-toothed, Jacinia mobilis bifid with 4
dentioukite weth on one side and 9 cuspate teeth and

al blunt terminal tooth on the other, accessory blades

of 3 toothed spines and 4 sctose inler-rakers, 3 short
plumoseselae and | narrow blunt spine toward the buse
ofthe molar, Molar with | sctose median, short. blunt
Spine, and long plumose seta,

Lefl first maxilla (Fig, 1): palp-article 2 with 10 thin
upigal spines, otherwise naked, Outer plate medially
setose. JO denteulate terminal spines, Inner plate ovate,
laterally and medially with sparse straw-like
pubescence, 5 upice-medial plumose setae, Right first
maxilla (Pig, Ds palp article 2 with 6 thick apival
spines. L disto-laterul moderately long spine und |
luteral Sub-upical curved spine. Outer plate with 10
denticulute terminal spines abd 1 aatero-medial
plurimose seta, Media area Witt long pubescence, Inner
plate ws for left side.

second maxilie (Pig. 1) syinnietrical, outer plate
laterally setose, sub-terminal row of (0 curved spines.
fertninal row ef many curved setae. Inner plate laterally
seluse, SUb- imirginally pubescent; medial margin with
fine sete proximally, row af setal spines distally.

Masilliped (Pig. 1): palp article 3 with 9 inedial
sevihe setae, l6-antero-frctal comb row see extending
from MO4 to the sub-upex, 4 long terminal setae, 2
nid lateral setae and 1 median sew. Dacty! bearing
[distil and | oanedial accessory spines and, dorsally,
4 short post-lacial setae basal to the comb row: outer
plate laterally sctose. apically bearing sub- terminal
row of 8 strong curved spines, a disto-medial sub-facial
row of [O Tooth spines and 13 setae, selue distally sub-
facial fo the teeth, proximally facial, Inner pliite
terminating in 5 strony tooth spines and & plumose
selves medially 6 long setal spines, the distal 4
plumose: basal to the inner plate g transverse row of
% mediviy to long naked setae,

First ynathopod (Fig. 2). coxa tuypered, 3 posteriar
spines, weakly setose marginally: carpus moderately
long. sub-equal to propadus, not lobed: propodus
reclangulir, pulmar corner prominent, extended
posteriorly, pal ucute, convex, daety! reaching cornet
of palm, bearing small bent yner tooth, 5 phumose
und | naked spines al palmer enrner: numerous short
spines along palin

Second gnathopod (Fig. 2)! larger than the first (Lett
2x, Right 1-78); coxa wilh row of 4 or 5 posterior
spines, few snail distal setae; Carpus short, about 0.5%
propodus: propodus reetanmular, longer than wide,
palmar corner with 2 strong, naked spines and 4
pluniose spines. corner prominent and slightly
extended posteriorly: dacty! not reaehing corner, but
reaehing tothe second naked spine. bearimy 2 inner
teeth at approximarely MOS; palin slightly convex with
HUMETOUS SHORE Spins

A NEW GENUS AND SPECIES OF CRANGONYCTOID AMPHIPOD 7)

Fig. 2. Totgummarus eximius, sp. nov. holotype, male 10.6 mm, Gnathopods and pereopods. Scale bars: gnathopods and
pereopods = | mm. dactylar enlargements = 200 pm.

72 J, H, BRADBURY & W. D. WILLIAMS

—

a a

Fig. 3. Jotgammarus eximius, sp. nov, holotype, male 10.6 mm. Pleopods, uropods, telson, gills and abdomen, Scale bars:
abdomen and gills = 1 mm, pleopods. uropods and telson = 200 pm. retinaculae = 50 pm,

A SRW GENUS AND SPECIES OF CRANGONYCTIOD AMPHIPOD Va

Pereopods (Fig. 2): coxa 3 with 9 posterior spines,
cox + deeply emarginate, small setae and spmes below.
no posterior spines. Coxa 5 bearing 3 posterior ventral
spines, coxa O with 3 posterior ventral spines and 4
small posterior setules. Coxa 7 with 4 posterior spines
Pereopods 3 - 4 length 1.2 x G2, subequal, article 2
ol both bearing long posterior setae, Article 5 of P3
apicu-posterior spine formula (proximal to distal):
|-2-2-2 article 6) 1-3-3 -3-2-2-2 article
Sof PA, 2-3 - 3- 4,artivle6;3-3-3-3-3-3 3.
Pereopods 5. 7 of approximately equal length,
Pereopod 5 articles 3 - 6 bearing long apico- posterior
selue, upico-anterior spine formulae 2-3 ~2 and
3-4-4-4-3-3-4-4. Pereopod 6 article §
bearing few long setae, article 6 many apico-pasterior
setae. spine formulae 4 - 3-4-0 und 2-4-4 -5 -
() - ~3 respectively, Pereopod. 7 upico-untenor spine
formulde: article 5S! 4-6-6 and article 6:
a-3-4-4-4-3,

Gills (Pig.3)> coxall gill 5 slightly reduced, gills 5 -7
bi-lobed. Slernal gills 2-7 lateral,

Epimera (Fig. 3): with few ventro-facial spines. -

posterior margins with few small setules. Epimeron
| slightly rounded posteriorly with single antero-ventral
spine, Epimeron 2 with 3 srnall mid-ventral setae only.
Epimeron. 3. naked ventrally.

Pleon (Fig. 3): pleonies 3 - 6 with dorsal spines
and/or sétac. Pleonite 5 with 5 spines in transverse
groups of 2 and 3, Urosomite 6 with 1 dorsal spine
on either side,

Pleopods! pleopods | and 2 bearing paired, hooked
retinaculae and paired aveessory retinaculae, pleopod
Slacking second accessory retinacula, Uropods (Fig.
3): lirst uropod; peduncle length [2 x ramt, outer
margin with | upico-facial spine, 2 medial spines, and
strong row of 5 dorsal spines, without setae. Ram sub-
equal, terminating in a cluster of S spines. Second
roped; peduncle length equal to rami, lacking spine
rows, bub-with a cluster OF T large and 4 short apico-

facial spines. Inner medial angle with elongate spine
0.5 x length of peduncle. terminally spoon shaped.
Inner ramus 1,3 x length of outer, lacking setae. Both
tami terminating ina cluster of 5 spines. Third uropod:
peduncle iength 0.35 x Jength of outer ramus, about
the same length as. urosontite 3, bearing median
transverse row of 5 spines. distal transverse row of 7
spines at the base of the outer ramus and u group of
4 apico-lateral spines, Outer ramus proximal article
strongly setose baso-laterully with 4 disto-lateral
clusters. of spines and setae, paired medial und single
Jateral trigger spines apieally, medially a single sub-
apical trigger spine and evenly spaced plumose selue.
Small distal article, 0.13 x proximal, terminating in 3
short and 2 long setae, Inner ramus of 4 single article,
equal to the length of the proximal article of the outer
ramus, marginally setose. the medial setae plumose,
6 lateral and 5 medial trigger spines distally; 2 terminal
spines and 4 setae.

Telson (Fig. 3): 1.25 x urosomite 3, cleft 100% -
Disto-lateral margins und apex. with sparse dorsal
setauion, paired penicillate setules sub-muarginal al
M.80, Single sub-apical spime on either lobe,

Distribution

Western Australia (south west), Scott River Road,
sands in a roadside ditch coll, K. Davies, B. Knott,
03 Oct. OBL.

Acknowledgments

The authors wish to thank Dr A. J. Boulton
(Universily of New England, NSW) lor his efforts to
collect further samples of the species in January and
February 1994. A.B.R.S. support during the
finalization of the manuscript is gratefully
acknowledged.

References

Bakar, tol & Barsarp, C, M, (1983) “Preshwarer
amphipndi: of the world. 1, Evolutionary. patterns, IL.
Handbook wd bibliagraphy” (Mayfield Associates. Mt.
Vernon, Virginia.

— & Wiiatis, W. Do fin press) The wixonomy of
Preshwatsr Amphipoda (Crustacea) from Australian Fresh
Waters Part 2. Ree. Aut. Mus. suppl.

Lincorn, R, 1o& Huguey, D, E, (981) The calecolus, a
sensory structure of gamimaridean amphipods (Amphipada:
Gammuridea). Bull Brit. Mus, Nar Hist, (Zool,) 40,
(03-116,

Wituiams, W. BD. & Barnaros. J, L. (1988) The taxonomy
of crangenyctoid Amphipoda (Crustacea) from Australian
Fresh Walters; foundation studies. Ree. Aust Mus, suppl
10, L180

ARROWIPORA FROMENSIS A NEW GENUS AND SPECIES OF
TABULATE-LIKE CORAL FROM THE EARLY CAMBRIAN
MOOROWIE FORMATION, FLINDERS RANGES,
SOUTH AUSTRALIA

By MARGARET K. FULLER*, & RICHARD J. F. JENKINS*

Summary

Fuller, M. K., & Jenkins, R. J. F. (1995) Arrowipora fromensis, a new genus and
species of tabulate-like coral from the Early Cambrian Moorowie Formation, Flinders
Ranges, South Australia. Trans. R. Soc. S. Aust. 119(2), 75-82, 31 May, 1995.

The recently discovered Early Cambrian tabulate-like coral Arrowipora fromensis
gen. et. sp. nov. occurs in the Moorowie Formation of the eastern Flinders Ranges. It
is found in an ancient reefal environment in association with Moorowipora
chamberensis Fuller & Jenkins 1994 and Flindersipora bowmanii Lafuste 1991.
Arrowipora fromensis has tabulate-like characteristics including the cerioid form of
the corallum, wedge-shaped to spine-like septa and strongly developed dissepiment-
like tabulae. Although unlike any other Early Cambrian coral, skeletal characteristics
are similar to some micheliniids, which have a time range from the Late Silurian to
the Late Permian. Arrowipora fromensis provides further evidence that the time range
of the Subclass Tabulata possibly extended to the Early Cambrian.

Key Words: Arrowipora fromensis, Early Cambrian, Moorowie Formation, tabulate-
like coral, Flinders Ranges, South Australia.

Transactions of the Raval Serer of 8, Aust (995), WA2), 15-82

ARROWIPORA FROMENSIS A NEW GENUS AND SPECIES OF TABULATE-LIKE CORAL
FROM THE EARLY CAMBRIAN MOOROWIE FORMATION, FLINDERS RANGES,
SOUTH AUSTRALIA

by MARGARET K. FULLER & RICHARD J. F. JENKINS®

Summary

FULLER. Mo K., & Jenkin. ROLE (1995) airrowiporen fromensis. a new penus and species of tibulate-like cor)
trom (he Burly Cambrien Moorowie Formation, Flinders Ranges, South Australia, Tran. R. Sac. 5. Anse, 19(2),
75-82. 31 Muy, 1995,

Ihe recently discovered Barly Carnbrian tabulate-like coral Arrawipord fromensis: gen. et sp. nov. Occurs ITT
ihe Moorowie Formation of the castern Flinders Ranges. tb ts found in ain ancient reebil environment in association
With Mooravipare Mamberensix Fuller & Jenkins 994 and Flindersipora bawntanil Latuste 199), Arrowrpard
fromenvis has whulate-like characteristics including the certo form ofthe corallum, wedge-shaped to spine-ke
septa and strongly developed dissepiment-lke tabulue, Although unlike any other Barly Carnbrian coral, skeletal
characteristics are silmikir to Some michelinnids, which have a tite range front the Late Silurian to the Late Permuan.
drrawiporea fromenyis provides further evidence that the time range of the Subclass Tabulata possibly extended
to the Barly Cambrian.

Key Worbs: drrenvipura fromensis, burly Cambriitn, Moorowie Formation. tabulate-like coral, Flinders
Ranges, Saute Australia.

Introduction

Arrowipora jromensiy gen. et sp. Noy. occurs in the
Early Cambrian Mouvrowie Formation in the eastern
Flinders Ranges of South Australia in association with
Moarawipora chambercasts Fuller & Jenkins 1994, and
Flindersipora bowmeni Latuste 199). tt is present in
slumped réefal blocks within a megabreecia at a. site
close to the disused Moorowie Mine (Fig. 1) described
in Fuller & Jenkins (1994). The corals. aré preserved
as upright coralla relative to bedding and clearly are
in lite position within individual slump blocks. They
occur in) association. with both fragmental and
engrusting remains of the caleimicrobes Kenulcts
Vologdin 1932, Girvanella Nicholson & Etheridge 1878
und Epiphvien Barneniin 1886, and current-deposiled
archaeocyaths. The ancient reefal systern may have been
established on a marginal fan comprising a coarse
breeeia (Savarese eral. 1993). The high energy murine
environment was responsible for the influxes of
sediment preserved within the framework of the coral
colonies. Arrewipyra fromens?s and the two previously
described corals from Moorowie have few skeletal
characteristics m common,

MIDDLE CAMBRIAN
WM Lower CAMBRIAN

Preservation

SE adtutaiaw
st

,
{

The available material, collected many years.ago by =QUORN Ny
Mr Brent Bowman, then a technical assistant al the

University of Adelaide, shows parts of probably one

Fig. 1. Location map showing fossil occurrence near the

* Departinemt of Geology and Geophysics, University of
Adelaide, South Australia 5005.

Moorowie Mine and the distribution of Early and Middle
Cambriin outcrops in the Flinders Ranges of South
Australia.

76 MARGARET K. FULLER & RICHARD J. F JENKINS

Fig. 2. Holotype SAM P34167 (complete specimen), illustrating rectangular shelves extending from a large colony (x1.0),

THE EARLY CAMBRIAN TABULATE-LIKE CORAL ARROWIPORA 17

Fig. 3. Holotype SAM P34167 (reverse side of specimen shown in Fip. 2) with shell-like projections across adjacent sediment
(x10).

is MARGARET K

coluny broken trom a large specimen (Pigs 2,3).
Duling lite the colony appears to have been repeatedly
hut partly covered by centimetre thick layers of Tine
sediment which now fill large spaces between lateral
expansions of the corallum. Many curallites: were
sinothered. allowing only wlimited number to continue
their growth, Subsequent corallites grew either inglined
or spread faterully above the tenses of sediment.
Transverse and oblique sections of small urchaeocyuths
lying on their sides relative to hedding are evident ip
eavities between extended shelves of the corallum (Fires
2,3}. The geapetal infilling of the urchaeucyaths lurther
indicates that (hey were transported into the cavities
with the sediment.

Culcureaus sediments filling small cavities hetween
the vorallites have generally been recrystallized, while
the calyces (lopether with larger cavities) wre usually
Mlled with very fine sand or silt, Laterally exvended
shelves of the specimen SAM P34167 are irregularly
reclingular or platy and project over the bioclastic
and/or culcarenite matin (Figs 2,3), Coralhites also
show unications of being eraded by rapid, energetic
influxes of course sand. Caleite-filled fractures appar-
endy related to post-diagenerie defornmution of the
corillim oceur rarely (Figs 4B,C)_

Reerystallization has affected wll of the colony and
sume of (he skeletal structures observed may be
artifacts of diagenesis, There are, however, domains
within the reerystullized fabric where sume evidence
ol the primary structure of the skeleton appears (a be
preserved, These relic, rather robust fihrous elements
which evidently formed the sclerenchyme (ealcarcous
skeleton of corallites), are seen as either lineations
across the walls of corallites (in transverse section)
{ind/or divergent bundles (in longitudinal section) (Figs
4&.5D).

In longitudinal section, upturned spines along some
corallite walls (Pig, $C). and spines situated on the
upper surtuce of some tabulae (Fig. 4D) are represented
by bundled tibres, giving both the wall and tabuiae a
bumpy appearance. [n transverse section, Most septa
uppear ty terminate in fan-shaped arrays of fibres, or
similar arrays arise from. the walls (Fig. SC). The
bundled fibres resemble primmlive trabeculae. However,
fan-shaped tufts in carbonates often result from
diugenesis (Oekentorp 1989).

Systematic palacontology

Phylum CNIDARLA
Class: ANTHOZOA
Subclass: 2TABULATA
Family: uncertain

Genus: Aprowipera gen_ nov,
‘Type speaes: Arrowipera Jronunaix sp, noy,

PULLER & RICHARD LP. JENKINS

Envriolony

For the Arrowie Basin, an Early Cambrian shallow
marine busin. extending over much of the area wl the
present Flinders Ranges of South Australia.

Digencyis

Corallum large. massive cerioid, comprisiny
polygonal corallites: corallites prismatic and irregularly
cylindrical, walls separated by a medial plane, thick.
wavy to crenitle. sometines almost straight; tibulie
numerous, rarely complete, commonly dissepiment-
like tabellaes septa nunieraus at absent, numbering up
to 35.in each corallite; where present septa form short
wedge- to spine-like projections into the lumens tinea
pores whsent,

Arrawipara fromensis sp. ny.
FIGS 2-5

Etvmalogy
Por nearby Lake Frome.

DiaENOsIS
As for genus,

Ape specimens; The specimens described in dis paper
ure held at the South Austrulian Museum (SAM),
Holotype SAM P34167, a polished skib of a broken
part of a corallum and thin sections SAM P34167 1.
SAM P34167-2, Puratype SAM P31962-1. SAM
P31962-2, counterparts comprising two triangular.
large, cut, polished slabs approximately 34 cm normal
to hedding and 28 em parallel to bedding. containing
either two coralla or more likely the disjunct parts af
one large corallum which formed numerous platy
shelves, Thin seetion SAM P34168-1. The material wus
collected from the Moorowie Formation, near the
Moorowie Mine in the eastern Flinders Ranges.( Fig. 1).

Description

Colony Jarge. more than 24 cm tall and extending
laterally Well in-excess of 23 em. In longitudinal section
the corallum may broaden upward, or more cormmenty,
forms wide shelves extending laterally over the adjacent
sediment, Shelves are either irregularly rectangular in
shape, with corallites tending to diverge slightly. or
are plate-like. Individual shelyes measure up to 70mm
high and {30 mm in width (Figs 2,3). The upper
surface of the shelves is irregularly horizontal to
concave, und culices may extend up to 7 mim above
the uppermost tithellae. In transverse section (Figs
5A,B,C), the ceriid corallites. are seen as 5-8
(generally 6) sided polygons, varying belween 10 and
l4 mm in diameter,

Walls ure relalively thick. varying between 0.1 min
and 1.0 mm, and are wavy to almost suuight. The inner
Surfaces of the wally are irregular, due ti the msertion

THE EARLY CAMBRIAN TABULATE-LIKE CORAL ARROWIPORA 79

Z wy ee

Fig. 4. Longitudinal sections of Holotype SAM P34167 A,B. Adjoining sections illustrating general shape of the corallites,
tabulae, vertical and basal corallite walls (x2.4). C. The irregular surface of the walls and upper surface of tabulae. Two
fractures which post date growth are observed mid-to lower-right of figure, together with the recrystallized fabric within
the corallite (x10.6). D, Enlarged section (x2.4) of corallite (lower right Fig. 4B) illustrating tabulae with possible septal
spinules on the upper surface. E. Higher magnification (x40) of a corallite section illustrating diverging fibres of a vertical
wall, a; and the similar structure of the basal wall, b; and tabulae, c.

80 MARGARE] K, PULLER & RICHARD J, F JENKINS

Fig. 5A, Transverse section of Holotype SAM P34167 (x38), B, Transverse section of Paratype SAM P34168 (x45) showing
variition in comtlite shape and septa, Tabulae are observed as irregular lines crossing the corallite: the midline of the
wall (arrowed) may be seen it some adjoining corallites. C_ Enlarged section (x10,5) of SA illustrating septa. wall irregularities,
Widline a und fabulae b, The reerystallized fabric is observed within the corallite. D. Corallite walls (x40) showing the
bundles of fibres which cross the wall (irrowed) in seations of the specimen.

TH RAKRLY CAMBRIAN TARULATE-LIRE CORAL ARRUB IPRA RI

of numerous tabellae and septal spues. In thin sections.
aw medal line divides the walls of adjoining eorallites
(Figs 5A.C}.

In transverse section al low magnifications (up to
X 40) straight to slightly diverging fibres crossing the
wills between adjacent corallites are Comunotily
disruplead by the medial line (Figs 5A,B.C°,D). tu
longitudinal section, fibrous elements diverge outward
and Gpwards fron the medial line and often protrude
inte the lumen giving the walls an irregular appeurance.
The walls which truneate parent corallites and form
the base of subsequent corallites, are composed of
vertical lo slighuly inclined fibres. These partitions arise
from the vertical walls and ure usually V-shaped, but
may be undulating, horizontal or inclined (Figs
4C DE},

In longitudinal section (Pigs 2.3,4A,B,C), individual
conallites are prismatic to irregularly cylindrical und
upto 14 min Wide and 47.5 nun long, Corallites vary
lithe in diameter and length, prior to the addition of
new coralfites (increase). Increase is both lateral ani
peripberal intracalicular. parricidal wilhin the
established budy of the colony (Figs 2,44,B).

‘Tubolae are numerous, comuionly formed of
invommplere. globose and dissepiment-like Labelle,
Occasionally some wre Continuous weTOss Very TUTTOW
cyrullites, Tabellae may arise from the wall. ot ftom
adjacent tabellae, extending jmwand and curving
downwards to pest upon other tabellae. They are very
thin, yenerally less than 0.06 mm. often wayy wnd
rarely strdight. Small projections often occur on Lhe
upper surface of tabulae (Fig. 4D). In transverse
section, tabellae are seen as wavy and crenate, srising
from the walls and dnastomosing with adjacent fabellae
(Figs 5A-C). At low magnification, the fibrous
structure of fhe tabellag is sinmilar to that of the walls,
with some bundles extending to give the srmull
projections on the upper surface, In longitudimal
section, the fibrous elements are normal to the byse
of the tabellac.

In transverse sectioyi, septal spines vary fram
nuMeryus (about 35) to absent and are often diffieult
to distinguish from other irnmegularities on the wall (Figs
SA.B.C). Where present they arc short (up to c¢. 1.25
ith), generally equal in length, blunt triangular or
spine-like in shape and equidistant trom each other
(about 0.25 to 0.5 mm), They are commonly present
on some walls while absent on others within a single
corallitg, Septal spines appear to be continuations of
bundles ot fibres of fhe fibrous wall, usually
rerniinating us, or being seen as fan-shaped tults (sec
above - Preservation), Lo longitudinal section, the
generally upturned septal spines are observed lo
oocasionally form short verlical rows on corallite: walls,

Discussion

4. fromensis is unlike the bw previously desorbed
Harly Cambrian carals from the same location,
Flindersipora bowmarni Laluste 1991 (e.g. Lafuste of
al. YA) and Movrawipara Chamberenyiv Puller &
Jenkins 1994, 4. fromensis is distinguished from F
bewmani by the size and general form of the colony,
(he position and shape of tabulae and septa, as well
ag the mode of inerease. In FE bonwwncn/, tabulae are
mostly complete and concave proximally, there are 6-16
strongly devoloped slightly curved septa, the edges of
which bear very short blunt spines; the walls are very
short Segments between the sepra. Increase is by
longitudinal fissien

The main differences between 4. fremmensiy and M,
chambereisis are in the size und form of the colanies,
the size and shape of the corallites and the arrangement
and shape of tabulae, Although bow are cerioid im
colonial form. the former is much larger amd usually
has parallel corallites, while those in M. charburinsix
are generally divergent. Corallites are prisinatiy to
cylindrical and up to 14 mim in diameter and 47.5 nim
in length in 4. fromensis. bul much srualler tup to 5
mim in diameter and {9.5 mm in length) ine tuberond
(yy integularly cylindrical in M. ckamberensis. "The
presence or absence ul septal spines /s common ta hott
corals: When present [hey are about the sane size and
shape

Tshulae differ greatly, being incomplete, globose and
dissepimentlike (abellue) in) df. fremmensyis and
coniplete, undulating and horizontal to concave upward
in M, chamberensis, Although the microstructure bas
not been studied at high magnification, there are sone
sumilurities between lhe aboye corals at low
mapnification. These toclude the purillel Abrus
elements of the selerenchyme evident in trunsverse
section, and the parallel to diverging fibrous elements
in longitudinal section, Fan-like arrays of hres. are
not present in M. chaniberensis, A medial Tine within
walls of adjacent corallite occurs in both corals. Medial
lines in the walls are common in iabulate corals. and
represent the external epitheea (Hill JOT).

A. froonensis 1s unlike ony of the previously deserined
Cambrian corals suggested by Serutton (1979) 1o have
tabulate affinities, but does have skeletal characteristics
in common with some of the Late Silurian to Lite
Permian michelininds,

‘The diagnestic characteristics for the genus
Michelinia De Koninek [841 inelace thin ty moderately
thick walls with & medial suture, short septal
trabeculae, tbulie incomplete and globose sometimes
with septal spinules on lhe upper surface. and large
mural pores (Hil) 1981). The walls und fibulwe ure
similar to those seen in 4. fromens/y hot ihe presen|
taxon Jacks mural pores.

42 MARGARET K, PULLER & RICHARD 1 F IENKINS

Micheline expanses White 1883 | Tabelleepliyiiun
peculiare Sturm \948) (Stumm M48) from the Early
Carboniferous of Arizona, is sintilar to A. frorerists
with respect to the form of the colony, the size and
shape of corallites and the arrangement of jabellae.
Corallites are up ta [S mm in diameter in the larmer
ane enim inthe latter, Corullites are wso of a-similar
shape. being generally 4, 5 or 6 sided, but difter by
the lack of septa in ML expanse. A most noticeable
simukarity between the two is the placement, size and
shape of the tabellae, They are incomplete and globose
and ure arranged in similar manner in both taxa, arising
from either the walls or adjacent tabellae. The tabellae
in A. fromensis appear to be less. glohose, spaced
slightly tirther apart, and havea more irregular and
wavy surface.

Aluiough 4. /ramensiy most closely. resembles some
of the mivhelinids, beeatise of the long time separation
between them (120 million years) itis highly unlikely
that they ure relisted and more probable their skeletal
similarities result from convergent evolution

Conclusions

The three described corals from the Moorowie
Formation, A. framensis, Mo chamberensiy and F,
Hewnani’, are very different in fori and arningement
of the skeleton, The diverse nature of the corals from
his ancient reefal environment indicates that during
the Early Cambrisn, variubility in polyp form and
skeletal morphology was well established.

The genus Lichenurie has been recognized as the
carhiest tibulate com, witha lime range from the base
Of the Barly Ordovictan tothe eurly Late Ordovicuin

(Serutton 1979, 1984. J992) Hull INST), Tt has beer
described as primitive, cerioid, of simple morphology.
usepture, but with tabulae and rare mural pores (Bassler
1950, Flower 96); MeLeod 1979, Serutton 984: Liah
1984). Although A. fromensis lacks mural pores, it hits
asimilar skeletal structure (0 some micheliniids which
post-datte Lichenaria. Most of the skeletal aspects ol
A fromensis are characteristic of Palueozoie tabulate
corals. These are (1) the cerioid form of the colony:
(2) walls separated] by a medial line reflecting
individual corallites (Scruton 1987); (3) the spine-like
lo wedge-shaped seplt occasionally situated in
longitudinal rows (Hill 1981); (4) individual corallites
which Spread above the pockets of sediment within the
colony, this. habit being usual tor enidarians following
influxes of sediment (Serutton (979): (5) lateral
increwse common, with peripheral intracalicular
increase being described in some Favositidae (Hill
W981). Although tabulae dre incomplete and dissepn

ment-like, they are consistently and strongly developed
both within individual and between adjacent vorallies
in AL fromensiy.

A. fromensis has anthozoan structural characteristics,
most of which are evident in tabulate corals. It should
thercfore prohably be included in the known group af
tubulates, thus extending the tine range of this group
to the late Early Canibrian,

Acknowledgments
The authors gratefully acknowledge Brent Bowman

und John Hart who collected the studied material, We
alsa thank Gino Snider for his phalographie work.

References

Bassinet. ROS (1950) Faunal fists and description of
Palucovore Corals Mem. Geol, Sa Anr a4, 1Al5,
Prowek, R.A (1961) Marleya and relitled colonial corals

Nee Mec Inst. of Ter Mem. 7, 1-97,

PoieR ML RK, & JesKiIne, RoE BE (00d) Moeprny pera
chemberenisis. a comal tron the Burky Cambrian Moorawie
Permation, Flinders Ranges, South Australia Trans, A
Sea S. luyé W864), 227-235.

Hi), D (ISAT) Rugosa and Tabulata pp, 1-762, fe Teicher.
C (Ed) “Treatise of Inyertebrite Palaeontolowy. Part F,
Coglenteritta, Supplement 1 (Geol. Soe Am. & Uno
Kansits, Boulder, Coloradey-

Larusie, J, Drogisxt, Bo Gaspin, Ao, & Gravesruck,
1, (991) The oldest tabulite goral and the associated
Archucoeyalbit, Lower Cambrian. Flinders Runes, South
Ausrnilta, Geublos 24, 697-718

Lape, R. 8, (i984) Liehendrty WINCHELL &
SCHUCHERT, 1895! Lamotttia RAYMOND, 1923, and the
carly history of the tabulate corals. IV Liternational
Svinpasium on Fossil Cidaria Puluewmniovr am, Sd,
154-163

Oerkesmore, K (1989) Djagenesis in corals: syntirvl
cements as evidence for post-mortem skeletal thickenings
V Internavooual Symposium on Fossil Cnidaria. Massif
Cnideiria 5, V9-177.

Mereap. J BD, 1979) A Lower Ordewiciin (Cunadiin
livhenarid Coral trom the Ozark uplift area, 2 Paleont, 33,
§05-5U6.

Savarisi, M_, Meine, JF, SoKave, dE & Bucks IN, I
(1983) Paleobiological and paleoenvironmental context Of
coral bearing Eyrly Cambrian reefs: implitations for
Phanerozuie reef devclopment. Geology 24, 917-920.

Scrotton, © T. (1979) Barty fossil Cnidarians pp. (6i-207
In House, M- R. (Bd ) “The Origin of major invertebrate
groups” (Academic Press, London & New York).

—— WOK) Origen and carly evelution of Tabulate Comets
WV International Symposium on Possil Cnidaria
Palavaniiolagr Ane Sd, WO-1R.

—— (1992) Findersipora boven: Lafuste. ane the early
evojulion of the tabulate comals: Fossi/ Cnidaria & Parifera
21, 29-33,

Srunm, B. R (1946) Upper Devonian tetracorals trom the
Manin Limestone, Felaennir 22, 40-47

CHARACTER AND INTERPRETATION OF THE REGOLITH
EXPOSED AT POINT DRUMMOND, WEST COAST OF
EYRE PENINSULA, SOUTH AUSTRALIA

By E. MOLINA BALLESTEROS*, E. M. CAMPBELLYf,
J. A. BOURNET & C. R. TWIDALET

Summary

Molina Ballesteros, E., Campbell, E. M., Bourne, J. A. & Twidale, C. R. (1995)
Character and interpretation of the regolith exposed at Point Drummond, west coast of
Eyre Peninsula, South Australia. Trans. R. Soc. 8. Aust. 119(2), 83-88, 31 May, 1995.
The weathering mantle developed on granodiorite at Point Drummond, Eyre
Peninsula, South Australia is examined using thin section and XRD analyses. Stages
in the alteration of the granodiorite can be deduced by examination of the zonation of
the regolith; release of oxides and hydroxides from the parent rock; removal of iron
and kaolinisation; new concentrations of haematite in micropores; development of
nodular structure and renewed removal of oxides and hydroxides. The possible age
relationships of this profile with laterites and Plio-Pleistocene ferricretes from other
South Australian sites are discussed. The age of weathering is uncertain but it
predates the calcarenite (? Pleistocene) and is probably post Permian, with Plio-
Pleistocene the most likely.

Key Words: regolith, ferruginisation, Point Drummond.

Transactions af the Reval Savielx af S. Aust, W995), WY) Ba-RR

CHARACTER AND INTERPRETATION OF THE REGOLITH EXPOSED AT
POINT DRUMMOND, WEST COAST OF EYRE PENINSULA, SOUTH AUSTRALIA

by BE. MOLINA BALLESTEROS* E, M. CAMPBELL, J. A. BOURNEY & C. R. TWIDALEF

Summary

Mouna Bariisteros. E.. CAMpRrui. Bo M.. Bourne, JI A, & Twibsuby CR. (995) Character and
interpretation of the regolith exposed ai Point Druinmond, west coast oF Fyre Peninsula, South Australia. Trans;

Ro Noe. S. Aust, W9(2), 83-88, 31 Muy, 1995,

The weathering ante developed on granodiorite at Point Drummond, Eyre Peninsula. South Australia is
examined using thin section and XRD analyses, Stages in the alteration of the granodierite cam be deduced by
examimation of the zonation of the regolith; release of oxides and hydroxides from the parent rock; remoyal of
iron and kaolinisation; new concentrations of huemutite in micropores: development of nodular structure and
renewed removal of oxides and hydroxides. The possible age relationships of this profile with Laterttes and. Plio-
Pleistocene ferricretes from other South Austrahan sites are discussed, The age of weathering is uncertain but
i predates the calearenité (2 Pleistocene) and is probably post Permian. with Plio-Pleistocene the most fikely.

Kiey Worps: regolith. ferrivinisation, Point Drummond.

Introduction

The west Coast of Eyre Peninsula is characterised
by high cliffs eroded in dune calearenite (also known
as ueoliuniles see e.g. Crocker 1946) of Middle and
Late Pleistocene age (Wilson 199] !). The calcarenite
rests unconlormably on Precambrian rocks, mostly
igneous and metamorphic. with granite and gneiss
prominent, but including sandstone and conglomerie
neur Talia. The uncontormity is uneven and the base
of the calearenite commonly extends below sea-level,
Elsewhere. the Precambrian basement is exposed m
rather irregular shore plattorms and in the lower
sections of the cliffs which. however, are composed
mainly of the culcarenite, At several sites remnants of
the pre-calcarenite regolith are developed on the
Precambrian basement. One of the best exposures of
the regolith, in terms of thickness, completeness and
luleral extent, occurs at Point Drummond, a westerly
projecting promontory located on the west coast of
southern Eyre Peninsula, some 80 km north-west of
Port Linvoln (Figs | & 2).

The purpose of this short paper ts to describe the
mineralogical variations between horizons within the
regolith, and to discuss their genetic nnplieations, The
site is on the southern portion of Point Drummand,

* Department of Geology. University of Salamanca, 47008
Sulumarica, Spain,

+ Department of Geology and Geophysics, Unversity of
Adelaide, South Australia, S005

1! Witsow, ©. C. (1991) Geology of the Quaternary
Bridgewater Formation of southwest and central South
Australia Ph.D, thesis, University of Adelaide, Adelaide
(Unpubl).

Two profiles. one from a south-projecting peninsula
and one trom the cliff adjacent lo the access steps
several hundred metres to the north, were examined.
The samples were selected on the basis of colour and
textural variations, The profiles are so similar both in
appearance and upon analysis that they can be treated
as one,

Pont
Augusla
/
Paint
Brown . Streaky Bay
EYRE PENINSULA
Paint
babhart Cowell
Point

N Drummond

JS AUSTRALIA
Port
Lincoln TSA ADEE
0 bie) 100 Kms Uv

Fig. 1. Locution Map.
Description of the profile

The profile, approximately 18 m thick, was sub-
divided from the base to the top into five horizontal
zones on the basis of their colour and texture (Pig. 3).
Mineralogy and (exture have been studied in thin
section (using. samples impregnated with a thinned
aruldite to prevent any disturbance of the original
structures) and by XRD analysis of bulk samples.

na }, MOLINA BALLESTEROS. E M CAMPBELL. J. A. BOURNE & C. R. TWIDALE

ee . os -
Fig. 24, General view of Point Drummond, Eyre Peninsula, South Australia. 2B. The weathering profile sumple site, Point

Drummond peninsula, showing the coastal plaviorm developed in granodiorite, the goethite-rich zone at the base of the
cliff-and above this the white kaolinised zone, the haematite zone and the calcarenite remnant ut the surface

REGOLITH AT POINT DRUMMOND RS

Zone |

The parent rock is a gneiss of granodtorilic
composition consisting mainly of quartz, plagioclase,
biotite and muscovite with some orthoclase. Secondary
minerals resultiag from hydrothermal alteration of the
rock prior to weathering mclude sericite, chlorite (from
biotite), epidote-zoisite and calcite (from plagioclase
- Sample |, Fig. 3, Munsell Rack Colour N&. white
to. N3, dark gray - dry. The colours of other samples
are Munsell Soil Colours - 1994). The granodiorite is
intruded by amphibolitic and quartzitic veins. Schistuse
shear zones are also present. These various rocks are
all members of the Sleaford Complex, dated al
2,700-2,300 Ma and thus of late Archaean or Palaeo-
proterozoic age (Flinter al, 1984; Parker er.a/, 1985)
though the shear zones may resull from the Kimban
Orogeny (~1700 Ma - Thomson 1969).

A
25
a
c
2
iss]
5
20>
oO
o
= 10
—
Ee a=
9
a2)
2
)
B

s
F’s Old fissure
\

Stages in the weathering of the gneiss can be traced
by examination of the zonation of the regolith or
weathered mantle, assuming that the weathering Front,
or Jower limit of weathering, has descended into the
rock mass from the surface. Hence, in these terms,
the mitial stages of weathering are represented by the
zone immediately above the weathering front and the
most advanced by the near surface horizon.

The coastal platform eroded in granodiorte and
located between high and low tide levels, is. irregular
with many blocky and bouldery rises and intervening
clefis, Many of the outcrops are superficially altered,
wilh rinds of ferruginoys oxides. and hydroxides
developed at the margins of blocks, boulders and other
exposures. The rinds are also found bordering fissures
(Sample 2, 7.5YR 7/6, reddish yellow to 7.5YR 7/3,
pink). In this zone the thickness of the rinds increases
up the profile, but nowhere exceeds 5 cm.

sainjonijs Mau
jo Juawdojecsq

sainjonis pjo
By yO UOI}eEAIasUOD

¢ New void

Fig. 3. Schematic diagram of the weathering profile on the peninsula, Point Drummond, A. Profile. -V, Zones of weathering
(see teat for explanation); 1-11, sample numbers (sample 5 similar to 6). B. Principle processes: a, Unweathered parent
rock. b. First stage of weathering (goethite-rich zone). C. Kaolinisation. d. Formation of new structures and conventration
of haematite, @, Removal of iron and plasime separation. £. Development of peds and nodules.

tia) LC. MOLINA BALLESTEROS, E M. CAMPBELL, J. A. BOURNE & C. R. TWIDALE

The first stage of weathering discernible is the
development of the rinds around the corestones. In
these the epidote-zoisite minerals are dissolved and the
alteration of plagioclase is manifested by the
‘ppeurance of zones of rundomly orjented clays. The
clay minerals are prelerentidlly developed along
fissures presumably beenuse the latter allow
penetration of water. Biotite changes colour Irom green
ty hrownish-yellow, reflecting a release of iron which,
as goethite, is concentrated in cracks and lissures,

Zone i

Here. though the original rock structures are
everywhere distinguishable. some of the blocks and
boulders are entirely discoloured and the rinds ure
Uneker than those in Zone L (Samples 3 & 4, IWYR
8/4, very pale brown to (OYR 7/6, yellow) with a white
and (N8) developed on the outside, and a sirong brown
(7.5YR 5/6) interior, Again the rind increases in
thickness up the profile und, as in Zone f, ss also found
along purtings. The contact between the weathered rind
and the interior of the corestones low in the profile
is sharp but is more diffuse at higher levels.

‘The rinds are pale in colour and voids are apparent
in thin section. Plapioclase is progressively reduced
higher in the profile and essentially isotropic clays,
beheved to be mainly kaolinite, become doyniient.
With the XRD method used, it is diffigult to
dilterentiate any other polymorphs of kaolin. Resistarces
such as quartz and muscovite are present,

Fone Lit

Almost the enure rock in this zone 1s white (Sample
6 N&) but most of the original textures and structures
ure preserved and remnants of the yellow and red iron
oxWes occur as spots in the upper pant of the zone
At the top of Zone IH loss of material has led to the
formation of voids. The weathering plasma (in the
sense of Nahon 1991, p. 63), derived from weathering
Of the parent materials, begins to appear anisotropic,
especially im areas close to voids [the vosepic plasma
separation ot Brewer (1964. 1976)]. Some, thuogh not
all, oxide concentrations are related to voids.

Zone IV

The yellow and red spots present in Zone U1) here
iierge to give a mixture of Oxides and hydroxides of
iron in differing degrees of dehydration and
erystallisation (Sample 7, SYR 4/4. reddish brown lo
TSVR 6/8, reddish yellow io JOR 4/3, weak red;
Sample & IOYR 8/8 yellow to IOR 4/3, weak red;
Sumple 9, IOR 4/3. weak ted to IOR 5/6 ted). The zone
is up to four metres thick. None of the original
structures survives.

XRD analysis shows that haematite is dominant in
the weak red patches, kaolinite having been removed,
On the other hand, where there is no iron oxide or
hydroxide, the kaolinite altéroplasina is well preserved

Muscovite remains. alber! weathered to varying degrees
and quartz. is corroded and in some instances clearly
disapgregated, At the lop of Zone LV voids are
common,

This zone is Similar to the “mottled clay™ horizon
from a laterite profile deseribed by Nahon (1987), of
which alumina (not analysed in this study) is a typieal
component

Zone V

The lower part of this zone ts characterised by
irtegulir Ul-defined fissures in the mottled clay, Also,
anew structure occurs in the form of polyhedral peds
some centimetres across, They bevorme stoaller and
more rounded upwards, where they take the form of
soft nodules 0,5-1.0 cr in diameter, reddish vellow in
colour but with red oxide conceatrations 1m the interior
(Sample !0, 75 YR 6/6, reddish yellow to 1OR 4/6, sed)
(Fig, 4). The partings which define the polyhedral! pecs

Pig. 4. Thnn section (crossed polars, plain light. x25) of upper
partof Zone Vo Kaolinite plasma surrounds the ferruginous
nodule (itrk ured). The light-coloured material in both the
nodule wnbthe surrounding plasma is quan.

are preferential zones of leaching. Removal of oxides
and hydroxides has resulted in zones of concentration
of skeletal grains (mainly quartz) fram the purent
matenial, and in the appearance of 4 kaolinitic chiy
plasma (a pedoplasma in the sense of Boulet 1974),
It grades trom asepie, where oxyhydroxides are
abundant, to skelsepie, vosepic and even masepic
where the oxyhydroxides have disappeared (Brewer
64, 1976). The development of the nodules is a
centripetal process involving the removal of oxides ond
hydroxides from the margins of the peds and their
concentmtion in the aodules, the redistribution of clays
and the concentration of skeletal grains in the leached
Zones.

Both the promontory and the cliff profiles are over
lain by a calcareous crust (calerete - Sample }t, 2.5Y
7/2, hyeht gray), located at the base of the calcurenite
but developed on the weathered material and including
nodules like those found ta Zone V CPi 4)

REGOLITH AY POINT DRUMMOND gy

Interpretation

Various stages in the weathering of the granmtivrete
pre eviderd;

\. ‘The penetration of metearig walters trom. the
surface und release of vaides and hydroxides fron
minerals in the parent rock, especially biotite, ts the
initial process in evidence The iron oxides are
concentrated in fissures at Lhe weathering front, at the
margins of blocks and houlders waned adjacent ty jount
purtings. The hydrothermally altered minerals are
dissolved and the plagioclases are weathered Lo clays.

2. Removal of ifon aad kaolinisation are represented
by the appearance of white rinds. These processes are
usually achieved in acid reducing solutions, which
wppear to have leached most of the calcium, iron and
sodium and some of the silica and produced the newly
formed mass of isotropic to insepic materials. (see
Brewer 64, p, 309), mamly kaolinite, Inside this
isotropic plasmic material are resistates such ws quarks
and muscevite {skeleton grains, of Brewer (1964,
1976)), On the whole, however, kaolinite is dominant,

3. Ferruginigation results from (he progressive
development of & kaollnitic plasma, and a new porasity
plus the destruction of the original features ancl
structures. Micropores, especially, become the sites
of new concentrations. of oxides (weak red spots),
particulurly haematite (Pig. 4), due to the decreased
mobility of the solutions in these pores (Dichter ef al.
1983; Tardy & Nahon 1985), This.is the origin of the
‘mottled clay horizon’ typical of protiles develaped
under seusonally wet and dry climates,

4. The development of nodular structure is related
to the removal of oxides and hydroxides by solutions
probably emanating from an overlying soil, A new
k#olinitic plasma is developed as the oxyhydrox ides
are remoyed. The mobility of the materials 1s governed
by the amount of oxyhydroxides: the less oxyhydroxide
the more plasmic movernent and hence better ree
organisation of the soil mass.

5, The removal of oxides and hydroxides requires
nod salutions (i.e; those which are poor in carbemuales).
so thit the processes described in paragraphs 1-3
inclusive predate the development of the calerete and
the deposition of the dune culearenite

The age of the profile, and the events ta whicl tt
relales, are mainly problematic. 1is clearly younger
than the Proterozoic rocks on which itis developed,
and predates the calcarenite which, according to Wilson
(1991). is Middle and Late Pleistocene i age (maxi-

2 Hossre.o, P. (1926) The pcology of portions of the
Counties of Livht, Epre, Sturt and Adelaide, MSc thst,
Univwmiry ob Adehide, Adeline (Unpubl),

mum-¢. 700,000 years). But allocating at to an hiatus
bl sore 1500-2000 Ma is neither precise nor inform

ative, The extent of the hiatus can be reduced if two
general arguments are accepted, First. although
regoliths have survived the passape of ice shivets (see
ey, Fogelman 1985), the profile, which has apparent
equivalents al several points along the west coast of
Fyre Peninsula (e.g. Point Brown, Point Labatt, Talia),
is anlikely to have survived the Early Permian
glaciation, which evidently affected most of the present
state ol South Austratia (e.g, Ludbrook 1969), and
subsequent erosion: for the reyolith has readily been
eroded by marine agencies and by gullying, In these
terms the regolith under debate is less than 250 Ma old.

Second, the hiatus is farther reduced if il is conceded
thin the revohith is most likely immediately to predate
the cover material, that is the calcaremte, This last
suggestion assumes that even if the development of the
regolith began long before the deposition of the dune
lunestone. it would have continued to evolve (see ee.
McFarlane W986; Bourman 1993) up to (and even
beyond) being covered. In these terms the young date
for the regolith is of the order of 700,000 years, though,
because ibmust have develgped over a long period, it
ought to be assigned an age range ane could reasonably
he labelled Plio-Pletsiocene.

Broader considerations support this suggestion.
First, with what other regoliths might the Point
Druinniond profile be retites!? Perruginous regoliths
are known from various parts of South Australia (e.g.
Hossteld 1926-5 Northcote 146. Miles 1952; Horwitz
& Daily 1958: Campana 1958; Cilaessner & Wade
1958; Horwatz 1960, Wopfner 1967; Daily et al. 1974;
Twidale eral, 1976; Wright 1985; Milnes eral. 1985;
Bourmanh er af. 87), They hive been variously
defined and interpreted (see e.g. Bourman 1993 - but
see also McFarlane c.g. 1986: Firman 1994), Some,
characterised by » sandy Or silty Achoraton, a massive.
commonly pisolific, ferrnginous horizon averlying a
thick bleached zone, have been labelled lanerite, Others,
consisting af cither a [erruginous crust aloe, of a onust
resting ona thin bleached horizan, huye been lermed
ferricretes (¢ 2, Lumplugh 1902; ‘Twidale 1976, p
196-197), The Point Driuniond profile does. not sit
eusily with either of these, bul is perlaps closer to the
ferricrete than to the Ialerile. particularly af it is
considered logether with other stratigrapically
comparable regoliths such ap that exposed at Point
Brown. Ferricretes in southern Soulh Australia have
been dated by various means. but oo Yorke Peninsula
(Horwitz & Daily 1958) local stratigraphy indicates
a Pliucene age. Equally, occurrences in the interior ol
Eyre Peninsula have, on stithgtaphic grounds, also
been attributed to the Teniary, some being considered
Eocene but others clearly Pliocene or post-Pliocene
(Rankin & Flint 199); Flint & Rankin 1991; Flint 1992),
Given the Middle Late Pleistocene age of the overlying

4H bh MOLINA BALLESTEROS, B, M. CAMPBELL. J. A. BOURNE & © R. TWIDALE

caleurenite. a later rather than an earlier Tertiary age
scems appropriate for the Point Drummond exposure,
und on balance a Phi. Pleistocene attribution is in
keeping with the available evidence,

Acknowledgments

The authors thank Dr MJ. McFarlane for comments
and suggestions during fleld work and Mr M. J. Wright
und) Iwo referees for constructive suggestions on ar
carly draft. Professor Molind’s-work in the Universily
of Adelaide in the period August-December 1992 wats
mide possible hy a grant trom the Direecion General
de Investigacion Cientitica y ‘Teentea (DGICY'T).
92-176, Ministerio de Educacion y Ciencia, Spain

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A REVISION OF THE GENUS TIKUSNEMA
(NEMATODA: ACUARIOIDEA) WITH THE DESCRIPTION OF
A NEW SPECIES FROM THE FALSE WATER-RAT
XEROMYS MYOIDES FROM QUEENSLAND

By LESLEY R. SMALES*

Summary

Smales, L. R. (1995) A revision of the genus Tikusnema (Nematoda: Acuarioidea)
with the description of a new species from the false water-rat, Xeromys myoides from
Queensland. Trans. R. Soc. S. Aust. 119(2), 89-94, 31 May, 1995.

The genus Tikusnema Hasegawa, Shiraishi & Rochman, 1992 is redescribed. The
species Molinacuaria indonesiensis Gibbons, Cranshaw & Rumpus, 1992 was found
to be synonymous with Tikusnema javaense Hasegawa, Shiraishi & Rochman, 1992,
the two species having been described almost simultaneously from the rice field rat,
Rattus argentiventer. A new species of Tikusnema from the false water-rat Xeromys
myoides is described. Tikusnema vandycki sp. noy. can be distinguished from T.
javaense by the size of the adult male and female, the shape of the cuticular leaves on
the pseudolabia, the length of the male tail and spicules, the length of the female tail
and size of eggs. The implications of the presence of acuariid nematodes, normally
found in birds, in a range of small mammalian hosts, are discussed. The significance
of the presence of Tikusnema in Indonesian and Australian hosts cannot be
determined until its presence or absence on the island of New Guinea is confirmed.
Key Words: Tikusnema, Nematoda, Acuarioidea, Xeromys myoides, false water-rat,
mammalian hosts.

Transietions bf re Hewal Soler af A Maye (19057, TEC) 89.94

A REVISION OF THE GENUS TIKUSNEMA (NEMATODA: ACUARIOIDEA) WITH

THE DESCRIPTION OF A NEW SPECIES FROM THE FALSE WATER-RAT
XEROMYS MYOIDES FROM QUEENSLAND.

by LESLEY R. SMALES*

Summary

Svares. dh. R995) A revision of the genus 7iaisnema (Nematodu: Acuarioided) witht the description of a
new species Prom the false water-eat. Neronvs vryaides trom Queensland. Trans. Re Sue. &. Aust. E92), 894A,
41 May, t995,

Nie genus Twsene Hasevawa. Shiraishi & Roehman, 192 is redeseribed, The species Melineenarta
iIndonesiensis Gibbons, Crawshaw & Rumpus, 1992 was found lo be synunyrnous willl Tikisndvte Jd vdense
Hasta, Shiraishi & Rochman, 1992, the two species having been desorbed almost simultuncously fremy the
fiw (eld ray, Aaais urveriventen A new species of Tkusrena trom the false water-nt Aeron mveides ts deseriled,
Tikusnema yandyeki sp. noy, can be distiawuished From 7 jeveense by the size of the adult male and lemale,
he shape of the cUlicWar leaves on tie psetidolabia, the length of the male tail and spicules, the leagih of the
jumule tailand size of egys. ‘The implieauons of the presence of acuariid nematodes, aormally found in birds,
ina range of small mammalian hosts, are discussed, The significance of the presence of JrAwsnenta in Indonesian
vind Austrian hosts cannot be determined until ts presence or abserice on the island of New Guinea 1s comiirmied.

Key Worps: 7ikusnenta, Nematoda, Acudriodea, Xeraumyys nevetdes, false water-ral, mammalian hosts

Introduction

The lalse water-rat Xeremius myoides Thomas. 1889
isa small dark grey semi-aquatic rat whose preferred
habitul 15 shallow coastal wetlands, such as swarnps,
mangroves, forests, lagoons. or sedged lakes (Van Dyck
1994). ‘They are currently known from only six sites
in north-centoul and north-eastern Australia. Their
current Conservation status is valnerable und likely to
progress lo endangered because of human proclivity
to drain and develop swamps (Van Dyck 1992) They
forage on the mud flats for food items including aquatic
invertebrates, such as crabs, mud-lobsters, niussels.
midrine pulmonates and polyclads (Van Dyek 1994).

Nematodes dissected from specimens of X, yrvoides,
collected by staff of the Queensland Museum were
found to be species belonging to the Acuarioidea. The
genus Tikusteme was erected for specimens from
Rarus arveniventer (Robinson & Kloss, 1916), the rice
field rat from West Java by Hasegawa ev al. (1992).
Almost simultancously anew species of Molinaeuaria
was described, also fram Ro argennivenrer from Java,
by Gibbons er al, (1992).

Comparison of material fram X. niyeides with type
specimens of both species deseribed trom Autrity
argentiventer suggest that the nematodes from X.
wiveides are new species of Tikusheia while all the

* Biology Department, Central Queensland University.
Rockhampton 4702 Queensland,

material from R, urgentiyenter ws cop-specific.
Molinaeuaria indonesiensis therefore falls ay a
synonym of Fikusnenta javaense.

Materials and Methods

Six lalse water-rats, Neronny anvoides. were collected
from Myora Swamp, Stradbroke Island, Queensland
during 1992, Fuecal pellets from two hosts were
examined. The ulimentary tracts of the tour other hosts
were dissected for helminth parasites alter the bodies
had been fixed whole in 10% formalin. The nematodes
so collected were cleared in lactophenol for
microscopic examination. Figures were drawn with the
aid of a drawing tube. Measurements, of 10 specimens
in jem unless Otherwise slated. with the range followed
by the mean. were made with the aid of an ocular
micrometer, drawing tube und measuring wheel.

Specimens of Molinacugria indonestensis Gibbons,
Crawshaw & Rumpus, 1992 and Tikusnema javacnse
Haseyawa. Shiraishi & Rochman, 1992 from Rattus
areentiventer were also examined for comparison,

The terminology used for morphological features in
the deseriptions is that of Hird and Bird (1991) and the
taxonomic system of Anderson (1992) is followed.
Abbreviations. aré: Queensland Museum QM:
Australian Helminthological Collection South
Australian Museum SAM: United States National
Museum Helminihologicul Collection USNM:
International Institute for Parasitology 1"

90 LESLEY R

Systematics
Order Spirurida
Suborder Spirarini
Supertumily AcuarioiWed
Family Acvariidae
Sub family Seuratiinae
Genus Tikusnenia Hasegawa. Shiraishi & Rochman,
1992
Type Species Tikusniemea javiense Hasegiwa, Shiraishi
& Rocha, 1997,

Tikusnema
Hasegawa, Shiraishi & Rochman, 1942,

Revived veneric diagnosis

Cephalic end inflated and set off from body by
consuicuon, Oral opening laterally compressed.
Poeudolubta large, triangular in latery] view, each with
two cephalic papillae and an amphid, Pseudolabia
attached 10 each other apically. separated dorsally anc
ventrally by cordoas and indented deeply at level of
wimerior extremity of cordons. Posterior end of each
pscudolabium forms two cuticular leaves euch sub-
divided into teeth, Cordons small, net extending
postenorly. Buccal cupsule long, culicular wall thick-
nol striated. Pharynx divided into anterior muscular
and posterior glandular portions. Deirids small,
bicuspid. A pair of cuticular ornamentations: present
laterally, posterior to deirids, Parasitic in the stomach
or intestine of rodents.

Tikusnema javaense
Hasegawa, Shiraishi & Rachman, 1992,

Synonym Molinacvaria’ indonesientsis Gibbons.
Crawshaw & Rumpus, 1992; pp. [75-181
Material exaniined

From R. atgenrivener: \o allotype Tikusnema
javaense USNM 822235 Pusakanagata, West Java,
Indonesia: 1 O° paratype IP BIOSSB from Sukamandi,
West Java, Indonesia.

Description

As in Hasegawa ef al, (1992), From the combined
Measurements of both Hasegawa ef al, (1992) and
Gibbons ef al, (1992) the dimensions become as
tollows:

Male: length 9-2) mm, width 277-440, Buccal
capsule 359-490 Jong, muscular portion of pharynx
410-560, glandolar portion 1130-1980 long, Deirids
296-440, nerve ring 450-560, excretory pore 525-830
from anterior end, Right spicule 190-210, left spicule
491-570 long; tail 556-990 long,

Female: length 11.0-24.5 mm, width 293-510. Buccal
capsule 330-490 long, muscular portion of pharynx
402-630, glandular portion 860-2030 long, Deirids
273-430, nerve ring 410-630, excretory pore 502-870
from anterior end, Vulya 5.02-12,21 mm from anterior
end, Tail 230-520 long. Egys 28-31 by 38-44

SMALES
Remarks

Tikusnema javaense and M. indonesiensis were
deseribed almost Simultsnewusly by Hasegawa et al
(1992) and Gibbons e7 al. (1992). the descriptions
appearing in different journals, Both cescriptions
referred tw material collected on the island of Java from
Rattus argentiverser, A caretul examination of (he
deseriptions given by each group of authors, together
with a comparison of the material they exawiined, has
revealed that they gre of the same species. Any
differences In measurements benween the two sets of
materiul relate only to the fact thar the specimens
described by Hasegawa ez.a/. (1992) were smaller thin
those described by Gibbons eral, (1992). The lemales
described by Gibbons er a/. (1992) for example ure
lurger, gravid females containing embryonated epys
while the smaller females described by Husegawa et
af. ()992) contain unfertilized epgs.

Hasegawa er a/. (1992) describe and figwee a pair
of cuticular ornamentatiuns much larger then the
deirids in the adult worms and even more prominent
in the 4th sutge larva. Gibbons et al. (1992) did not
mention this feature in their description of their more
mature worms. Therefore it appears that the cuticular
OTmamentavions may he 4 more prominent leature ol
juvenile than mature worms, This would account for
their upparent absenee in the specimens examined by
Gibbons et al. (1992).

The interpretation of the cephalic ends of the
specimens, in particular the origins of the cuuicular
leaves. by Hasegawa er al, (1992) appears consistent
with both sets of muterial. Ay discussed in Hasevawa
et al, (1992) the cuticular leaves of 7ikusnema originate
directly tron’ the pseudolabia without separating
furrows. Tikusnema also has small cordons nol
extending posteriorly. By contrast the genus
Molinacuaria, although characterized by the absence
of pseudolabia (Wong & Lankaster 1985), does have
Brooves. located immediately anterior to ptilina,
Molinacuaria can be further differentiated from
Tikusnema by a lack of cordons, Moalinacuaria
indonesiensis lherelore falls ws a synonym of Tikusnenn
Jovaense. The species name javaense has prioriry
because it was published in October 1992, while
indonesiensis did notappear until in November of that
yeur.

Tikusnema vandycki sp. nov.
FIGS 1-16

Material examined

From Xeremys myeides. 54 mmmature adults and
fourth stage larvae, 31 anterior ends, 40 mature & cr,
26 mature 9 2 from Myora Swamp, Stradbroke
Island. Queensland.

TIKUSNEMA SP, NOY FROM THE FALSE WATER-RAT ww

Pescriprian

Long, Slender worms with tapered extremities,
clticle thm, with fine annulations, Lateral alae absent.
Cephalic culicular leaves each divided into 4-5 teeth,
lateral rooth targest (Figs 2.3.5,16), Cordatis rod-like
in dorsoventral view (Fig. 2). Cordons and dorsal and
ventral rim of pseudolabia faintly striated (Figs 2,8).
Muscular portion of pharynx narrower and shorter than
glandular portion, pharynx aboot 1/7 body length (Pips
1,15) Nerve ring near anterior end of muscular portion,
excraory pore posterior ta nerve ting. Derrids Liny,
bifid, between nerve ring and exeretory pore (Fig, 1),
A pair ot culictlar ornainentatieons, saruil.
inconspicious at about mid level of plandular ponion
of pharynx (Fig. 12).

Mile: Length 27 (25. 30) mm, width at midkxidy 412
(317-470), Cephalic end 177 (156-245) long, 240
(215-260) wide, Posterior end of cuticular leat 220
(208-26) [rom anterior extremity. Buccal capsule 269
(260-287) long, muscular portion of pharynx 435
(370-680) long, glandular portion 3229 (2975-3872)

long, Detrids 307 (186-325), nerye ring 377 (338-410). °

excretory pore 499 (442-559), cuticular ornamentation
(one measurement only) 2685 from anterior end.
Poslenor region curved ventrally. Caudal papillae
arranged in 10 pairs, 4 pairs pre-anal, 6 pairs post
anal, large pedunculate, Ist and 2nd pairs grouped
logether, 3rd and 4ih pairs grouped together, Ist anc
3rd more lateral, Sth and 9th pairs grouped together
Sth more lateral; th pair close to tail tip. Longitudinal
cuticular ridges present anterior to cloaca (Pig. 19)
Spicules dissimilar, right spicule short robust, rounded
distal tip 241 (208-266) (Mig, 9); lett spicule, trifid
distal tip 746 (682-813) long (Pigs 4.14), longest spicule
about 1:36 body length, tail 721 (598-845) (Pip. I).

Female: Length 34.8 (30-41) mm, width at mid body.
555 (510-629). Cephalic end 188 (156-201) long, 255
(240-273) wide. Posterior end of cuticular leaf 238
88-260) [rom anterior extrenuly. Buceal capsule 282
(266-292); muscular portion of pharynx 461 (325-650)
long. glandular portion 4040 (3111-4675) Deirids 336
(273-383), nerve ry 399 (357-422), excretory pore
500 (455-546) from anterior end, Vulva circular.
without Jips, 16.5 (3.6-19.4) mm from anterior
extremity. Ovejector amphidelphic: vagina vera
directed transversely, 550 (one measurement), vagina
utering 250 (oné measurement) parallel to body wall
(Pig. 6). Tail 621 (510-748) (Figs 7,10). Eggs thick
shelled, 33.8 (32-34) by 48 (44-53).

Etymology

The Specific name vandyeki is given in recognition
of Steve Van Dyck who first noticed the presence of
these worms in the host,

Host; Xeramyy onyeddes
Location, Stomach

Locality; Stradbroke Istaned, Queenstand

‘Type specimens: Holotype male. QMIL9I5; Allotype
female, QM211926,

Paratypes:! QM2\1927-30; SAM24832,

Remarks

Tikusnema vandycki can. be distinguished from T:
javaense by the shape of the cuticular leaves. In T
vandycki the leaves are subdivided at the edge into 45
teeth, but in 7 javaense Ute leaves have three teeth,
the middle one being the most prominent. Jikusnema
vundyek? can be further distinguished by its larger size;
miles up to 30 mm tong, lemales up to 4t mim as
compared with 21 and 24.5 in 7% javwense respeetively:
The spicules of 7 vandweki are lonper (208-266 and
682-813) than those of 7) javeense (90210 and
SOK)-570). However since 7 vardyek!is.a larger worm
than 7. javiense the proportion of left sprcule to body
length ts smaller for TL vandvek? (36) than tor 7
javeense (E18). Male @ vandveki have a sberter tail
(598-845) than. do T javacnse (B40-990). The eyes of
T. vandychi (44-53 x 32-34) are larger than those of
T. javaense (38-44 x% 328-31), Comparative
measurements of 7. javaense and T vandyeks are given
in Table J. Since the specimens exanuned by Hasegawa
er al. (1992) are smaller immature adults only the
measurements from Gibbons ef al. (1992) of mature
specimens are used. "This allows an easier comparison
of the relative sizes of nouture adult specimens of cach
species. The paired cuticular ornamentations at the
level of the glandular portion of the pharynx are tiny
and difficult to find in 7. vanevekt but more prominent
in 7) javaense. The vagina vera of 7) vandycki appears
to consist-of nwo parts, # globular heavily cuticularized
part leading into a more tubular less cuficularized part,
which in turn opens ilo vagina uterina (Fig, 6), The
vagina vera of T jawaense is similarly figured in
Gibbons er ad. (1992), Further investigation is needed
to determine whether the vagina vera is actually
bipartite or whether the distal, globular part is. actually
an ¢laboration of the yulya,

Fourth stage larvae and immature adults of 7
vandyeki show similar morphological features to. those
of 7 javaense. A detailed comparison and anilysis,
panticularly of the development of the cephalic
structures, 38 being prepared for a subsequent paper

Discussion

The spirutid superfamily Acuarigides is usually
found in birds. However an increasing: number of
genera has now been reported from mammals, The
genera Starimerinemd Osche, 1955, Antechiniella
Quentin & Beveridge, 1986, Chandlerancma Lute &
All, 1980, and now Jikusrtenta occur exclusively in

92 LESLEY R. SMALES

TIKUSNEMA SP. NOV. FROM THE FALSE WATER-RAT 93

TABLE 1. Mean measurements, in wm unless otherwise idicated, of Tikusnema species. Measurements of T. javaense are
from Gibbons et al. 1992. Standard deviations are given for the measurements of T. vandycki.

T. vandycki T. javaense T. vandycki T. javaense
Male Female

No. of specimens measured 10 4 10 2
Length in mm 27+ 2.15 20.19 34.8 + 3.16 23.5
Width 412 + 43.39 410 555 + 43.67 445
Length buccal capsule 269 + 13.9] 465 282 + 31.55 515
Length muscular pharynx 435 + 95.09 540 461 + 89.07 585
Length glandular pharynx 3229 + 383.86 1870 4040 + 578.37 1935
Deirid to anterior end 307 + 48.85 390 336 + 33.15 410
Nerve ring to anterior end 377 + 22.09 550 399 + 24.96 585
Excretory pore to anterior end 499 + 37.45 765 500 + 27.58 825
Right spicule 241 + 19.73 200 - -
Left spicule 746 + 53.62 535 - -
Tail 721 + 67.32 915 621 + 65.41 505
Vulva to anterior end in mm (one specimen) - - 16.5 12.21

Figs 15,16. Photomicrographs of the anterior end of Tikusnema vandycki sp, nov. lateral aspects. Fig. 15. optical section.
Fig. 16. showing the cuticular leaves of the pseudolabia. Scale bars =100nm. Arrows indicate cuticular leaves.

Figs 1-14. Tikusnema vandycki sp. nov. Fig. 1 Anterior end, lateral view. Fig. 2. Cephalic region, dorsal view. Fig. 3. Cephalic
region, lateral view. Fig. 4. Left spicule. Fig. 5. Cephalic region, enface view showing cuticular leaves. Fig. 6. Vulva,
vagina and uteri, lateral view. Fig. 7. Posterior end female, lateral view. Fig. 8. Cephalic end, enface view, optical section
showing cordons, Fig. 9. Right spicule. Fig. 10. Female tail tip. Fig. 11. Posterior end male, lateral view. Fig. 12. Cuticular
ornamentation in pharyngeal region. Fig. 13. Posterior end male, ventral view showing cuticular ridges arising anterior
to the cloaca. Fig, 14. Left spicule tip. Scale bars: Figs 1,6,4,=100 um; Figs 2,3,5,8,=50 wm: Figs 7,11,13,=200 am;
Figs 9,10,12,=50 pm; Fig. 14,=25pm.

4 LESLEY R, SMALES

mammils. (Gybbons er ul 192) while others
Synhimantus Raatliet, Meory & Sissotf. 1912-
Farcewuria Rao, 195) and Skt vebinuclave Sobolev.
1043 withourh primarily found in birds, ulso occur in
mammals, Various arthropods and fish serve as
intermediate hosts lor the life cycle stages of acwariids
(Anderson 1992). The link between mamunalian Host
and acuariid parasite therefore may be one of dietary
habit (Stnales 1991), A particular set of dietury
preterences and habits of a few mammals thus allows

these odd oecurrences of iifechon by deusriids of

byologicully unrelated host species. in geographically
unrelated regions of the world. Slirews from Bulgaria,
Isracl, Alaska. Burope: mice rats, raccoons, muskrats,
from USA, rice Tield rats from Indonesia) pyreuean
desmans from Spain: Anrecitinus species, water-rals
and false water-rats from Australia are all able to be
parasitized by acuariids under apptopriate cireunt
stances (Quentin & Beveridge 1986; Hasegawa ef a.
1992; Alvurez eral, 1994; Anderson & Wony 1994),
The precise nature wf the link would probably differ
from one maromalhian host to another, For example the

diet, including crustaceans, and semi-aquatic habits of

WN. myeides appear to fit the required pattern,

Australian rodents are all included within the farrily
Moaridac Their ancestors are believed ty hive: evolved
in South-east Asia about 25 million years ago (Watts
& Astin 1980, Then some 15-20 million years aye
members of the lineage colonized the Indonesian and
possibly some Melanesian islands. Geological changes
during this period :solated the islands Jor greater or
lesser periods of tirae allowing further speciation to
occur, By 5-10 million years ago Australia and New
Guiiea had moved close enough to these islands to
allow colonization by what has becore Koowe as the
old endemic rodents.

The Australian water rat group, the Pydromyint fony
part of that old endente fiuna, The suggested perics!
of divergence Within the group (Warts & Astin L981)
would have the Australian and false water rats ovalvine
along separate lineages belure cheat arrival in Australia
Both genera have closer affinities. with Vavious New
Guinean rat species, in body torn god ecoloyieal niche,
than they do willyeach other, The fact that both genera
have acuariid parasites can be seen as a refleerion of
their aquatic fo semi-aquatic life-styles und the
inelasion of crustaceans in their diet. However, the
acuariids found mf chryogaster, Anrechinella and
Swrhimantus are also found im Australian casyurid
marsupials whilst Vikwsnenma from XN myoldes also
vecurs in R. arvenrventer from Widonesia. "This 96
consistent With the scenario proposed by Watts & Astin
(M981) that XL mvaides is a more reeent arrival om
Australia than ff, chrysexester, How the radiation of
the Hydrontyint is related to the murids of Southeast
Asia is unknown (Watts & Keniper 1989). A survey
of the prtrusites of the Papua New Guinean Hydromini
needed to determine whith, if any, acuariid parasites
are present. Conclusions may then be able to be crawn
as to whelher (he appearonce of Tikusnema in RK.
argentiverter antl X nryoidey has any sigmilicunt
bearing on murid telauonships in South east Asia
Papua New Guinea and Australia

Acknowledgments

My thanks to Steve Van Dyck and Lester Cannot
of the Queensland Museum for giving ine access be
the material from the false (water-nat, Dr Ralph
Lichlentels of (he Biosystematic Parasitology
Laboratory and Dr Lynda Gibbons of the International
Institute for Parasitology for the loan of the material

References

Anderson, B.C. (1992) “Nematode parusites of vertebrates
Their development and trapsinission” (CAB International,
Cambridge)

~—— & Wong, PL. (1994) Skrjubinovlavay Kinsellai wsp.
(Nematoda: Acuarioidea) front the clee wat Oryeanivy
palustris. in Flonda, Syst, Fevasite!, 28, |-+

ALVAREZ, FB... Guor-Boreni A. WH. Quinieino. PB. Rey. Lb,
Loprg-Romaw, Fo & Sanmartiiy, M. bL. (1994)
Paracuaria hispanic asp. (Nematoda: Acuarildae), a
stomach pariate of the pyrenean desman Galery
Pyeenuicus Geotly, Cinseetivera:Tulpidae). with a
oder of the genus Peracwanta Bao, 195i, did, 29,

S412.

Biko, A. Fo& Byrp, J. (1991) "The Structure of Nematodes*
2nd edtn, (Academie Press, San Dieu).

Crmmons, J. M_. CRAwsHAW, MT. & Rumeus. A. 1. 11992)
Malinieuaria indonesiensis tsp. (Nematoda Acuarioitesy
Tron Rats arventiventer in Indonesia, Syst Aarasital 23,
175. 1BI.

HasnGaws, H, Strats, S d& RocHiman (1992) Tidteseerne
favaenve nosso rsp, (Nematoda: Acuarinidea) and other
nematodes from Rattus araenrtvener collecred in Wess dasa
J, Panastinl, TR. 800-404

Quentin, JC. & Brverince, L (86) Comparative
morphogenesis of the cephalic structures of the aeuariid
nematodes Starunerind sorties (Tiner, 951), datechinietls
Snffodiax (Beveridge & Barker, 1975) pyen,, meomb and
Shrjabineclava thapari (Teixeica de Freitas, 1953). dysr,
Parasitol. 8, 63-171,

Smaces, L. R_ (1991) A new species of dareciiniella Quentin
& Reveridge, 1986 (Nemutoda:Acuarndac) trom the
Australian water mit Jivdrgmus chiyyeyescen Geotlroy, (ROS
Trans, R. Soe, 8, Ast, W5, 217-220,

Van Dyex, & (1992) Parting the reeds on Myora'’s Nevrrnvy
kibbutz, Wild Aust 29, 8-10,

11994) The rats at nepiune’s table. tase, Mar. His?
24, 30-37

Waris, CHS. & Astin, Ho F981) "The Rodents of
Australia” (Angus & Robertson, Landon),

—___— & Keweek, CM, (1989) Muridae pp, 949-957 Jn
Walton, D. W. & Richardson. 8, J. (Rds) “Faun of
Aasiratia” Vol 1B Mammalia (Ausr Cievi Publishing
Service, Canberra)

Wona, PL. & Lancaster, M. W. (985) Revision of the
genus droyracanhopsts Dicking, 1861 and deseription of
a new genus Molintucudria nawen. (Nematoda:
Acwurtdoidea). Can, 4. Zool, 63, (S501564,

MASTOPHORUS MURIS (NEMATODA: SPIROCERCIDAE)
FROM THE MUSKY RAT-KANGAROO,
HYPSIPRYMNODON MOSCHATUS

BRIEF COMMUNICATION

Summary

The Musky Rat-kangaroo, Hypsiprymnodon moschatus Ramsay, 1876, the smallest
and most primitive of the macropodoids, occurs exclusively in the rainforest of
northern Queensland." It forages in the leaf litter of damp areas of the forest for fungi,
fallen fruits and invertebrates. H. moschatus has a sacciform stomach intermediate in
structure between the simple stomach of the phalangerids and the complex,
compartmentalized stomachs of the potoroos and macropods.” Apparently the Musky
Rat-kangaroo has not adapted to a diet with a high content of cellulose dependent on a
fore-stomach fermentation chamber, but rather has retained an omnivorous diet of
higher nutritive value.’

Tronsactions af tke Royo) Suctery of S Aust, (995), L9(2), 95~96,

BRIEF COMMUNICATION

MASTOPHORUS MURIS (NEMATODA: SPIROCERCIDAE) FROM THE MUSKY
RAT-KANGAROO, HYPSIPRYMNODON MOSCHATUS

The Musky Rat-kangarov, Aypsiprymnodon maschaius
Ramsay, 1876, the smallest and most primitive of the
macropadoids, occurs exclusively inthe rainforest of northern
Queenshind,! It forages inthe leaf litter of damp areas of the
forest for fungi, fallen fruits and invertebrates. H. rescharis
has a sacciforn? stomach intermediate in structure between
the simple stomach of the phalangends and the complex,
compartmentalized stomachs of the potoroos and
macropods.* Apparently the Musky Rat-kangaroo has not
adapted to a dict with a high content of cellulose dependent
on a fore-stomach fermentation chamber, but rather has
retained vn emniverous diet of higher nutritive value!.

A-small colony of H. maschatns has been maintained, for
research purposes, at the Queensland National Parks und
Wildlife facilities, Pollarenda, Townsyille, One of these
animals died in its pen in 1994 and was subsequently made
available for dissection. Twenty imale and 17 female
Mastaphoras muris (Gmelin, 1790) were found in the stomach.
The worms were fixed in 10% formalin, stored in 70% ethyl
alcohol and then cleared in lactuphenol for microseopic
examination. The males measured up to 25 mm and the
females 63. mm Jong. The worms appeared maturé and healthy,
the females being pravid, Mastopharas mariy is a nematode
from the family Spiroeercidac, cosmopolitan in rodents of
the families. Microtidae and Muridac*, [t has previously been
recorded from rats, a “mouse” and eats in Australia. 2" A
range of insect species including cockroaches has been finind
lo be surtable intermediate hosts.”

Both cock rmaches and rats are attracted to human food stores

'Jonnson, P.M. (1988) “Musky Rat-kangaroo” pp, 179-180
In Strahan, R, (Ed.) “Complete Book of Australian Mammals”
(Collins, Angus & Robertsen, Ausiralia).

“Hume, 1. D. (1982) “Digestive Physiology and Nutrition of
Marsupials" (Cumbridge University Press, Cambridge),
Vanderson, R. C. (1992) “Nematode Parasites of Vertebrates
Their Development and Transmission” (CAB International,
Wallingford),

4Mackerras, M. H. (1958) Proc, Linn. Soe. NSW 83/2),
101-160,

S5Obendorf, D, L, (1979) Aust, J. Zool, 27, 867-879.
6Smales, L. R. (1992) System. Parasitol. 22, 73-80.
7Gordon, H. MeL. & Sommerville, R. 1. (1958) Aust, J,
Science 21, 148-149,

such as those kept.to feed the captive animals at Pallarenda,
If M_ inuris were established in an infective cycle, including
cockroaches and rats Itving in close proximity to the pens,
then such cackroavhes when caten by Musky Rat-kangaroos
could be the link between nomal mdent hosts and the
accidental macropodoid hos, Once ingested M. muris ts
apparently able to establish itself in the.simple non-fermenting
Ral-kanuaroo stomach. The environment here could niore
closely reseroble that of rodent stomach than that of the more
complex fermentative stomach of macropods and potoroos.

This is the first record of M. murixy occurring in a
macropodoid marsupial. The ealy other records of M, mauris
fron) marsupials are [rom a phalangeroid, the Broshtail
possum, Trrehasyrus vidpecula (Kerr, 1792). Worms fram
infected possums were: first described front Queensland)”
and then reported from New South Wiles! as Propaspinera
marsupialis Baylis, 1927. This material was subsequently
reexamined and deterniined to be M. muriy'*,

The diet of the Brushtail possum, including fruits and micat
if offered.2-® is more similar to that of the Musky Rat-
kangaroo than to that of other macropodois. The non-
fermenting environment within the possums stomach is
probably alse similar to that of A. meschatus. Therefore it
is likely that, as with H, moschatus, To vulpecuia could also
become accidentally infected wilh M. muris from time to time,

1 am indebted to P.M, Johnson for the opportunity to
examine the Musky Rat-kangaroo. All the nematodes collected
have been deposited in the Australian Helminthological
Collection, South Australian Maseum, Adelatde.

SQuentin, J. C. (1970) Ann. de Parasitol, 45(6), 839-845.
°Baylis, H. A, (1927) Ann, Mag, Nat. Hist, (95)20, 214-225,
Johnston, T. H. & Mawson, P. M, (1938) Ree. 3, Aust,
Mus. (2), 187-198,

"Baylis, H. A. (1434) Ann. Mag. Nat. Hist. (108)14,
142-153.

2 Johnstan, T. H. & Mawson, P. M. (1939) Trans. R, Soc,
S. Aust. 63(2). 209-209.

UWertheim, G. (1962) Trans. Am- Micros, Soc, 81(3),
274-279.

Mew, R. A (19881 “Comnion Brushtail Possum” pp.
147-148 Jn Strahan, R. (Ed_) “Complete Book of Australian
Mammals” (Collins. Angus & Robertson, Australia)-

L. R. SMALES, Department of Biology, Central Queensland University, Qld 4702.

OTOLITHS AS POTENTIAL INDICATORS OF AGE IN
COMMON CARP, CYPRINUS CARPIO L.
(CYPRINIDAE: TELEOSTED

BRIEF COMMUNICATION

Summary

The common carp, Cyprinus carpio L., was among the first species of fish for which
techniques of age estimation were developed’. The annuli of scales (seasonal zones of
slow growth) have been used as growth indicators in carp from Asia, Europe and
North America”®, Opercular bones®*, fin rays’ and spines'®"' have proven useful, and
the eye lens may also have value” although it is unreliable for older fish’**, Otoliths
have been used successfully for the cyprinid Phoxinus phoxinus’*”’, but not for
carp!!!

Transactions ef the Royal Society of S. Ausi., (1995), W942), 97-98

BRIEF COMMUNICATION

OTOLITHS AS POTENTIAL INDICATORS OF AGE IN COMMON CARP,
CYPRINUS CARPIO L. (CYPRINIDAE: TELEOSTED

The common carp, Cyprinus carpio L., was among the lirst
species of fish for which techniques of uge estimation were
developed!. The annuli or seales (seasonal zones of slow
growth) have been used as growth indicators in carp tron
Asia. Europe and North Aimerica?>, Opercular bones’, fin
vays “and spines!!! have proven useful. and the eve lens may
also haye value! although iis unreliable for older fish.
Otoliths have been used successfully forthe eyprinid Phavinis
phoximes'® 2) bat not for carp! 2!

It is not clear whether the authors considered the full
complement of otoliths in these studies. Carp. and teleost fish
general, have (hree pairs of ulricular. saccular and lagenar
Otoliths, respecuively named the fapilli, sagittic and asterise!
Given the peculiar morphology of The Jower part of the nmer
ear of eyprinids (and other ostariophysan fish), the asterise
ynd the lapilli are much larger than the comparatively thin.
elongate sagittue?? Lapilli have been used to distinguish
daily growth increments in the fllfsh Semrefitis corporaliy’
and the rose biterling Rhiadexs vecellarts necellatis. and
adiurmi rhyihin of valeiim deposition has heen ceported in
the asteriscr of youny poldfish Caragsyitey auras?
Unspecified otoliths have been used to age tench, Titer tinea
in Burope?’, hutitus-likely that these were asterised or lapilli
mither (han sagittae,

fn revent work on the cology of carp We have heen able
consistently co recover well-formed onoliths with patterns thar
upper to represent a chronolopieal record, As validation is
necessunily 4b protmacted procedure we belweve that 4
prehmingry eommunivation i warranted This work hus
special sizniticunce in Austruliy because carp are an
Introduced species thutis widely believed responsible for the
degradation of wetlands throughout the Murray-Darling
Rasin2’. 1 the impact of carp is to be evaluated the ability
to estimmite the ages of individual fish, providing for
ireusurements of wrowth. feceuitment and other popularion
parameters,

Samples for thas study were obtained by wilbnetting
backwaters of the River Murray al Swan Reach and at Gurr
Gurra Lukes near Berri, from January 16 April 1994, Body
weight (to within 0.1 wand fork leneth (can) were measured

is oraentl

heliire recovering the ofoliths und recording (he weights of

the asterisc: (LE mu).

Annuli (translucent bands) were counted on the distal side
of whole asterise When more than three annull were present
the earhescones, particularly (he first and second, were olen
obscured by culaium deposition, When more than 5.6 annul
Were present the outermost Ones were more eusily diseernible
in transverse thin sections (hun in whole otoliths. The lapill
proved Useful only when 2-3 annuli were present, and
otherwise underestimaled the counts trom the asteriscr, The
sigittae showed no recognisable pattern. For these redsons.
aMensct appear to be potconully inere usetul us indicators
of age in this carp population.

In Figures live the asteriscus weight, fork length and body
weight of 63 curp are plotted apainst the numbers of ammuh
on the asteriscus, Strong correlations are evident in cuch plot
(respeenvely, Spearman rank correlation coellicient r= 0.873,
P<O00)2 ro=O851 POON Fo=O842, POON,
sbggesting that the annuli are related to the size und weight
of the fish and the weight of the usteriscus, The pattern of
alternating translucent und opaque bands on the asterisch ts
comparable to that of other fish The lapith may provide
complementary readings. especially where there are few
annuli, We conclude thatthe asterisel. and to 4 lesserextent
the lapilli, may provide a chronological record, Validation
procedures aré now Underway to obtain data from fish of
known uge.

This paper is partola PhD progniny being undertaken by
the senior author, The Division of Fisheries, Department of
Primary Industries, Adelaide provided! u permit to ner fish.
We are grateful to Mr fan Petiman, Institute of Freshwiter
ELeology, England. for library dssistanee, and to Me John
Pillut, Berri, Jor assistanve in the field.

' OG Nea ye

Oroiith welght (ma)

Tak Length (my

Weight tg

Annull

Fig. |. Relationships between the numbers of annuli on the
Otoliths (asteriscl) of carp from the River Murray, Junuary-
April 94, (a) ofolith weight (mg) (b) body size (Fork
Length. nim). (ce) beady weight (wd.

'Carlander, K. D. (1987) pp. 3-14 /n Summerfelt. R. C. &
Hall, G, E. (Eds) “The Age and Growth of Fish” (fowa State
University Press, Ames, Iowa).

2Swee, U. B. & McCrimmon, H. R. (1966) Trans. Am.
Fish. Soc. 95, 372-380.

Crivelli, A. J. (1981) J. Fish. Biol. 18, 271-290.

‘Johal, M. S., Novak, J. & Oliva, O. (1984) Vest. cs.
Spolec. zool. 48, 24-38.

5Fernandez-Delgado, C. (1990) Hydrobiologia 206, 19-28,
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English, T. S. (1952) Iowa State Coll. J. Sci. 24, 527-540,
8Redher, D. D. (1959) Lowa State J. Sci. 34, 11-26.
‘Meunier, F. J. & Pascal, M. (1981/1982) Aquaculture 26,
23-40.

"English, T. S. (1952) Prog. Fish Cult. 14, 36.
''Wichers, W. F. (1976) “Age and Growth of Carp (Cyprinus
carpio) from Pathtinder Reservoir, Wyoming, 1974 and 1975”
(NOAA, National Marine Fisheries Service, Washington,
D.C.).

Carlton, W. G, & Jackson, W. B. (1968) Copeia 3,
633-636.

Crivelli, A. J. (1980) J. Fish Biol. 16, 469-473.
4Boehlert, G. W. (1985) Fish. Bull. 83, 103-117.
SBullough, W. S. (1940) J. Exp. Zool. 85, 475-497,

‘Frost, W. E. (1943) J. Anim. Ecol. 12, 139-162.
"Myllyla, M., Torssonen, M., Pulliainen, E. A. &
Kuusela, K. (1983) Aquilo (Ser. Zool.) 22, 149-156.
'8Mills, C. A, & Eloranta, A. (1985) Ann. Zool, Fennici
22, 1-12.

9SMills, C. A, (1987) Freshw. Biol. 17, 53-67.

20Mills, C. A. (1988) J. Fish Biol. 33, 545-561.

Hume, D. J., Fletcher, A. R. & Morison, A. K. (1983)
“Carp Program Final Report.” (Carp Program Publ. 10,
Ministry for Conservation, Victoria).

22Secor, D. H., Dean, J. M. & Laban, E. H. (1991)
“Manual for Otolith Removal and Preparation for
Microstructural Examination” (Baruch Press. University of
South Carolina).

3Victor, B. C & Brothers E. B. (1982) Can. L Zool. 60,
2543-2550.

24Solomon, G., Matsushita, K., Shimizu, M. & Nose, Y.
(1985) Bull, Jap. Soc. Sci. Fish. 51, 55-62.

25Mugiya, ¥., Watabe, N., Yamada, J., Dean, J. M.,
Dunkelberger, D. G. & Shimizu, M. (1981) Comp. Biochem.
Physiol. 68A, 659-662.

26Wright, R. M. & Giles, N. (1991) J. Fish Biol, 38, 17-28.
27Fletcher, A. R., Morison, A. K. & Hume, D. J. (1985)
Aust. J. Mar. Freshw, Res. 36, 31-327.

LORENZO VILIZZI and KEITH F. WALKER, Department of Zoology, University of Adelaide. S. Aust. 5005

VOL. 119, PARTS 3 & 4
30 NOVEMBER, 1995

Contents

Transactions of the

Royal Society of South
Australia

Incorporated

Li, Q. & McGowran, B. Comments on some southern Australian foraminifera and
description of the new genus Parredicta = - - - - -
Cappo, M. The population biology of the temperate reef fish pons jacks i
in an artificial reef environment -
Schluter, C., Bye, J. A. T. & Harbison, P. The most vigorous South Australian tide
Bird, A. F. Studies on Eutobrilus heptapapillatus (Nematoda: Tobrilidae) the
predominant nematode inhabiting the bottoms of Lakes Albert and
Alexandrina, South Australia = - - - - - - -
Skerratt, L. F., Beveridge, I. & Durette-Desset, M. -C. Distribution of species of
trichostrongyloid nematode parasites in the small intestine of the bush
rat, Rattus fuscipes - - - - - - - - -
Barker, S. Eight new species of Australian Buprestidae (Insecta: Coleoptera) -
Richards, S. J., Burton, T. C., Cunningham, M. J. & Dennis, A. J. A new species
of Callulops from New Guinea and comments on the status of C.
humicola comptus (Zweifel) (Anura: Microhylidae: Asterophryinae)
Stewart, A. C. & Nicholas, W. L. Manunema pectenophora sp. nov. (Peresianidae,
Leptolaimina), a nematode possessing unusual male supplementary
organs - - - - - - - - - - -
Kolesik, P. Asphondylia dodonaeae, a new species of Cecidomyiidae (Diptera)
damaging leaves and branches of hop-bush, Dodonaea viscosa
(Sapindaceae) in Australia - - - - - - - -
Kolesik, P. Contarinia bursariae, a new species of Cecidomyiidae (Diptera) infesting
fruits of sweet bursaria, Bursaria spinosa (Pittosporaceae) in Australia
Williams, D. J. & Brookes, H. M. A review of the scale insect subtribe Andaspidina
(Hemiptera: Coccoidea: Diaspididae) and a new genus, OSes
for two Australian species - - -
Connolly, R. M. Diet of juvenile King George whiting Sillaginodes punctata (Pisces:
Sillaginidae) in the Barker Inlet - Port River estuary, South Australia

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

113
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191

TRANSACTIONS OF THE

ROYAL SOCIETY
OF SOUTH AUSTRALIA

INCORPORATED

VOL. 119, PART 3

COMMENTS ON SOME SOUTHERN AUSTRALIAN
FORAMINIFERA AND DESCRIPTION OF THE
NEW GENUS PARREDICTA

By QIANYA LI* & BRIAN MCGOWRAN*

Summary

Li, Q. & McGowran, B. (1995) Comments on some southern Australian foraminifera
and description of the new genus Parredicta. Tran. R. Soc. S. Aust. 119(3), 99-112, 30
November, 1995.

The distribution and relationship of 20 benthic foraminiferal genera from southern
Australia are reviewed, and ranges of some stratigraphically useful species are
revised. Among these, Crespinella, Crespinina, Hofkerina, Maslinella and Wadella
are endemic to the Australian-New Zealand region. Others contain species which are
either endemic or cosmopolitan forms or those migrating into the region at various
times. The new genus Parredicta is described to include two endemic species,
Planulina kalimnensis Parr (early Miocene-late Pliocene) and Valvulineria porifera
Parr (Pleistocene-Recent).

Key Words: benthic foraminifera, Eocene, Oligocene, Miocene, southern Australia,
Parredicta, new genus.

frnsactions uf the Rowil Seriete ofS. Aids HONS). TIMI). 99112

COMMENTS ON SOME SOUTHERN AUSTRALIAN FORAMINIFERA
AND DESCRIPTION OF THE NEW GENUS PARREDICTA

by QiANYL LI= & BRIAN MCGOWRAN*

Summary

hi Q & Metinwess. B. (1995) Comments on some Southern Australian foraminifera and description of the
new gers Pormdict Tres, Ro Sav. S. Aust W934), 99412, Ah Nevember, 1995

The distribution and relationship of 20 benthic tordminiferd! genera from southern Austraba dire reviewed. and
randes (1 serie stratigeaphieally aseful species are revived. Aimonp these, Creyatuelio, Crespinind, Hufkeriau,
Mastinetla aol Madelly are endemic iw the Auswraliin-New Zealaml region, Others contain specjes which gre
vilber endemic or cosmopolitan forms Or those migrating rato the regioa af sarous tines, The new genus Parredieta
is desevibed to inelude two endentic specits. Plaaalina Kalinmensis Pare (early Miovene-hite Phioveme) and

Falvalineria partiera Pare (Plemtovenc-Recent)

Ky Words) benthic foraminifera, Eocene, Gligoetne, Miovene, sQuthern Austin. Parredion:, new genus.

fntvoduction

Poraminitera are singlescelled protozoiugs widely
employed in Stralizraphy and marine ecology for age-
dutin: and palueoenyvironmental interpreration
Howchin's (1889, 1891) work. which appeared in this
Transactions, laid the Joundation for surveying local
lorminiteral assemblages, Foraniniteral studies (i the
varly part of vhs cenlury in Australia were cultivated
parucukirly by W. J. Purr Like his New Zealand
eounterpurt Ho J. Finlay. Parr published many papers
on recent and fossil toraminitera and supplied
numerous specimens lor J. A. Cushman to deseribe
(Glaessner I950) Recent and modern students,
including Carter (Y58. 1964), Quilty (1974, 1977, 1981,
1982--mainly small benthics) and Chaproniere (I984--
larger beathies), tend to emphasize the furaminileral
biesutigraphic applicauon. as well 4s lineage
classification, Systematic treatments of southern
Australian foraminifera, however. have not yer reached!
the standard of Hornbrook et af. (1989) from New
Zealind. Locvd! marine sequences have been correlated
with standard hidstratigraphy (e.g. MeGowran [979),
but the correlation lacks cross-reference to the
geornagnelic record, and the range ol many species
is not well delined. Confusion aver synanyinis adds
(iM Ulies Lo any Uttempt lor systentitic compilations

As a prelude ty such a compilation, this paper
summarizes current knowledge of some important
Cenozoic taxa bused on material from) several southern
Mustruliin basins. The recerds of these tixa, as
reported elsewhere (Loeblich & “Tappan 1987), are
revised. The new genus Paurredicte is: proposed to
accommodate alvadinena parifera Parr and Planaling
Audimmensis Part,

The Material

The lute Kocene to early Oligocene samples were
taken mainly from two localities, Mastin Bay on the
southwest coast of Adelaide, South Australia and
Browns Creck, Ane Districtin Victoria (Fig. 1). The
Maslin Bay sequence has been deseribed and discussed
in greal detail by MeGowrin & Beecroft (986) and
MeGowrun (1990), and boih sections by MeGowrat
eval. (1992). The Lakes Entrance oil shalt section [rom
Gippsland Basin spanning the late Ohgovene-lute
Midwene was the focus of var study ol faunal overturn
and ccostratigraphy (MeGowran & Li 1993, 1995: Li
& McGowran 1995), and thus forms part of the
material here. We also examined 29 samples [rom the
early middle Miocene Morgan-Cadell section from the
Murray Busin (Ludbrook 1961). As well, we reler to
(he material during d recent biolacies study of dredged
sumples [rom the Lacepede Shell (Li ef al. 1995).
Other material includes random sumples from Castle
Cove (neur Browns Creek) and WMC eore 703 in
Kingston, South Australia, Relevant type specimens
deposited in the South Australian Museum and in the
Department of Geology & Geophysics, The University
of Adelaide, were also examined, Several scanning.
photographs, onginally taken by J. M. Lindsay on
material from the South Australian Department ol
Mines and Enervy, dre also reproduced.

Localities are shown in Fiz, |, and ranges of taxa
discussed in Fig. 2. The generic references to these
taxa refer Lo Loeblich & Tappan (1987). Appendix |
alphabetically lists all genera and species mentioned
m this report.

Systematic Remarks

Order Foraminilerida Eichwald. 1830
Suborder Rotaliina Delage & Herouard, 1896
Family Almaenidue Myatlyuk, }959

Genus Almaena Samoylova, 1940

100 Q, LI & B, McGOWRAN

Synonymy and Type species: see Loeblich & Tappan
1987, p. 622.
Remarks

In their compilation of A/maena, Loeblich & Tappan
(1987) apparently overlooked its record in southern
Australia. From the Gippsland Basin, Carter (1964)
described Almaena gippslandica (Pig. 4, la, b), a form
tindoubtedly belonging in this genus.

Carter (1964) correctly indicated that Almaena
eippslandica Was restriced to the region, as it has to
date never been reported from any other localities. 4
ringes from 347 m ~ 320 m in the Lakes Enlrance
section, in an interval equivalent to planktonic
forannmiferal Subzone N4b, earliest Miovene (Li &
MeGowran 1995),

It is not known whether A. gippslandica is
synonymous with any European taxon or whether it
represents a migratory species from Paratethys where
the genus first evolved in the later Eacene (Loeblich
& Tappan 1987)

Family Asterigerinatidac Reiss, 1963
Genus Asterigerinella Bandy, 1949

Synonymy and T\pe species: see Loeblich & Tappan
987, p. 606.
Remarks

According to Loeblich & Tappan (1987), this genus
differs from the similarly stellate Aseriverinea
d’Orbigny in having a flattened lenticular test and a
high aperture. The wall is papillate on both sides of
the lest, in contrust to the smooth surlace in
Asterigerina. Loeblich & Tappan (1987) found the type
species 4. gallowayi trom Alabanva to be the only
tecord.

Howchin’s (1891) taxon. 7runcatulina margarttiferc
var. adelaidensis, bears every feature of A, gallewayi
and must be a form of Asterigerinella, Lindsay (1969,
pl. 2. fig. 2. 4) illustrated a topotype of 4. adelaidensis
and Lindsay (1985, p. 203, as Asterigerina) indicated
that the species ranged from the top of South Maslin

AUSTRALIA

EUCLA and
GREAT AUST. BIGHT

ST VINCENT

ANTARCTICA

CHRON 19, 42 Ma, MIDDLE EOCENE

Fig, |. Southern Australia in the later middle Eocene, showing major Tertiary sedimentary basins (adapted from Falvey
& Mutter 981), [t was not until this time that sedimentation became widespread along the southern margin. Numbers
(7 are section localities: 1. Lakes Entrance, Victoria, 2. Browns Creek and Castle Cove, Victoria. 3. WMC core 703,
Kingston, South Australia. 4.. Morgan and Cadell, Murray Basin, South Australia, 5. Maslin Bay (Tortachillu, Blanche
Pow), South Australia. 6. Yorke Peninsula (Port Vincent), South Australia. 7, Lacepede Shelf, South Australia, 8. Nanarup,

Western Australia,

SOUTHERN AUSTRALIAN FORAMINIFERA

Sand to Perkana Member of the Blanche Point
Formation, Zones Pi3-P15 in modern biostratigraphical
correlation (McGowran ¢/ al, 1992), This view is
confirmed again here in our recent observations,

AS illustrated in Fig. 4, 2, 3, A. adelaidensis can
he differentiated from the slit-apertured 4. gallawayt
by its almost circular aperture.

Family Bronnimanniidae Loeblich & Tappan, 1984
Genus Bronnimannia Bermidez, 1952

Synonymy and Type species: see Loeblich & Tappan
1987, p. 563.
Remarks

Forms of Bronnimannia are rare in southern
Australia, although the closely related Diseorbinella
und Planulina have been widely reported (Parr 1950,
Carter 1964: Quilty 1977). Our record of B. halioris
from Cadell Marl section (Zones top N8 to lower NO,
carly middle Miocene) thus confirms the occurence
of the genus in the region, Quilty (1994, pers, comm.)
recently informed us that he has found sirnilar forms
in the Swan River estuary, Western Australia.

Bronnimannia haliors (Fig. 4+, 4a, b) is similar to
the type species B. pulmerae in the auricular biconcave
test, Unlike the latter taxon, however, the South

iuran) events
southom Australia
’ a

Win
bec tainncvlitnoonann
wre

geochronnlag
Mo

SCH guvvieTNTUiLane

& refulgens

21d planet
t

JAA id wort tetnre eel

Manna naliate

10
AN urswniunss

eS eu

4 onpace

Lit paunerimerieinea

reulprirr
Pee slr
Fo qyesrrinniss
A Tail
NP) Got jalevanereailula

respinalia uinbonifsta
vee |) Vintdrionsis

MIOCENE
— Aronnn

i dhungla

atir praasalale
Ge Irtynis
oY) ie
eal by noiMelna
tw eT Wy

—. Almaena opsiengics

f= Lig Giedlisiaaen

+. subnaldinger

pie |= Ci Aare

=) Gi mena i uharaaite=
fab
bre raids,
Cory

= 5 nnagporntann

Crespinina kingscotensrs
senees Pscomabovinas cuvillist

C. mediacris

(CO. perforalus

a! Gig disor
5 nadie

"7 vaciawun (Pri

OLIGOCENE

A drevoralis

Gah inate
uo ley meal

se TY cabryaiag (154
A cerned

—— Aslenperineia adeleensis

SC cubist

EOCENE
MIDDLE
Chicidaidas spp
Heterolzea spr

Cjbicides sop

A ante

Australian species is much flatter and Jacks a distinct
marginal keel. The strongly concave, evolute (ventral)
side is coarsely perforate, with limbate. impertorate
sutures. All these suggest that the illustrated [orm is
a distincl, perhaps endemic. species.

Family Cibieididae Cushman, 1927
Genus Cibicides de Montfort, 1808

Synonymy and Type species: see Loeblich & Tappan
1987, p. 582.
Remarks

The cibicidids are one of the most abundant and di-
verse foraminiferal groups found in many parts ol
southern Australia. This group includes. trovhospiral
forms with an extraumbilical aperture which muy ex-
fend around the periphery and onto the spiral (dorsal)
side, With these features. Cibicides, Cibicvidoides and
Heterolepa may be lumped as cibicidids in a classical
study of hiofacies (e.g. Hornihrook ef al 989). Al-
though Loeblich & Tappan (1987) demonstrated differ
ent hyaline walls between Cibicides and Cibicidoides
and classified them in two different superfamilies, these
two genera ure always associated ina faunal commun-
ity and some of their species show transitional charae-
ters, particularly in the flat to convex dorsal side.

loual slages
| Wernkawiar
Fe on
> 3 Kalimnaty
z = .
= a & Cheltennam
€ 4 Sspef se] & -
as Papi = | 2a E Michallian
. 2 eS ae ga 5
5 2 aJSa] bs 5 :
= 2 E =e 5 2 &
= Bs 2a a
* 2 5 3 = z Barmsaahian
[-4 © nu = Sa S
4 e a 5 eS bee
£ 1" | Balcombian_
! id | = Batefordian
= 5
: 5
; 4 6
; = Longtordiary
4 a
4 [=
a
q an fis =a
& Beri -
Nossal ® & w “| .
e = @ 5
2 FI & E = : a danjukian
i ae ee
a = ? g : 3
ah 2 = = = z
= a” & 3 “Ee
es 8 e ; —
2 3 & = 2 : = Willurgar
£2 a a = : 2 eye eet
+= a : 2 :3
_— - Toe. = ¢ Hi
: ~ Aldingan
1
(hat narned)

Fiz. 2. Ranges of the taxa discussed. Geochronology after Cande & Kent (1992) and N/P zones after Berggren e/ ul, U98Sa,
b). Correlation of local planktonic foraminiferal data and regional stages follows McGowran er al, (1971), Heath & McGowran

(984), and McGowran & Li (1993, 1994),

12 0, LI & B, MeGOWRAN

Quilty (982. p. 10) listed over 20 cibieidid species
known from the Tertiary of southern Austmliiun and
New Zculand, Together with the meonyex allied Cibr-
cidiides. the planovonves venus Cibicidles averages
20% - 50% Of total fauna in most suniples, Typical
vibicidid forms tnchide Cibieides thungia (Fiz. 4. 7,
8), GC mediverts (Fig. 4.9, 10), C. vortex (Fig. 4, 6a,
bh), Cibicidoides perforatus (Fig. 4. Wand C. pyeud-
nuverianus (= Cibicides neeperforams) (Fi. 4. 12.
1a). The evolution at ©. pseddoungerianus trom C
peifardns was in the lite Bucene. by a reduction ot
coarse perforations trom both sides (an CL perforarus,)
und restriclion jo the spiral side of the test, The
stranyraphically most useful species is Co karreriformix
Hornibrouk, oecurring in the Oligoeene (Fig, 2), Other
previously described species, such as C. subheidieert
and C. opacis, are now placed in the genus Heterplepe
(see below),

Three cibteidids characterising the modern biolicies
vin Lacepede Shelf, South Austalia, ure Crbivides
refidvens (Fig. 4, 5), C. nweiovriv and Cibieidoides
preudoungerianuy (Li etal, 195),

Fig. 3. Ja, b. Crespinells umboniferd sketches of the holotype
of 2Qperculing wanbonifera Howehin & Purr (1938) (see
also the Sedrining inicrographs in Fig. 4. nes. ia, by. 2,
b, Mastinella chapmani Gluessner & Wade (1959): sketehes
of the holotype. Both types are deposited in the South
Australian Museu, Note that the final chamber on both
qests is missing, bul umbilical openings (uo) sre present.

Family Epomuidae Hotker LWAl
Genus Cresprned/ia Parr 1942

Synonvery arad Type species, see Luebhch & Tappary
N87, p. S7Y,
Remarks

Pary (942) erected Ihe Gully Miocene taxon Oper
culina unibonifera Howehin & Parras (he type species
Of his genus Crespinedla. separating (his simple tarm
from similurly planispirul bot internally comples
Operculina

Crespinella wax monospecihe until Quity (YS)
udded to it another species. C. per’. with alow troch-
ospiral (other than planispiral) coiling. The overall
hrorpholugival similarity between C. uwnbentfera ind
C. parri ted Quilty (980) to imply that both C. parri
and C. umbenifeéra are phylogenetically related, with
C. parri being the predecessor, Loeblich & Tappan
(1987), however, rejected tis statetient on the basis
ol the distinet trochoespiral coiling and supplementary
suluinitl openings i Quilty’s species. Such contusien
over the generic stats at CL pared needs (0 be clarified

Li has Mspected the holotype of Co wabonifera,
which was made availible from the Seuch Australian
Museum, and found that it also possesses dn opening
on the umbilical side (Fig. 3. la, hy Pig. 4. 4a. by.
It is an incomplete specimen wath the final chamber
missing, und a solall opening can be observed at the
buse of the relic part of the missing, chamber, close
ta the margin of the pronounced umbilical boss, Ne
umbilical openings, however. were found related (0 ary
previous chambers. We thus conclude thal the species
C. parri is correctly assigned to Crespinella, a genus
having species with a very low frochospiral to
planispiral coiling and one or tigre supplementary
openings on the umbilical side.

Genus Hefkerinag Chapman & Parr, 193]
Synonymy und Tipe species: see Loeblich & Tappan
1987, p. S3L,

Remarks

Geographically Hofkeriia semicrnara (Pix 9, Wh, 12
is similar to dimaena gippslandica. as both ure
confined to the southeastern corner of southern
continental margin (Carer (958. 1964) Almaena
gippslandica is an earliest Miovene form und
apparenuy has affinities with species trom Paratethys
(see ubove). whereas Hofkerina semiornaiet seems i
be entirely endemic ty the region wilh w pang: Prony
the curly Miocene lo early nniddle Miocene.

itis notewarthy that both Mofkering semiornata andl
Crespinella vmbonifera, above. have a similarly thick
wall, which mimics the wall inthe Eocene Mastinelle
chapmani (see below). Unlike H. semiemata. however,
C. uinbonifera and M. chapmani. have also been
recorded from South Australia and Western Australia
(Quilty 1980, 1981), [tis nol clear whether the thick
wall in these endetme taxa signals a high CaCo,
buildup in local waters during the warming phases of
the later Eocene and eurly-iniddle Miocene.

SOUTHERN AUSTRALIAN FORAMINIFERA 14

i4a wee,

met 21)())1 omer 2001

Fig. 4. Scale bar = 100 pm, unless otherwise indicated, Ta, b, Almaena eippslandica Caner: two views of a single specimen,
latest Oligocene, Lakes Entranee. Sample 156. 2, 3. Averiverinella adelaidensis (Howehin): two specimens, later middle
Eocene. Tortuehilla Limestone, Maslin Bay. Sample AB-Tor, 4a. bo Aronnimannia haliotis (Heron-Allen & Eurland): single
specimen frons the carly middle Miocene, Cadell Marl section, Sample C9. 5. Cibicidey refulgens de Montfort: Recent,
Lacepede Shelt, Sample 89-1, water depth 171m. 60, b. Cibieides vertex Dorreen: single specimen, early Miocene, Lakes
Eniqince, Sample $08, 7,8. Cibicides thangia Finlay, Wo specimens, carly Miocene, Lakes Entrance, Samples 992 and
732.9, 10. Cibicides medineris Finlay: two specimens, late Oligocene and carly Miocene. Lakes Entrance, Samples 828
and 1196, [L, Cthicideidey perforaius (Karrer); late Eocene, Blanche Point Formation, Maslin Bay, Sample A3-091 12,
13. Cibietdoides preudoungerianus (Cushman): two specimens, late Oligocene and eurly Miocene, Lakes Entrance, Samples
1196 and 956, Ida. b. Crespinella umbonifera: lwo views of the uncoated holotype of ?Operculina umbenifera Howehin
& Parr, using a Philips &L20 scunning electron microscope at the University of Adelaide (CEMMSA),

ht Q LI& B. McGOWRAN

P49

es 5)

Fig 5. Scale bar = 100 am, unless otherwise indicated. | Crespinina kingycotensis Wade! axial section, carly Ohgovene,
Port Vincent Limestone, Yorke Peninsula. Sample BSI. 2) Lepideevelina howehini Chapman & Crespin: axial section,
early middle Miocene, lower Morgan Limestone, Mannum. Sample Li/93-1. 3, 4. Exearnebaving cuvillieri (Poinant):;
Wo specimens. carhest Oligocene, SADME bare Ad), western Murray Basin (3), F805, and SADME South Parklands
Bore, Adelude (4), FP 808, both from Lindsay (I98L, pl. 44. figs 1, 3). Note that Lindsay’, 1994 (pers, comm.)
considered the form inno. 4 nota typical specimen of that species, 5, 6, Halkyardia barrrumi Parts wo specunens, late
Eocene, Custle Cove, Sample RIPI 19. 7a. b, Aelerolepa opaca (Carter): single specimen, carly Miocene, Lakes Entrance,
Sample 724. 8-9. Heternlepa hrevoraliy (Carter); (wo specimens, early Miocene. Lakes Entrunee, Samples 788 (8a, b)
and 984 (9). 10a. b. Heleralepe subhendingeri (Parr); single specimen, early Miocene. Lakes Entriince, Sample 852. 11,
12. Hajkerine semiornate (Mowehin): two specimens. curliest Miocene. WMC 703, Samples 45.35 m and 45.65 m

SOUTHERN AUSTRALIAN FORAMINIFERA es

family Chapmaninidae Thalmann. 1938
Genus Crespinine Wade. 1955

Synonymy and Type species. see Loeblich & Tappan
1987, p. 668.
Remarks

This genus, together with its only species C.
kineseotensis (Fig, 5, 1), apparently represents ope of
the numerous laxa endemic to southem Australia. It
hay heen recorded in South Australia (Wade 1955;
Ludbrook 1961). Victoria (Carter 1958) and Western
Australia (Quilty 1981), Quilty (198i) also noted that
lests of C. Aingscotensis became larger and more robust
from cast to west, indicating a warmer temperature
towards the western part of the southern continental
margin.

Crespinina Kugscotensis occurs mainly in the later
middle Egeene to early Olipocene (Wade 1955). In the
Port Vincent Limestone from Yorke Peninsula (Pig
1). Vis associated with some planktonic foraminiterst
such as Guembelitria, an catly Oligocene marker m
local biostraligraphy (McGowran & Beecroft 1985),
ynd its last appearance precedes the first appearance
of Amphistegina. The latter daium in the region was
within the fate Oligocene (Lindsay 1985).

Family Gavelinellidae Hofker, 1956
Genus Excornebovina Butt, 1966

Svnonviny and Type species: see Loeblicly & Tappan
1987, p. 633,

Remarks

Specimens referable to E. cuvilliert were found in
the basal Ettrick Formation (Oligocene) from the
western Murray Basin and eastern St Vincent Basin
(Lindsay 1981'), but this record has never been made
public These specimens were compared with the near-
topotypes of FE. cuvilliert from Escornebéou, France.
supplieu lo Lindsay by Professor C, W. Divoger
(Utrecht), This record thus extends the peographic
distribution of this taxon from Paratethys Lo southern
Australia.

Two of Lindsiy’s specimens are shown in Fig. 5, 4
4.

Furmily Cymbaloporidae Cushman, 1927
Genus Hulkvardia Heron-Allen & Harland. 191%

Svnonviny and Frpe species: see Loeblich & Tappan
987, p. 593.

Remarks

‘The conical Halkyardia barirumi (Fig. 5, 3a, 6) has
been widely recorded in New Zealand from where it

' Livpsay, J. M. (i981) Tertiary Stranyraphy and
Foraminifera of the Adelaide City Area. St Vincent Basin,
South Australia, Unpatl M.Sc. Thesis, The University
of Adee,

was unginally named (Homibrook er al. 1989). In
southern Australia, Ludbrook (1961, as Halkverdia sp.)
found similar forms in the western Murray Basin, and
Quilty (1981) recorded it in the Nanarup Limestone near
Albany, Western Australis (Pig. 1). On the castern
margin of the St Vincent Basin, this species imakes Ewe
brief appearances. in Ihe Tortachilla Limestone and the
basal Port Willunga Formation (Lindsay 1967)
McGowran et al. (1992) recently Correlated these twa
intervals as from top PM to early PIS in the later middle
Eocene, and upper PIS in the early Oligocene
respectively.

No record of this taxén has been reported to date
from the eastern corner of southern Australia,

Family Heterolepidue Gonzales-Donoso, 1969
Genus Heterolepa Franzenau, 1884

Synonway and Type species; see Loeblich & Tappan
\987, p, O32.
Renusrks

Many species of Heicrolepa were previously
recorded as Cibicides in southern Australia. The genus
Heterolepa difters from the radially walled Cibicides
in having a granular wall and an aperture which does
not extend lar onto the spiral side (Loeblich & Tappan
1987). Cibicides breveralis (Fig, 5, 8-9), C. apacis
(Fig. 5, 7a, b) and C, subheidingeri (Fig. 5. Wa, b)
all appear to have these features, and are accordingly
(ranslerved to the genus Hetérolepa.

Also included in this genus is Cibreides victortensiv
(see also Lindsay 1969, 1981), a species confined t>
the middle Miocene, or Zones N9-N1I3 equivalents.
Morphologically, H. victoriensts is similar to both A.
brevoralis and H. subhaidingeri. but differs from the
latter two in the strongly limbate sutures on the spiral
side

At the Morgan-Cadell section, western Murray
Basin, Herervlepa decreases from the lower Morgan
Limestone, disuppears in the Cadel! Marl, and
reappears in the upper Morgan Limestone. The Cadell
Marl is composed mainly of bioskeletons including
abundant milinlid and discorbid foraminifera. and
represents a restricted. but highly productive.
environment, Phe marly sequence is dated at about
}S Ma, in the Jater part of the Miocene chmatic
optimum (Li & MeGawran 1995). Its absence froin
the Cadell Marl indicates that Heierolepu may be an
open marine genus only. in contrast to the ubiquitous
Cihicides,

Family Lepidocyvlindae Scheffen, 1932
Genus Lepidoeyelina Giimbel, 1870

Synonymy and Type species: see Loeblich & Tappan

1987, p. 614,
Remarks
The last oceurrenee of Lepidocyelina sensu lata was

LOG Q, LI & B MeGOWRAN

in the middle Miocene (Zone N9), if dor the late
Miacene or early Pliocene (Adams 1992). This has
heen apparently misquoted 10 he in the Aquitanian (Nd.
eurliest Miocene) by Loeblich & Tappan (1987).
The local representalive of this genus is L. hesvehini
(Fig. 5, 2), a species widely reported fron various
localities in southern Australia (Ludbmok 1961;
Lindsay 1969: Lindsay & Giles 1973; McGowran [979;
Quilty 1982, Chaproniere 1984; Lindsay 1985)
Associated with many other larger forms. ib was
conlined (a the litest early Miocene tu earliest nddle
Miocene, or Zones N¥ and NO equivalents, Us
occurrence in the region has been hailed as a sigrval
of the Miocene climatic optimum (MeGowran 1979:
Frakes e7 a. W987, McGowran & Li 1993, 1995),

Fanvily Linderinidae Loeblich & Tappan, 1984
Genus Linderina Schlumberger. 1893

Swenvew dnd Wyre speciess see Loeblich & Tappan
1987, p. 645.
Remarks

The species Linderma glaessneri is large, discoid
and internally complex with numerous chumberters
(Pig. 6, 1). Like Halkyvardia bartrami, above, it wis
restricted to the central and western parts of the region
and has never been recorded from either Gippsland
or Bass basins in the southeasrern corner. The
stratigraphical oecurrence of Linderina wlaeysneri is
also similar to that of A, partrumi in two short
intervals; later middle Eocene (Zones top Pl4tower
PIS) and earliest Oligocene (upper PI8). This record
thus extends the range of that genus into the early
Oligocene from the originally middle and late bocene
(Quilty 1981).

Family Elphidiidae Galloway, 1993
Genus Parrellinad Thalmann, 195)

Synenvmy und Type species; see Loeblich & Tappan

1987, p. 677.
Remarks

Wade (957) emended Purrellina, a plunispiral
elphidiid which appears to have been restricted to
southern Australian waters during its early evalutionary
history. It first appeared in the middle Oligocene, Zone
P21] equivalents, abour 15 Ma ufter the evolution of its
trochospiral ancestor Nereraralia Finlay,

The New Zealand taxon, Discwrotalia, ts similar to
Parrellina in many morphological aspects except the
distinct evohite spiral side, and both sre believed to
have evolved from the trochospiral Noreratalia
(Eocene-Recent) in the late Oligocene. Ir ty difficult,
however; to separate Parre/ling from Discoratalia, as

same of our Oligocene-early Miocene specimens of

Parrellina vrespinae and Poet imperdrety vend to be
alse Tow-trochospiral (Fig. @& 5-6). ‘Typical planispical
FP iinperdtrit (Fig. 6 7) seems to have decurred only
from the curly Miocene to Recen|. Modern speciniens
SUP Tmperairis Vor offshore southern Australia may
tow at test> Dommi in diameter while is allied form
F verriculata is much smaller and without peripheral
spines. A large. typically plamispinal species existing
in the early to middle Miocene (NG@-NIQ) is P
eraticulatifornis (Fig. 6, 8),

Fainily Siphonidwe Cushman, (927
Genus Siphonndes Cushman, 1927

Syaronyery aud Type species: see Loeblich & Tappan
1987. p. 572.
Remarks

A smooth form deserihed by Huwehin (1889) as
Truncatulind echinaia vat. laevigata is appanently a
Siphoninoiles (Pig. 6 4). Whether the smooth wall
has been subject to the efter of cold waters js not
known, This consistemt feattire gueanlees Ural the
laxon 1s a distine| species, The genere description of
Siphoninoides, as in Loeblich & Tappan (987), should
be revised to embrace this feature.

We found nurnerous specimens of S laewpare in
suinples from the Cadell Marl section, western Murray
Basin (Fig. 1). Theage-of these samples is wilhin Zanes
top N& (o NY equivalenis, early middle Minvene-
Quilly (994, pers: conuy.) mndicuted that uw similar
form exists in the modern Swan River estutiry, Western
Australia,

Family Uvigerinidaé Haeckel, 1894
Genus Siphouvigerina Parr, 1950

Synonyms and Type species: see Lochlich & Tappan
187, p. 525.

Remarks

This genus was supposed to oecur only in the
Holacene (Lochlich & Tappan 1987), However, we
recently discovered torms similar to the type species
S. fimbriata trom (he Lakes Enptranee section.
Gippslind Basin, ina level correlated to the carlest
Miovene,, One of the specimens is ilustrated in Fig.
6, 9). Our record thus extends the punge of this venus
down te the early Mohocene, although the form was
found only sporadically;

Revels (1993) recently found the type specimen of
3. fimbriata ta be biserial throughout, a finding
contrasting the conventional definition of the genus
(e-g Parr 50, Loeblich & Tappan 1987), However,
omany uvigerinid and angulogerinid forms are triserial
intuilly and change to biserial a any later sage, The
triserial part of the test would be difficult to define it
carly chambers ure loosely epiled, a case most likely
CXIStiNE iS. fimibriata,

SOUTHERN AUSTRALIAN FORAMINIFERA {7

Fig. 6. Seale bar = 100 pn, unless otherwise indicated. |. Linderina glaessnert Quilty: late Eocene, Case Cove, Sample
RIE! 19, 2, 3, Mastinelle chapmani Glaessner & Wade. (wo specinens, late Bocene, Adelaide area (Children’s Hospital).
Sample 19.2-19.5 m, FI 955 und FI 956, both trom Lindsay (98T', pL 48, figs. 1, 4), 4. Siphoninoides laevigatus
(Aowehin): later early Miocene. Lower Morgan Limestone, Sample LM2. Sa, b, Parrellina crespinae Cushman: single
specimen, earliest Miocene, Lakes Entrance, Sample 140. 6a, b. Parrellina cl inperatrix (Brady): single specimen,
eurly Miocene, Lakes Entrance, Sample 992. 7, Rarrelfina imperarrix (Brady): Recent, Lacepede Shelf, Sample 89-60.
water depth 82m, 8, Parrediina crancularijormis Wade. hater early Miocene, Lower Morgan Limestone, Sample LM2.
Y. Siphoyenerina fimbriata (Sidebolom): earliest Miocene, Lakes Entrunee, Sample N40, 10. Tbulogenerina ferax (Heron-
Allen & Earland): later early Miocene, Lakes Entrance. Sample 700. 1k. Tibulogenerina mooraboolensts Cushman, later
carly Mivcene, Lakes Entrance, Sample 868.12 Cifel/ia cestata (Heron-Allen & Barland); later early Miocene, Lukes
Entrance, Sample 708. 13. Ficrortella convidea (Rutten): carliest Miocene. WMC 703, Sample 45,05 m. 14. Hadella
hamiltonenyis (Glaessner & Wade); lite Eocene, Blanche Point Formation, Maslin Bay, Sample 099, 15a, b. Parredicta
kalimnensts (Parr): single specimen, ater middle Miocene, Lakes Entrance, Sample 416,

108 O, Lt & Th Mectiwaan

Family Siphogenerinoididae Suidova, 198i
Genus Tubulogenerinag Cushman, {9297

Synanvery ane Type species: see Loeblich & Tappan
1987, p. $20.
Remarks

Gibson (1987. 1989) Gibson et al. 1991) conducted
a series of studies-on the evolution and distribution of
Titbulogenerina and related taxa. Two main
conclusions from His studies are= (1) this genus ranged
from early Encene to Plincene, with Europe being the
site of its first evolution, and (2) species seem to haye
migrated westward from Europe, through the Atlantic.
to Pacific and Indian Oceans, According to Gibson
(1989). mid-latitude Miocene species were largely
confined to the laler early Miocene tw early middle
Miocene. or Zones N6 to N& equivalents.

Quilty (1977) reported T picerubealensix rum the
carly Miovene in Tasrnania, In the Lakes Entrance oul
shaft, we found Liree tubulogenerinines (Fiz. 6, 10-12):
1. ferax, T mooraboolensis and Cifellia costar. The
combined range uf these species is fron 263 ry - 157
m inthe seetion, which is mid-N5 qo early NID in our
cortelation (McGowran & Li 1993, 1995).

We follow Gibson (1989) in considering & vosteater
a tubulogenerimd without a toothplate. Revets (1994),
however, classified Cifellia and Tubulogenerina into
two different superfamilies, on the absence and occur
rence of toothplates in these two genera respectively.
Whether the touthplate ever exists in the early part of
C. costiia is not known, and tittle evidence has heen
found to resolve problents such as the developinent and
reduction or function of foraminiferal toothplates
(Revets 1993)

Family Victoriellidae Chapman & Crespin, (930
Genus Muslinella Glaessner & Wade, 1959

Synonymy end Type species: see Loeblich & Tappan
1087, p. 596,

Remarks

Similar to several other endemic taxa, this zenus is
also monospecific. Mastinella chapmani (Fig, 6, 2.3)
isa large but internally siinple form ranging from the
later middle Eocene to earliest Oligocene. Although
not mentioned jn {he original description, sutural
Openings occur on the umbilical side of some
specimens. (Fig. 6, 2), possibly resulting from relic
apertural extensions. This feature can be seen even in
the holotype. sketched in Fig. 3 (compare Glaessner
& Wade 1959, pl. 1, fig. 7).

Crespinella parri Quilty, above, is morphologically
similar to Masdinella chapmani at least in the
following: (1) a large, low trochospiral test which tends
0 be plamispiral in (he final stage. (2) a. distinet

peripheral keel. (3) sutural openings on the umbilical
side, and (4) a thick. Jantinwtect wall theugl
perlorations on M. chapmeani were much coarser. All
these indivale that C, parr’ is marphologacally. if por
phylovenctically, closely related 19 Maslinella. The
peeurrence of C. parri im the late Gligocene is
cryptogenic, and penuing sidies of its relationship with
M. chapmuni are necessary.

Genus Merariell¢ Chapman é& Crespin, 1930

Synovviny and Type species: see Locblich & Tappan
1987, p, 59.

Remarks

Glaessner & Wade (j¥59) emended this genus and
discussed its affinities, They found the type species
Fictoriella plecte io bea jumor synonym of Carpenteria
convidea, now KF comnidea (Rutten) (Fig, 6, 14). The
total range of K concidea in southern Australia is From
the latest Bucene (P17) tw cartiest Miovene (M4).
Ludbrook (1971, p. 64) noled the tansition of V/
convidea trom Carpenieria hamilionenyis (now Wad-
eller hamiltonensis, see below), in the carliest Olio
cence Glohiverina angiporoides dangiporoides Zone

The Bocene-Oligocene record of (hal species, hiow-
ever, is relatively rare. Only in the latest Oligocene
and earliest Miocene did F conoidea become common
und soulhern Ausiralia-wide, us well as fram north
easiern Australia (Quilty 1993). Tt is conspicuous in
the curbunate-chert association of the Gambjer Lime-
stone in the Otway Basin (G. Moss, 1994, pers
cont),

Genus Wadella Srinivasan, 1966
Syronymny and Type species; see Loeblich & Tappan
1987, p. 596.

Remarks

The genera Wadella and Vieroriella are similarly
large and high trochospiral, However. Mudella
hwniltonensis (Fig, 6, 14) can be distinguished from
F conotdea by its smooth test lacking pillars and less
regular coiling.

In the later middle to late Envene, Hiadelhe
hamiltonensix was one of many large species endemic
to southern Australia and New Zealand. Prior iw the
lute Eocene, in southern Australia, MWadeflu
Aaniltonensiy achieved a wider distribuuoa than (
convidea (Cooper 1979: Quilty 1981; Lindsay 1985)
Inthe Maslin Bay seetion, W hami/tonensis was found
in the Tortachilla Limestone and basal Blanche Point
Formation. in an interval equivalent io Zones upper
Pi4 i PIS (McGowran er al, 1942). Hiidella
glebifarnus also evelved in the late Bocene, and ranged
into the early Miocene, Unlike % humiltonensiy, W
slobifarmis developed a law trochospiral test and
globular chambers)

SOUTHERN AUSTRALIAN FORAMINIFERA 109

Camiuy Bapeinidae Cushman, (927
Genus Parredicta gen. ov

(FIG 7)

Type species: Valvulinerta porifera Parr, 1950

2 3
—_—

Fig. 7. Parredlicta porifera (Parr), Scale bar = 200 po, la-
c, Scanning micrographs of the unvoated halotype of
Vilvadineria paritera Purr, 2. 3. Twa specimens from
Facepede Shelf, Sanyples 49-3 and 491. 7 water depths
123m (2) und I71 om. (3) respectively

Enyirmulaey

This geous is named in honour of W. J. Parr, who
was one of the most influential and prolific
foraminiferal students in southern Australia m the early
part of this.century, and who originally described the
species on which this few genus. is based: edrorue
(Latin = proclamation or decree,

Deseripiton

Test medium to large, low trochospiral, biconvex
chambers high, enlarging regularly, more than 6 1 the
final whorl; 1% to 2% whorls in adult tests, surliace
smooth, sutures radiate to strongly curved, depressed
or flush on ventral side, flush and limbate on the dorsul
(spiral) side; urnbilicus small, depressed or closed with
shell material, but wilhoul a distinct umbilical boss,
periphery narrowly rounded lo weakly keeled: wall
culeareous hyaline, distinctly perforite excepra small
area immediately above the apertural lip: aperture
large, arched or slit-like, extending from periphery to
marginal urea of the umbilicul depression. apernural
lip distinet, regular or irregular: supplemeniary

opembgs copimoen, resoliing trom cither irregular
growth of the lip or relic extension of the aperture on
the uinbjlical side.

Rennttks

This genus differs trom Walvwiineria mm haying an
ovul test outline, angular periphery and supplementary
openings, and lacking apertural Naps. Valvudineria
Cushman has a pronounced apertural flap whieh
proyects over the umbilicus (Loeblich & Tappan 1987,
p. 547), Many species of Madvd/ineria are rounded in
outline, with # distinctly lobate margin which ts broadly
rounded in peripheral view, und have no supplementary
openings On the umbilical side

Parredichas introduced to accommodate two species
which were originally considered as Planutina
kalimnensis Parr (Fig. 6, \Sa, b) and valvalinerta
povifera Parr (Pig, 7). Among others, Carter (1964)
und Quilty (980) recorded Parredicra kalinmensis
(both as Keladineria kelimnensis) in the Miocene of
Victoria and Tasmania, Inthe Lakes Entrance oil shaft.
it was found from 340 1m to the top of sampling level
(63.6.m)...e. the earliest Miocene to late Miocene (Li
& McGowran 1995). The younger oecurrence of P
kalimmensis was veported by Quilty (1985) from the
Pliocene in Flinders tsland, Bass Strail, Parredivte
povifera (Parr), On the olber hand, seems to be a
Quaternary species. On the Lacepede Shelf of South
Australia, 2 poriferda occurs. frequently between 50 m
and 200 m, and some specimens grow up to about 15
nin (height) x Tam (width), with over 15 chambers
in the final whorl (Li er al. 1995).

Quilty (980) sugpested that Crespinella parrt was
the probable ancestor of both C. umbenifera and
kalimnensis. His view is upheld here. The evolution
of this lineage might have begun from C. puri in the
later Oligocene. but the radiation of both Crespinella
wrbonifera and Parredieta kalinuensis did not oceur
until the carly Miocene. This was probably
implemented by vy mearphological change Lron low
trochosprral ty planispiral (Co purr? > C. umbonifera)
and from keeled to weakly keeled or non-keeled (AY,
parri > P. kalimnensis), The loss of the umbilical
Hing (boss) alse took place in the early Miocene and
subsequently became a diagnostic feature in younger
specimens of P kalimnensis and, particularly, in the
much younger 2 purifera (Fig. 7).

Divirifution
Southern Australia, carly Miocene to Recent

Acknowledgments

Amanda Beecroft compiled and scunned most of the
Eocene species, J, M, Lindsay generously allowed the
use of his unpublished data and scanning micrographs,
and he.and P. G. Quilty read an carly draft and shared
with as their Knowledge on the distribution of many
species. The manuscript was reviewed by PG. Quilty
and S$. A, Revet, whase econments are acknowledged
We are indebted to S. Shafik Jor the Lakes Entrance

Liu D&B MecGOWwRAN

material aid ta Y. Bone for dredged samples fram the
Lacepede Shell. G Trevelyan did thin seetuons and AL
Shubber took several optical photographs BL McHelry
arranged the loan af type material from the Scvath

Australian Museum, J, Terletassisted in scaring the
Uncoated Lype specimens, R. Barrett reproduced Figs.
47. This work was supported by an Austrafian
Researeh Couneil crane.

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Appendix 1. A list of

Almaena Samoylova

A. gippslundicn Carter

Amplusteginag dOrbigny

Asterigerinella Bandy

A. gallawayi Bundy

A. adelaidensis (= Truncatilina marsaritifera vat,
udelaidensis Howchin)

Bronnimannia Bermudes

B palmerae (= Discorbis pulmerae Bermiidez)

BK hatiotiys (=Disearbis haliois Heron-Allen &
Harland)

Cifellia Gibson

C. cast (= Chrysalidina costar Heron-Allen &
Eurland)

Cihicides de Monttort

C. thungia Finlay

C. medtoerty Finlay

Cc. refulgens de Montlort

C. vertex Dorreen

Cihicidoides Thalinann

C_ neoperforas Hornibroak

C_ perforatus (=Roralie perforata Karrer)

C. pseudounvertanus =Trauncarulina

peeudoungerianus Cushman)

C. karreriformis Hornibrook

Crespinella Parr

C. parri Quilty

Co umbenifer (=? Operculina umbeanifera Howchin &
Parr)

Crespinina Wade

C. Kingscotensis, Wade

Discerotalia Hornibrook

Eycornebovina Butt

E. euvillieri (= Retalia cuvilliert Porgnant)

Halkyardia Heron-Allen & Earland

HA. bartrumi Part

Reteralepa Franzenau

H. brevoralis (=Cibicides brevoralis Carter)

H, opatca (=Cihicides opacns Carter)

H. subhaidinzert (—€. subhaidingen Parr)

(1985) A Phocene foramin(lerid fauna from blinders
Island, Bass Strait. /hid, 119, 89-91,

— (1993) Tasmantid and Lorn! Howe seamounts:
bioswatigraphy and paltcoceanographic significance
Alcheringa V7, 27-53.

Rivets. 8. A. (199]) The generic revision of the Reussellids
(Foraminiferida). Jour Micropalacaniol, 10. }-15.

(1993) The fonuminiferal toothplate, a review, /bud.
12. 195-168,

Wane. M (1955) A new genus of the Chapmaniminac from
southern Australia, Conte, Cushinun Bound. Foram, Res,
6, 45-44,

(1957) Morphology and kixenomy of the foraminuteral
family Elphidiidite. Jew: Washington Acad, Sei. 47,
330-339

genera and species

H, victoriensis (=Cibieides \Vietoriensis. Chapman,
Parr & Collins)

Hofkerina Chapman & Parr

H, sentiornara (= Pulvinilina semiornata Howehin)

Lepidoecvelina Giinbel

L. howehini Chapman & Crespin

Linderina Schlumberger

L. glaeysneri Quilty

Maslinella Glaessner & Wade

M. chapmani Glaessner & Wade

Nororatalia Finlay

Operculing COrbigny

Parredictat Li & MeGowran

P. kalimnensis (=Planulina kalimnensis Parr)

P. porifera (= bilvudineria porifera Parr)

Parrellina Thalmaun

FP. craticulatifurmis Wade

P. crespinae (= Elphidium crespinue Cushman)

P. imperairix (=Polystomella iimperatrix Brady)

P. verriculata (= Polystamella vericulata Brady)

Siphominaides Cushman

§. laevigata (= Truncatulina echinate var, laevigata
Howchin)

Siphouvigerina Parr

S. fimbriata (= Uvigerina perrecta var. fimbriata
Sidebottam)

Jubnilogenerina Cushman

T, ferox (=Bigenerina ferox Heron-Allen & Earland)

Lo mooraboolensis Cushman

Vicloriella Chapman & Crespin

Ko conoidea (= Curpenteria vanovidea Rutten)

Mictoriella plecte (=Carpenteria proteiformiy var.
plecte Chapman)

Wadella Srinwasan

Meo heamiltonensiy (= Carpenteria
Glaessner & Wade)

Ke globifarmis (= Carpenteria globifarmis Chapman)

hamilionensts

THE POPULATION BIOLOGY OF THE TEMPERATE REEF
FISH CHEILODACTYLUS NIGRIPES IN AN
ARTIFICIAL REEF ENVIRONMENT

By MICHAEL CAPPO*

Summary

Cappo, M. (1995) The population biology of the temperate reef fish Cheilodactylus
nigripes in an artificial reef environment Trans. R. Soc. S. Aust. 119(3), 113-122, 30
November, 1995.

Underwater surveys and observations of tagged fish were used to examine spatial
distribution, temporal variation in abundance, habitat use and agonistic behaviour of a
small population of Magpie Perch, Cheilodactylus nigripes, in a 1176 m* site beneath
a pier over two winters. A marked decline in numbers of small fish in the population
was observed in one year and the number of larger fish was more stable. The
unstratified density of fish was between 1.6 and 3.4 fish 100 m* but locations of fish
sightings were strongly positively correlated with two dimensional cover of hard
substrata within the site. Cheilodactylus nigripes was a diurnally active micro-
carnivore which used hard substrata for shelter and for feeding on benthic
invertebrates. Movement patterns were measured or inferred from spatial patterns of
distribution and were found to be restricted to small areas within the confines of the
pier. Home range was estimated to be 26 m’ for one juvenile fish. Only juveniles < =
12 cm TL defended space aggessively against intrusion by conspecifics and fish >19
cm TL engaged in lateral displays with colour changes in agonistic encounters. These
displays were considered to be related to maintenance of spatial patterns.

Key Words: temperate reef fish, habitat use, agonistic behaviour, feeding,
Cheilodactylus nigripes.

Transeetionsuf the Rav Soeers ofS. Aust (985). EG, 1-2

THE POPULATION BIGLOGY OF THE TEMPERATE REEF FISH
CHEILODACTYLUS NIGRIPES UN AN ARTIFICIAL REEF ENVIRONMENT

by MICHAEL CAPPOF

Summary

Capen. M1. (1995), The population. biology of the teniperate reef fish Cheiledacius migeipes in an artificial reel
envimanment Trans. Ro Sac, & Aur W984), 13-122, 30 November, 1995.

Underwater Surveys and observations ol fagged {ish were used to examine spatial distribution, temporal variilion
imabundance, habit use and agenisne behaviour of a Srull populition of Magpie Pereh, Chedtedecivtay nigripes,
inv 176 m site beneath o pier over wwe winters: A marked decline in numbers of small tish in the population
was observed in one year anu the number of larger fish was mere stable. The unstratified! density OF Tish) was
berween [fr and 3.4 fish (00m? buf locations. of fish sightings were strongly positively correlated with Wwo-
dimensional cover ob hard substrata within the site. Che/ldacilis nigripes was a diurna lly acuive mncro-curnivore
Which usect hurd Substrata for shelter and for feeding om benthic invertehrates Movement patterns were measured
or inferved tron: spatial patterns of disiribulion and were found to be pescicted to smal] areas within the confines
ofthe pier, Home range! was estimated ta be 2h i) far ane juvenile fish, Only juveniles <= cm TL
detended spave aggressively against inuslon by conspecifics and fish > 19 em TL engaged in lateral cisplays
wilh colour chutes (aivonistic encounters. These displays were considercd to he related ty majitendnee of spithal
patlenns.

Replenishment of he pier population (as observed wy oceur in spring fran eeccuininent of Sem TL juveniles
Vhese duta indicat’ the importance of refulively small marine protected dreds as refuges Tron spearfishing for

CO. niripes.

Key Worbs terapetate tect fish, habitat use, agonistic behawinur, feeding, Cheilodacr ius nigrines.

Introduction

Cheilodactyld fishes are a numerically important
component of the cool temperate reel fish funas io
Australia, New Zealand, South Africa, South America
and Jupan (Lincoln Sinith ef ad, W989) Branden er al.
1986. Loum & Chout 980; van der Bist 981; Nielsen
1963; Sano & Moyer 1985). They are relatively linge,
slow moving and can be casily approached underwater.
making therm very popular targets for spearlishers, La
south-eastern Australia (hey doniinate the catehes made
in Spearfishing competitions Johnson 18S, Lincoln
Smith ef al. J989) and there is evidence thitt
spedrfishing is a major cause of lovalised depletion al
chetlodactyhds in New Zealand (Cole et al 1990).

Assessment of effects of recreational speattishing on
cheilodactylid populations requires a knowledge of
habitat use by the fish, (heir pallerns of movement and
abundance und the sources of populuton
replemshient, These data are essential for the
inplementation of marine protected areas (Edy vane
993) at the proper sparial scale as uo means of
managing temperate rect fisheries, A knowledge ot
feeding habits is ulso desirable to determine (he role
of cheilodaetylids ut trophie dynamics and fo predict

r Australian Instijute af Marine Science, PMB 3. lownsville
MC, Old. 4810

any Subsequent Chunges in benthic community structure
which may result fram the effeets oF high fishing
inortality. Their behavioural traits make these fish ideal
subjects for underwater studies yet little is known of
Ihe patterns of distribution und ecology of temperite
vheiloductylids, especially those that inhabit the coastal
reefs of southern Australia.

Cheilodactylus tugripes is an abundant inhabitant ot
shallow (<30 metres) limestone and basaltic reefs in
southern New South Wales, Victona, Tasnkinia, South
Australia and southern Western Australia (Hutchins &
Swainston 1986). It is commonly Jound m assoetation
wilh the hard subs(rata provided by artificial tyre reels.
ship wrecks and piers. This species attains 4) em in
length and rarely takes a baited hook although i
comprises a major portion of the spearfishers’ catch
in South Australia (Johnson 1985a). Limits to the
speared catch are enforced by competition bag limits
and legishited closure of marine reserves und all piers
to spearfishing,

With the exception of counts of C. muripes in surveys
of reel faunas in the Great Australian Bight (Kuiter
1983; Branden er a/. 1986), and frequency in catches
ul Spearfishing competitions (lohayon 185a-b). there
have been no studies of the ecology of this species Mm
Australia.

The present study documents the patterns of spatial
und temporal abundance, habitual use and agonistic
behaviour in a protected populiion of Co migrpes

below a South Australian pier. Specific aims of (he
study Were to!

(1) map the spatial and temporal patterns of abundance
ob the population in relauou i ihe wricrohabitats
provided by the pier,

(2) deseribe the feeding morphulagy and diet of the
species and

(3) deseribe fhe agonistic behaviour amongst
individuals and its importance in the maintenance of
spatial patterns.

Methods

The study site

Edithburgh pier is located on the woslern side of Gult
St Vincent, South Australia at 35.5°S, 137.45°R (Fig.
1), The seabed below the pier sloped seaward tou
maxinun depth ol approximately 4.5 m at low tide
und consisted of a base of coarse sands, gravels und
shell, The predominant hard substrata were artificially
placed, inthe form of fallen pier niaterials, discarded
ship's ballast and debris and w limestone shelf produced

“T T

atl) he Js 138 As
| rae }
\
| SS VaR
Spencer Sul

Gulf Vincent

Adelaide

Edithburgh

Kangaroo Island
— 35 N
t o kin) tDQ
) N B.
f outer half of pier
pt oO) zo

Fig, 1. Location and aspect of Edithburgh pier in Gulf St
Vincent, South Australia,

M CATO

by dredyiny along the northern side. Large expanses
Of seagrasses Surrounded (he structure.

The configuration of pylons and major features vt
the hard substrate below the pier were mapped (Fig.
2). The entire picr was 168 metres long bounded hy
53 rows of timber pylons in 4-5 columns, The study
site Was under the outer half of the pier seaward of
row 23 (Fig, 1). Tt ecomprised un area of 1176 my and
was divided into 120 quadrats from 9-I] m° in area
with relerence to the grid of pylon rows and colunms
(Fig. 2).

Topogtaphic complexity (Leum & Choat 980) was
described for the site by estimating the (wo dimensional
“cover” of hard substrata within eavh quadrat, using
three ordinal categories of topographic complexity. A
total of ROS min Sl quadrats was “simple”
(cover =10%), 225 m° in 24 quadrats had
“middle” level of cover (cover = 11-25%) and 147m
in 15 quadrats were classified us “complex”
(cover’> 25%). The seaward end and the southern side

of the pier had the most hard substrata, in the form

of boulders, slabs and blocks thal were usually less
(han 0.8 m high,

Disrrihurion

Seven visual censuses of the population were nade
during March-September 1980 using SCUBA, On each
census (he whole habilal was searched und the positiun
of each fish recorded. The total length (TL) of euch
fish was estimated lo the nearest centimetre and a pote
was thade if the fish were feeding wheo sighted, Fish
were approached to within obe metre or less and length
estimates Were frequently made in direct comparison
to a 40 em plastic ruler, Four censuses of the size
frequency under the pier were made by another diver
in April-July 1981 and fish lengths were estimated to
the nearest centimetre (K, Wehr unpubl. data),

In analyses fish lengths (TL) were categorised as:
R recruits (<6 em); Cl (6-12 em); C2 (13-19 em), C3
(20-26 om), C4 (>26 em). The choice of distinction
between Cl and C2 was made to separate youny-ol*
the-year from older fish. These were biologically
meaning tul divisions Of the population, a8 colour and
morphological differences occurred amongst them, The
usual colour pattern of Co nferipes was three broad,
verticul, dark bands on the white background of the
body and caudal peduncle and a dark cheek stripe
through the eye (Figs 3a, 4a), Recruits had a bright
orange caudal fin and pale orange caudal peduncle with
black tips on the catidal lobes. The mouth was more
terminal than inferior and angled slightly apward
giving these lish u shorter snout and deeper chin than
larger fish, Size class Cl fish had the same caudal fin
colouration but this faded with size to dusky or reddish
in larger classes. There were protuberant crests on the
preorbital bane of the two largest size classes which
were not observed an sinaller fish,

POPULATION BIOLOGY OF MAGPIE PERCH 115

Fiz. 3 Summary of colour changes displayed by large C
nigripes. a. Normal banding pattern. b. Transitional
banding pattern, c, Modified banding pattern - note dark
midriff band and caudal pedunele, loss of posterior and
cheek bands, white iris and white demarcation between dark
bands.

Feeding habits

To avoid sacrificing the small population in the study
site a sample of 21 Magpie Perch (TL 21-36 cm) was
speared on 25 September 1980 under Port Giles pier
12 km NNE of Edithburgh pier (Fig. 1). Immediately
afer capture the alimentary tracts were removed and
preserved. The food bolus in each tract was dispersed
evenly in water and taxa in four field views (7) were
identified as far as possible, The yolume of each taxon
in the pooled contents of the alimentary {racts was
estimated by a water displacement method,

Observation of mavement and beluaviour
Underwater behavioural observations were made 1
60-minute periods at dawn, midday, dusk and midnight
on five days during August-September 1980 from a
single vantage point amongst limestone slabs at the
seaward end of the pier in pylon row 53 (Fig. 2).

Fig. 2. Comparison of pooled sightings of C. nigripes in 1980
and.an index of topographic complexity in quadrats (right)
with the mapped study site (left), Pylon rows are numbered
in ascending order seaward from row 23. “Simple” cover
of hard substrata< =10% (no shading): “middle” cover
11-25% (mild shading); “complex” cover >25% (dark
shading)

110 M. CAPPO

q i 3 ]
_ _ 1
_— = a
‘ . _ _ 2 7
: ]
—. _. _ _ -
is | ]

Fig. 4. Photographs of colour change and Jateral displays. a. Normal banding pattern. b, Loss of posterior dark band. e.
Transitional banding pattern, d, Circling and leaning displays. ¢. Anti-parallel orientation and modified banding. f Break

in encounter and transitional banding.

POPULATION BIOLOGY OF MAGPIE PERCH Wy

Observations began belore sunrise and finished alter
sunset lo (nelude crepuscular periods of activity. The
movernents of four fish at this location were monitored
simultaneously during the dawn observation periods.
Each fish was recognised individually by tags or body
markings. Further observations were made at a variety
of locations on an opportunistic basis during 1981-1987.

Data on the movement patterns of tagged individuals
were collected during 1980, Seventeen fish were tagged
with “T bar" anchor tags individually coded with
coloured paints. Fish were captured with a large hand-
et. tagged underwater, measured and released
immediately at the site of capture, and a numbered
stake was used to mark each release site. During
subsequent dives the sightings of tagged fish were
recorded on the site map, The larger fish in the
population evaded capture and the tagged sample (TL
10.5-23,0 em) did not include C4 fish,

Results

Temporal changes in abundance and size composition

The sightings of fish of the five size classes are
shown for euch sampling date in Fig. 5. The mean
numbers of total sightings for each census were similar
for the two years with 24,85 +4.30 fish sighted in 1980
(95% CL = 19,32-40,37) and 30,504 1.55 fish in 1981
(95% Cl = 25.55-35.44), but seasonal declines. in
sivhtings were different.

In 1980 sightings declined from a Mareh high of 53
fish to an August low of 18 (Fig. 5). This was partly

Tor! Langth
C1 c4 326cm

Ea ca 20-26cm

50
g (T] G2 19-196
£
oa
‘wo 40 Gl 6-12¢em
oo
2
‘5 30
g
E 20
3
c
10

census dale 198)

Fig. 5. Changes amongst months in (he number of five size
classes of Co onigripes at Edithburgh pier during
Underwater visual censuses in 980 ant 198!

due to the low visibility (< =0.75 m) encountered
during (he August census, The decline was evident tor
all size classes from March to April when both Cl and
C2 declined by one half. The steady decline in Cl
numbers may be partly accounted for by growth into
the C2 size class which showed an increase in abun-
dance. Recruits were first observed in early September
1980. and increased by late September (Figs 5, 6).
Predation may play a role in size-specific mortality as
a Southern Calamari squid Sepioteuthiy australis

20 March '80

25 April '80
a | |

td June ‘80
os i
44 July '80

{2 August '80

2 Saptember '80

26 Saptembar '80

Number of fish sighted
=
=
=
—

4 6 8 10 12 14 16 18 20 22 24 26 26 30 32 M36

| 1 | April ‘81
a ale oll obese

f May '81
TIA eee a

th. 0) ok bo”

duly ‘81

4 6 B 10 12 14 16 18 2 22 24 26 28 30 32 M4 36

Total length (cm)

Fiv, 6, Modal progression in estimated total length of C
nigripey sighted at Edithburgh pier during WSO and 1981.

I) MO CAPRO

Was Observed to make several atlaeks On some Hewly
recruited juveriles before capturing and consumme one
in 1982

This patter was not seen in the 198] winter surveys
wthough there was a slight decline in the proportion
of class Cl fish from April to July. The tune low. was
attributable purtly Wi low visibility (<=0,6 im)
encountered during that census. In both years the
winter papulation counts were al comparative levels
in the study site, Changes in abundance were relatively
sinall in the two largest size clusses amongst months,
and between years (Fiz. 5). However, the sample sizes
and numbers of fish were low, precludiag meaning ful
stufishicul comparisons. Caution is therefore used i
the interpretiGen of these data,

Modal progression is evident in Fig. 6 for the smuller
lish from. 1980 to 198] showing maximum growth rates
of ybout 0.46 em month! for Cl fish and about 0.77
cm month’ for recruits. These are in general
agreement with the measured rate of 0.46 cm month’
growth of a tagged fish from 13.7 em to 15.0 em over
five months in 1980. However, the aveuraey und
precision of length estimation Were noldetermined lor
either observer so further estimation of growth from
modal progression was not possible.

Spatial distribution

The overall density of fish in the study area was
estimated al J.o4-3,.43 fish per 100 min 1980 using
95% confidence limits for the mean number of Jish
sighted divided by the area surveyed. Localised
densities were much higher and (he quinber of ©
figripes sighted in quadrats under the pier wus
positively associated (Spearman R=0.5487, P= 0.000)
with the amount of bard substrtta within quadrats using
Spearnian’s rank correlation procedure (Zar J9R4)_
Fiflzen “complex” quadrats. comprised only 2.5% of
the study aren but gave 5596 of Sighlings compared to
30% in “middle” quadrats and only 15% in the
remaining 68% of the study area defined as “simple”
(Pig. 2). Nearly 18%, ofall sightings were made in one
quadrat on the scaward end of the prer where large
limestone slabs and other hard substrata occurred
These slabs. were the highest (0.6 - 1.2 m) afd aiost
rugose hard substrata in the study area and provided
extensive sheller (Fig. 2), Fish were rarely sighted in
quadrats with 10% or less of bard substrai. with the
exception of quadrats contaming tall three-dimensional
structdres not well described by the two-dimensional
method used to classify quadrats. For example, clusters
of buffer pylons.on the seaward corners ol the pier were’
used together by enerusuing organisms and provided
shelter siles,

There was no clear relationship between water depth
and fish size due to the shallow nature uf the habitat
and the overriding infloence of three-dimensioral reel
structure. However, the largest fish were generally

restricted to the deepest Water and were nor seen in
the area siloreward ol pier row St (Fig. 2) New
recruils were seen thronghput the study wea. hut
mainly dear the seu xurhive ont pylons at (he deepest
seaward end, amongst arhoreseent bryozoans,

Movement of fish in the smely area

Qbservuuions throughout the day and in jhe night
showed that individual Co nivrineys emerged Frew
shelter sites just after daw and retreated to the sante
sites during sunset. No fish were seen io he aetive al
night. In five might dives ove of the largest fish ithe
population were seen in the same sheler sie in a
crevice beLween close pylons. resting morianless un
vutstreiched pectoral fins with raised spinous dorsal
fins. ILappesared thal largest fish emerged earlier and
retreated later than smailer sive oclasses, The
movements of three fish Were moyilofed in each of tie
five dawn observation periods but the fourth and
smallest (CV) sb was only seen for the first three days.
The largest Fishy (C4) traversed an uverdge of 6.04 14)
metres, a 23.0 om tagged fish moved 78+0.37 metres
uid a 14.0 ein tagged fish nreved 2.5 40.55 metres nt
the 61 minules afer eniergenee frum their individual
shelter sites, The smallest individual moved only
OSSIAN metres Only the langest fish moved Jar Irom
the confines af the pier foraging amongst dow
limestung shelves within JO menes of the seaward end

Seven of the mags (41%) were nor seen after
applivauion, Inree orbers persisted for unity 9 days. atid
only five tips were sighted after Of divs The
movement of lane Mayple Perch could fut fe assessed
trom the tageing program because lhe mean length it
release (ir ihese remaining tive fish was only 15.4 em
(C1,€2) and the largest was 23.0 on (C3). Only four
tagged fish were present at the end of the 1981) study
period and euch of these fish was seen during evers
cersusy Mllowing lagging. The loss of lags from the
population was at least panty due to tag shedding On
two oceasions fish were observed to ¥eruh the tag ol!
agains! hard abjects, The sightings of all tagged tish
were restricted fe the habytat underneath the pler within
Timeines of the tagging site wilh the exceplion of We
individuals. Several Weeks after Lage ing these two sly
were fotind to be resident for |he remainder of Ihe stutly
in Quads about 10 metoes away fon the tagping site.
Tagged Nish were never seen to traverse thie seagrass
beds on the siuthern Sidé of the pier which appedres!
tu det as natural boundaries to the pier habirat

Wilson (1975) defined “home range” to be the ares
that an aairial lems thoroughly and habitually patrats,
and “core red” to be the area of Ieaviest regular use
within the home range Only a single tipged Tish was
seen frequently enough to confidently estimate these
Iwo areas. This smull C2 fish (13.7 cm) was sighted
18 umes Wethin 7 metres of the tagging site. The hame
range of this fish was estimated to be only 26 ny hy
Measuring the area ota polygon joining fhe outermost

POPULATION BIOLOGY OF MAGEE PERCIT 1h%

sighungs (Leum & Choar I980). Similarly, the corm
fired Was estinwied at 17 nv ja be the area
encompassing 50% of sightings. The focus of this area
was a sidall shelter Site within a cavern amongst
limestone blacks on the southern side of the pier in
pylon row number 28 (Fig, 2). This fish was only
sighted once outside the shaded confines of the pir
and the furthest displacements were northward under
the pier and westward along the pylons.

Feeding morpholaay and diet

Cheilodacrylus Wenpes bad a small moulli wilh tick
fleshy lips anda siogle row of widely spaged peg-lpke
teeth on the centary and premuxiilary. In the throat
there were a paje of upper, and a single lower
pharyngeal (ooth pauls covered with bands of villiform
vecth. ‘There were IS tong and fine gillrakers on the
Hirst will arch and the stomach was small with a large
pylone region containing five short pyloric caeen,

The fish were ohserved to inspeet closely poekets
al sediment fa erevices or amongst fronds ol
arborescent arganisms and fed jn a pecking mation
The mouth was rapidly opened. lorming a suction with
(he fleshy lips, and benthos was ingested with an
audible clicking sound, By rapidly openime und closing
jhe opercula, Fine silt way strained oul rhroaugh the gill
chambers and larger particles were ejected froyn the
moudt. Large polychactes were wrenched trom the
substratum und vigorously shaken to break thent up
inky pieces suitable for swallowing. When elose
observations were thade, no evidence of prey was seen
near the lish and they appeayed to select fecding
substrata, but not the henthic organisms within,
although fish directed repeated feeding “peeks” at large
prey such us polychaetes once they were detected.

The fish fed mainly on benthic invertebrate jaune
wilh gammarid umphipeds about 4+ mm long
predominuat in the pooled sample, Ol the 60'% of tood
volume identitable, the fourteen major taxa were
ranked as: gammuarids (25.6%): Polychaeta (10%);
ostracods (7.5%), Bivalvia (6.6%), Brachyura (3.2% 1:
Mysidacea (2.7%); and. Tammidacea, Arehaengastriy-
pada. Polyplacaphora. Gastropoda, Isopoda, Ophiuroi-
dea, and caprellid and tubiculous ampiipods (each Jess
than 1%}.

There did not appear to be any size-related diflerence
in the feeding behaviour of C vigrpes with the except
ion that small fish were observed to feed mure ofen
than larger ones, OF all sightings made in |981 the
Jollawing propertions were engaged in feeding when
sighted: R79: Cl 76%; C2 79%, CI 49% C4 ASG

Awonistic hehavianr

Wilson (1975) defined agonistic behaviout to be any
activiw related to fighting, whether ugeression or
caneihation and retreal. Agonistie behaviour in ©
nigripes Was difeeted toward only conspectics of a

similar Size and inree sizesspeciiic patterns were
deseribed for the smallest and largest size classes.

The grallest fish (< = 12 cm), elassified as recruits
(R) ant Cl. aggressively defended space. Most
common was a Short pursuit of incoming Ror! fish
away trom a foraging area Also observed was the
head-on spprouch of Cl fish te within about 4 em
followed by sustained pursuit, darting in small circles
with the dorsal fin fully raised. Biting was evident as
audible sounds and tai! damage from broken caudal
rays. This was termed “carausel fighting” by Chiszur
(1978).

Three stow lateral display palterns were observed
amonest fish more than about (9 cm long (C3, C4)
which did not involve such pursuit. ‘Uhe first involved
pairs of 3 {ish which approached each other head-
onand met head-to-tail in a parallel orientation, often
within centimetres, and one or bath fish tilted upward
slowly. The two fish then swam in slow circles in close
proximity with seme lateral displays. but ne colour
chanye, belore parting,

At claburation of this theme occurred during (he
approach of the large C4 fish when a characteristic
colour Change occurred. The posterior band began to
pale und the white midriff darkened with the
development of a sharp white line between them (Pigs
3b.c, ded). When the fish met they assumed un anti-
parallel orientation separated hy nly 4 few centimetres
for abyut IS seconds, during which they slowly tilted
30 degrees (oa head-up, tail-down position (Fig. de)
The posterior band blanched and the darkening of the
midnlf and caudal peduncle deepened, Other colour
changes were also striking. including blanching of the
anal fin, eheck band, iris and peeroral fins. and
definilion of a biack patch around the pectoral axil (Fig.
3h.c). When one fish withdrew the other followed and
both began laceral “leaning displays” wilh lowered or
raised dorsal spines, described by Chiszar (1978) as
defensive posturing (Fig. 4d). The entire sequence
lasted one lowe minates and once the fish broke off
ihe encounter the midriff band quickly resumed its
former white state tml the other bunds were slower to
return (Fig. 4e.!).

These displays were cleat) identified to occur When
fish met and appeared &) relate to the position of the
fish within the habilai. although it was not possible
io define the home ranges and spatial boundaries
involved in eliciting the agonistic behaviour. The
sipnificance of a third type of hebaviour was more
obscure and involved the loose aggregation of large fish
into a slowly circling group above the seabed. Most
of the circling fish had a pale posterior band and some
of the fish exhibited a leaning display towards others,
‘There was insufficient intormation lo recognise charac
teristics of “winners” or “losers” and no atlempt was
rade t) identify the sex of the participams. but these
lateral displivs were considered to relate to habia use.

Ae) M CAPO

Discussion

The low. variability in-counts of C nivripes dimicnest
most months in winters of two yeurs. the very close
association bewween fish Sightings. and ropogruphic
complexity and the restricled movements of ttsved fish
were all evidence fora high degree of site specificity
oC. nigripes under the Bdithburgh pier. Ruyose, hard
substrara were used as sleeping und sheltering sites and
us lecding substrata and the eopeept of a scall home
minge may best desaibe the use Ob this Space by C
nrertyqes

These patterns of habitat ase indicute thal relatively
small urtificial habitats such ay piers can aer as
Important marine protected areas for this speeics.
Murine protected! areas can full] a numberof important
finetions in fisheries management including protection
of “eriteal habiuty” and provision of areas for stock

replenishment, lor fishery-independen| monitoring of

stock Moctuations and for resolution of conflict amongst
Competitors tor use of manne resources and hubitats.
(Edyvane 1993), Planning the spatial scale and habilal
composition of marine protected areas tor lemperale
reef fishes. requires knowledge of sources of population
replenishment, ontogenetic movement patterns, home
hinge size, habitat requirements and natural habitat
houndaries. Some of these can be inferred for ©
menpes fromthe simple observations presented here
supported by comparison with other detailed studies
ol {he eheiloduerylids associated with tees

Pisiburion and mavement

In translation of the pier observations to natural reef

populations of C. nigripes in South Australia i is
essential to recognise that major biological features at
the habitat have been shown to alfeet the distribution
oftenperate reef fish ata variety of spatial scales and
(hese patterns have been maintained over lore line
scales Jones 988: McCormick l989b), In cis regard
the artiticht buture ot the pier habitat us considered
to differ from nearby natural rects in two tain ways,
Unlike the algal-dominated feels surveyed by Branden
etal. (W986), the habitat beneath Edithburzh pier lacked
macrophyte cover, perhaps because of shading, There
Way thes be more suitable leeding substrata there tor
C) nipripes. as Choat & Ayling 987) found that larger
curmivoraus feel fishes, including cheilodaety lids,
forage preferentially in open reef arcas which support
greater densities of their invertebrate prey in
cornparison to areas dominated by laminarivn algae.
The amount of habit for feeding and refuge is further
enhanced under the pier by the presence of the pylons
und the Jouling communities that encrust them.
Secondly, the density of C. rigripes beneath the pier
(L6-3.4 per 100 m’) was six-fold higher than an
estimate for unexploited reefs oF the Great Australian
Bight (0.2 - 0.6 per WO mr) ealediated from the
survey data in Branden eval. (986), The tome range

Aves. overnent parieris ard agentistre hehaviewy oti
tatu! ceers nay be diflerent sis a Consequence,

The microhabliat requirements ot © aieripes. tor
Shicller sites and feeding subsimara were nor deseribed
wilh the simple habia vlassifieations used here bot
co determined by studving assoenujene berween
ubundunve lind habirat at syall spatial seates,
MoCormick & Choat (987) strilified estimates of
density of (he morwong Cheiladaetvis specrbitis in
New Zealand. by ten habicar types and depih, un
reprinted iverages af 0.25-2.09 fish per WOU nr. with
the exeepion of the wpugraphieally complex Yumble
boulderhank” habital where the density-wits [S87 Lish
pet WOW

There is wlsira clear role for ontogenetic movements
lung onviranmentil gradients in establishing patterns
of chejoddetylid abuindanee amd these should he
consmered in selection of reef areas for protection,
Atier First necruiuig in surge zones (Leum & Choat
VSO) eheiuductylids are known te eve to
prigressively deeper parts of the reet habitat as they
grow (Sane de Moyer 985; McCormick 1989uch)
Although the seagrass beds uround rhe pier appeurec
fo wet us habrur boundaries whieh smaller © végripes
did’ aot traverse. ik Was NOL pussihle to describe
WaMigratiin snd eibiteation of fish with (he simple
tech iydes Used in the study, Replenishrent ofthe pier
Populishon was observed to occur only through the
spring arrival of new reevults.

The auttiniy decline in nulnbers of these young-ol
the-vear io 1880 wis not abserved in I9K) censuses.
und i was nol possible to tesolve the roles af size
specific natural rortlitv. lageindueed mortuliry ar
counting biases tn the decline af such a sil
population. Properly replicated censuses stratified ley
identify Individual fish and deleet counting biases ane
diurnal and sedsonal diflerences in activity. sould help
clarify these temporal changes. The pier map and
census data presented here provide a buscline for future
surveys Lo examine long-tertiy variations in pallerns of
Ubundance oF Co aigripes al Edithburgh,

iis possible that Home ranges and movements or
natural reefs may be mure extensive amongst ull size
Clusses of C. nigripes, und it unknown whether the
Wistributions yeported here were more, ar less
restvicted diirmg warmer months outside the study
period, Leum & Choal (M980) uttribured signifivarit
winter declines in oumbers of ©. spectabilis sighted
to un extension of home ninge during cooler meanths
The estimures of hone range Jor Co aleripes were
relarively Strall in comparison to those constructed for
C speerabihs by Leuny & Choat (IY80) an nalural reels
and Qray depend on fish density aswell as habitat type
Juvenile C) syectabiliy hud home ranges =< = 100 m_
which were abuut three times that estimated here for
Co nripes ustig the same technique. Sinntirly the
jurgest Co vcgnpey were always sighted jp i relanvede

POPLLATION BIDLOGY OF MAGPIE PERCH (21

small atea atthe seaward end of tte Eaithbuesh pier
but Leum & Chout (980) observed that larger C
spectabilis moved large distances and had home ranges
up to 50-70 thousaral tm’.

Fevding leabits

Cheladactvius nigeipes were diumally active,
bentine cariivores feeding mainly on panunaridean
amphipods and other small benthic invertebnites, Thetr
mode of feeding is common ty ther reel-associnted
morwongs tn the same genus, Bell (1979) reported (hat
C. fuseus and C. speectabilis use ther thick fleshy lips
to Wrench and suck aninmils off rhe substratum. mainly
polychaeles, brachyutans, amphipods, gastrapaus, and
bivalves. Sano & Moyer (1485) reported that the
Japanese OC. cebra deeds mainly on epifouna. especially
simmardean Jimphipods and decapods, while the
syinpattic C. sonarus tends to wike both cpilauna
(mainly gammaridean wnphipods, isopods, spanges
und decapods) and intiunal polvchuetes.

Agunistic beltavieu

Tivenile CL agripes <= 12 em'TL were observed
(defend space ageressively, but such defence muy bave
been cnergelicully uneconomic lor laeer fish >t en
TL oceupying lurger home ranges, Lh is proposed here
that the lateral displays and colour-change during
agonistic encounters amongst larger C. aigripes were
related to the muinténance of some Undefined spatial
pallernt of overlapping horne ranges, Such patterns were
mapped by Lean: & Choat U980) for C speviabils
Which directed agonistic behaviour only towards
conspecifics of similar sve und only smaller size
classes Vigerously defended space outside ol the
spawning season. The habitat of C, spertabifis was
described as a mosaic of exclusive lertofies occupied
by sumaller fish through whieh the larger size classes
foraged in larger, overlapping home ranges (Leam &
Chout 980) and this may be a uselul model for turure
studies of habitat use by ©, pleripes

Using Chisear’s (1978) definitions of lateral displays
i agonistic behaviour the description of “gariusel
fighting” fits well the behaviour of CL nigripes
javeniles Whereas the varions colour phases of larger
fish cun be interpreted as varying degrees of threat in
a typical “colour fight” The anti-parallel orientation
adopted during these reciprocal lateral displays is
widespread in fishes, and some species wilh long dopsal
spines, such as the chaetudontids are reported to vilt
the raised spines towards the uther lsh ina defetisive
posture culled “rolling” or “leaning” if it approwches
to closely (Chiszar 178)

The possibihily ol a reproductive basis for same.
ngonishc behaviours cannat be discounted for all
observations of ©. vjeripes, as McCormick (9898)
found that darge male C. apectabilis ipgressively
defended territorwes during the spawning season by
“rolling” dow) On to Totriders Jind restrained visiting

fermales by “ight civeling, pursait and blocking” ot
chasing and tail-nipping. Seme of the lamer C nigripes

al Edithburgh pier wete observed to have fentacular

protuberances on the prearbital hones which have been
used in sthdying sexual dimorphism and separating the
seacs of CL spectabiliy and C\ fiseus hy external char-
acters in visual counts (MeCormick 19893; Schroeder
et al. 1994), The high population densities of C.
nigripes ac some deeper South Austrilism piers may
provide the best chances of clarifying the spatial and
sexual significance of agonistic behaviour of C. nigripes
through observation aad tiorphometric studies.

Fulnte research

The resihence ol C. vigripes to spearfishing and
recuvery Of depleted populations depend on growth
rates and the soufees abd sate of population
replenistiment, This study sugeests that widespread
mavement of C vignpes amongst habitats is not an
Important source of replenishment but further studies
aL uppropriate scules are necessary to determine the
contributions of recruilment and post-recruitment
processes in determining spatial patterns of abundance.
These data are needed to determine iF marine protected
areas should include shoreline surge zones as
recruitment sites with corridors of hard substrata
linking them to adjacent deeper reef, or if isolated
offshore habitats such ws artificial reets are adequate.

The magnitude and frequeney of changes in
population structure are likely ta be directly related
the longevity of © aigripes as variable recruitment
will have least elect and spearfishing the greatest effect
on the population size of long lived species. When
fishing mortality is absent in such cases age classes
accumulate and terapore! consistency in population size
may misk jin underlying instability in the age
composition (Jones M988) Consequently. future studies
of ©. nigripes population dynamics may require
uualyses of age camposilions of unexploited
populations im Conjunction with recrustment surveys
(Doherty & Fowler 1994) and monitoring of the
survival, growth and movement of individually
recognisable (ist from ume of recruitment in
permanent quadrats (Connell & Jones 1991). The
results presented here forin a basis for such studies und
for longer-term assessment ol temporal consistency in
{he patterns of abundance of C_ nigripes in a mapped
habilat,

Acknowledgments

Most of this study was undertaken whilst the author
was a BSe(Hons) student in the Department of
Zoology, University of Adelaide. A.J. Butler provided
supervision, logistic and financial support and use ol
the Coobowie Marine Research Station. The author
also gratefully acknowledges the contributions of K,
Weln for use of unpublished data. C, Proctor and A,

122 M, CAPPO

Davis lor assistance with diving and revision of the
pier map. and M, Keough, K. Branden and K. Handley
for photography. The comments of M. McCormick,

1. Suthers, D. Booth and especially lwo anonymous
reviewers greatly improved earlier drafts of the
manuseript,

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CuHiszan, D. (1978) Lateral displays in the lower
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habitat complexity on postreeruitment processes in a
temperate reef fish population. . Exp. Mar. Biel. Ecol.
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Doverty. PJ. & Fowrer, A, J, (1994) An empirical test
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Epyvane. K. S. (1993) An ecosystem-based approach to
murine fisheries management pp. 21-27 Jn Hancock, D. A.
(Rd.) “Sustainable fisheries through sustaining fish habitat,
Australian Society for Fish Biology Workshop. Victor
Harbor, SA, 12-13 August, Bureau of Resource Sciences
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VAN DFR Etst. R. (1981) “A guide to the common sea fishes
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Hotcuiss. B. & Swamstox, R. (1986) “Sea fishes ol
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Joxes, G. P, (1988) Ecology of rocky reef’ fish of north
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22. 445-462.

Kurrer. R. (1983) An unnotated list of fishes of the
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Fish, (S, Aust.) 7, (-12.

Leum, L, L., & Cyoar. J, H. (980) Density und distribution
patterns of the temperate murine fish Cheilodacrytis
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Biology $7, 327-337.

LincoLn Samim, M. P, Beir, J, D.. Porntarp, DB. A, &
Russent, BoC. (1989) Catch and effort of competition
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temperate reef fish Cheilodeactylus speciahilis
(Pisces : Cheilodactyliae). Mar Beal, Proer, Ser, 35,
113-120.

_—— (1989b) Spatio-temporal panerns in the sbundance
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& CuHoat. J. H. (1987) Estimating total abundance
of 4 large temperate-reef fish using Visual strip-transects,
Marine Binlogy 96, 469-478,

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ScurRorper, A.. Lowry, M, & SuTHERS, [. (1994) Sexual
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THE MOST VIGOROUS SOUTH AUSTRALIAN TIDE

By C. SCHLUTER*, J. A. T. BYE}, & P. HARBISONE

Summary

Schluter, C., Bye, J. A. T., & Harbison, P. (1995) The most vigorous South Australian
tide. Trans. R. Soc. S. Aust. 119(3), 123-132, 30 November, 1995.

Harmonic analysis of tidal records for the region between the city of Port Augusta and
Yorkey Crossing in the upper Spencer Gulf indicates that the most vigorous South
Australian tide probably occurs just north of the Whyalla Railway bridge and has a
maximum range of about 4.1 m, just short of being classified as macrotidal. The
special property of South Australian tides, that the semi-diurnal constituents (M, and
S,) have about equal amplitudes, results in very interesting shallow water tidal
interactions, in particular the generation of a large amplitude quarter-diurnal
constituent (MS,). The intertidal environment of mangrove forests and especially the
samphire flats of the upper Spencer Gulf is shown to be finely tuned to this shallow
water tide.

Key Words: tides, Spencer Gulf.

Transactions of Me Kevul Seciery af) S, Atest (OOS), MICS), 52

THE MOST VIGOROUS SOLTH AUSTRALIAN TIDE

by C. SCHLUTER*® J. A. T, Brey, & PL HARBISONT

Summary

Striloier. Co. Byrd QT. ke Haneison. P1995) The most vigorans South Ausvalian tide, Tras, Ro Soe

4. dist: V3), 123132, 30 November, 1995,

Hurmonic amilysis of Gua) records for the region between the cy of Port AuLUSE amd Yorkey Crossing in
the upper Spencer Gulf indicates thal fhe frost vigorwus South Austrtham nee probably oecurs just north: of the
Whyallt Ruilway bdge and has a maximun mings bf about 1 un asi short of being classified as maerotidal,
The special property of South Australian des. thal the semi-diurm! constituents (M, and Ss) have about equal
aunplitudes, results in very interesting shallow water Hdal interactions. in particulue the generation of a large amplitude
quarter-diurnal constituent (MS,). The interbdal enviroamene of mangrove forests and especially the samphire
this of the upper Spencer Gulf isshown w be finely tuned (o (his shallawy water tide.

Kry Worns: fides, Spencer Gull

Introduction

The tides of South Australia have altracted interest
for aver 100 years (Chapman 1892; Easton 1970),
However there appears 10 be no accaunt of (he region
in whieh the lurgest tide oeeurs. This region. is oF
interest lo dal theory because both Gulf St Vincent
and Spencer Gull have large semi-diurnal (ides at their
heads and alsa because the major lunar (M,) and
solar ($3) constituents are of similar: magnitude. The
diurnal (ide progresses from west to east along the
Southern Shelf usa Kelvin wave which enters the South
Australian seu where its amplitude and phase inerease
regulatly and gradually towards the head of the gults.
The semi-diurnul tide. oo the other hand. displays i
tnuch iiore energetic response,

Tidal characteristics in Gulf St Vincent
and Spencer Gulf

Ap-almost progressive waye enters Investigator Strait
and becomes converted jnto & standing oseilauon
within Gull St Vincent (Bye 1976), Bowers & Lennon
(1990) investigated the tidal character using the classical
model (Bowden 1983) in whieh an incoming waye ts
refleeted at the head of the gulf in the presenve of a
(rictional force linearly proportional to the tidul current
velocity, Particular fitention. was given to the
importance of Backstarrs Passage in this process. The

+ Department of Environmental Bayineerings Cenrre for
Water Research. University of Western Australia, Nedlands
WA. 6009,

t The School of Barth Scyences, The Flinders Universily
of South Australia, GPO Box 2100, S.Aust 5001,

£ pH Govironment, 26 York Street, Adelaide S Aust. SOO,

1 Scuveter, ©, G, (1994) The Generation of Shallow Water
Tides within at Mangrove Enyjronment, MSc Thesis. The
Tlinders University of South Austrabia (unpubl, )

system can be best described as a quarter-wave
resonance af [he open sea tide,

In Spenver Gulf the tidal resonanee is more complex
and lies closer tou three-quarter resoounce in which
a tidal node oceurs between the head and the mouth
of the gulf This behayiour results in a minimum semi-
diurnal tidal amplitude near Wallaroo, beyond which
there is a rapid inerease in amplitude towards the head
of the pulf. Easton (1978) bas given an elegant
mathematical demonstration of this resonance which
also uses a frictional (er linearly proportional to the
Gdal current velocity. Numerical models of the tides
of Spencer Gulf bave been developed by Noye ef al.
(981) and Bills and Noye (986), including fine
resolution models of tidal eddies in upper Spencer Gulf
(Noye 1984; Noye e7 al,, 1994),

In both gulls mangrove forest und samphire flats are
extensive, especially near Port Wakefield, Port
Adelaide! (Schluter 1993), Franklin Harbour and Port
Augusta. The des are-of great ecological sigmficunce
to these areas, The most vigorous tidal system in South
Australia occurs between Port Augustit and Yorkey
Crossing in upper Spencer Gull This distinctive region
has the character of a brine estuary (Bye & Harbison
1990).

The action of tidal cureents is responsible for internal
mixing of the waler column, Stabilising forces, sucli
us Surface heating and horizontal salinity gradients,
tend to result ina stratified water column, the lower
stratum being denser than the upper, The dynamics
Of the mixing process have been extensively studied,
inilially in temperature stratified environments such
as the shallow European seas (Simpson & Bowers
981). and more recently in the salinity stratified
enviroment of the South Australian sea (Samarasinghe
1989). Stratification oceurs when the ratio of the
horizontal density gridient multiplied by the depth and

{i C SCHULUTER, JA
divided by the density and the root mewn square of the
slope Uf the wetter surtice die to the bde exceeds a
critical valué (Bye 1990),

In Upper Spencer Gull the tidal currents ovainian
a vertically well mixed water column whereas in mid
Spenver Gull a Very detailed ond extensive
Observational programme fas shown that transient
stratification occurs (Nunes & Lennon 1987), At the
mouth af Spencer Gulf the hortzoncal density pradierit
Maintains a siranfied exchange for about nine months
ot tie year (Bye & Whitehead 1975; Lennon er ad.
1987) In the summer Months, however, Ite horizontal
density geadien! 1s reduced due io the reversal of sign
in the temperature yradicnt and vertical mixing can
also occur ‘These considerations highlight the
sigmilicance of the tidal regime jy the dispersion ol
dissolved material introduced inte (he wafer column
inn he cowstal provinces of upper Speneer Gulf,
especially m the rine estuary,

The Site

Previously, meastutements af the ude in Spencer Gulf
extended nly as lar north as Port Augusta. Recent tidal
measurements (Bye & Harbison 1987, 1991, 1994) north
of Port Augusta (Fig, 1), indicate thata very vigorous
tidal regime exists. The site of the investigations was
yoold wooden bridge on whieh an abandoned mineral
railway Co a salt works crossed Over Spencer Gull) On
(he caster shore of the gulf the salt bridge was
Originally commected with an embankment which cuts
(hrough the mangrove lorest and into the samphire Hats.
The maximum span height of 4.4 m is just greater than
high water springs, and atlow water springs, the water
by confimed lou lew central spans where the max anit
depth js about 30 em (Pig. 2a), The piers and spans
we ideally sulted for instrument deployments ‘Tide
fanges were secured to the piets and Current meters
were suspended from the spans or mounted hy poles
driver into the ground. On the western side, beyond
4 narrower fringe of mangroves, the bridpe toads
directly to the salt works, The tidal observations
vndertaken it (his site extended trom 28 February lo
2% March, 1986 (Bye & Harhisen 1987)-

Tide gauges und current meters were also deployed
for shorter petiods between the Central Australian
Railway bridge and Yorkey Crossing. This station lies
uppreximately 4 kin further north of the salt bridge
and is bordered on both sides by samphire fats (Fig
2b) Which are covered at high waler springs and also
Wuring the rare floodings which originate on the Pirie-
Torrens plains north of Yorkey Crossing (Bye &
Harbison 1994).

Since the above observations were of limited rime
spar, second mire extensive period was initiated! in
1993, The choice and location of tal equipment was

HYF & P HARBISON

based upon the pilot investigations. Aa Lnter-Ocean Sd
current meter wits located within the town of Port
Augusta at un abandoned road bridge using a cradle
mooring device, The current meter is based upon an
clectvomagnche ax measurement and thus has tea
moving parts as do the conventional current roeters

This makes the current meter immune to the effect of
alval growth, The current meter obtained data between
27 July 1993 and 27 August 1993, A tide gauge located
at this sie proved faulty, The tidal constants in Table
T were obtained from the Port Augusta power statien
tide gauge tot the same period as the salt bridge
Jdeploviment.

A bottom mounted pressure fide gauge was deployed
atthe Whyalla Railway bridge north of Port August
between 28 July and 30 October 1993_ Another bottom
mhounted tide gauge was deployed at the salt bridge,
but this gauge was destroyed by vandals. A botton
mounted {ide palige Was deployed at the Central
Australian Railway bridge between 18 September anc
S| October 1993. This gauge cleurty indicates the
unusual tide of the tegion.

Tidal Analysis

To distinguish benween a progressive und standing
wave Situation the phase differences between the tidal
corrents and clevations must be kfown. One of the
most inportant aspects of tidal systems is the definition
ob standing anid progressive waves. With standing
waves. the phase difference between the tidal elevations
and ciltrents is 90° while for 4 progressive wave the
phase difference js O°. A standing wave may be
considered as being the sum of two progressive waves,
one directed landward and one of equal amplitude
directed seaward. The Jandwatd energy Hux associated
with the incident wave is, in this case, balanced by the
seaward energy flux of the reflected wave and thus there
is no pet energy flux, In dissipative situations the
reflected wave will be frictionally attenuated and must
be smaller in amplilude than the inbound wave and
(hus a perfect standing wave is impossible, As the tide
propagates from the open ocean. various nonlinear
distortions occur in the tidal signal, These distortions
ate primarily influenced by nonlinear mechanisms
including frictional interactions and interactions with
surrounding bathymetric features, as well as
atmospheric effects and continuous freshwater
discharges. The interactions of the primary
astronomical tide discussed above may be represented
by the growth of the shallow water tides whieh indicate
the degree of nonlinear distartion af the primary signal.

In the analysis of tidal signals, it is common practice
to decompose the tidal signal into a harmonic series
of amplitudes and phases, The total tidal elevation or
current may thus be represented by the sum of the

MOST VIGOROUS SOUTH AUSTRALIAN TIDE 125

BA Yorkey

Central Australian Crossing

Railway bridge .*.*.)
1 bottom mounted—
tide gauge

salt bridge
1 bottom mounted
tide gauge

samphire salt bush

Abandoned /‘.: } *% | mangroves

Salt Worksalf 4". road

railway

Whyalla
Railway bridge
1 bottom mounted
tide gauge

1 bottom mounted
tide gauge

1 InterOcean S4

current meter

Fig. 1. Locality map of upper Spencer Gulf.

126 C SCHLUTER, | A. T BYE & P HARRISON

Fig. 2. (a). View of the salt bridge looking westward Note un investigator (Pat Harbison) on the fringe of the mangrove

forest, and the abandoned salt works in the background, (b), View lnoking southward trom just before Yorkey Crossing
Note the samphire flats on (he westem side of the chinnel, and the Cental Australian Railway bridge in the backeround
Both views were taken a short time alter low wurer

MOST VIGOROUS SOL'TH AUSTRALIAN TIDE 137

Tante ( Vidal constituents for the upper Spencer Gulf.

a = amplitude; g = phase in degrees. The tidal elevations are in vim and the currents are m ems. Currents are related
1 the magnetic bearing. The record lengths (sed with the analysis are given in the text.

Elevations Currents
Constituent Port Augusta Whyalla salt bridge Central Aust Port Augusta Port Augusta
(986) Railway (186) Railway (Bast) (North)
bridge bridge
a a a £ 4 E u z a g a g

O, 248 39 165 42 263 SI WO 75 UW.75 322 2.98 337
Ky 483 73 462 74 45) 80 308 90 U.68 | 2.14 4
M; 617 206 oll = 210 559 223 245 227 2.14 140 928 139

in td 257 104 267 AB 275 301 288 2.62 198 10.8 202
3M3S, 4a 24005 5 278 35 «WW G18 #3 O58 109
Meus x 224 1 400 92 30 33 1R9 0.28 218 (15 274
25M, 49 «93 107 78 Ih 12 330 4 52 42 245 ST
35M) 42 165 13 Is M 75 26 ORS O19 347 42 350
MO, v] 91 7 63 ld 175 28 284 003 79 138 172
SO, 3h at 200 Ol 242 Su 356 O45 200 345 219
SK, 3 94 13) 208 28 = 459 76 35% O1K 337 li 276
Mg 2 Sis 3 317 24 ST 50 132 25 59 118 U8
MSy 34 9 12 295 7 U4 1360 38 029 [42 I41 32
Sa 1 302 ta 523 5A 73 78 IRS O45 276 0.54 307
4MS,, 3 229 4 345 7 48 | 337 oos 7I 0,47 39
My Uf) 9g Q 497 5 (29 1 64 O13 355 0.26 349
2MS,, K 40 14 2 27 l49 2 44 O17 WY O46 45
25M, 18 139 Il 93 ad 2 \4 76 26 [68 163 169
Se 2 2K6 9 130 2 245 4 205 013 36 0,39 27
4SM, 4 242 6 35 4+ 408 4+ OW 202 O56 221

predicted harmonic series and (he residual signal, i.c. Results

Galt) =] Cyveut) + Cyst)

where ¢€om(t) is the observed tidal elevation or
current, ¢re(t) is the predicted tidal elevation or
current, (ri(t) is the difference between the ohserved
and predicted tidal elevation und current,

The predicted harmonic amplitude is given by
i]
Cult) = Lyacos(ort— 2.)
=|
where a) 1s the amplitude of the ith constituent, o is
the frequency of the ith constituent, ¢ is the local time
of |he dita and g; is the corresponding, phase lag,
The residual signal includes all tidal frequencies
which ure nol harmonically analysed as well as
atmospheric storm surges. These storm surges usuully
persist for 2 Wy 4 days depending, on almospheric
conditions,
The harmonic analysis of all tidal constituents (Table
1) utilising programs developed by the National Tidal
Facility shows the importani aspects of the upper
Speneer Gulf vide

2 Only part of py is 4 shallow water conshtuent, joy also is
aw minor primary tide,

The four major primary constituents (M,, S., Ky,
und ©)) are the maim energy source for the region,
und the interactions of the dominant semi-diurnal
doublet (M, and $5) generate suites of quarter-diurnal
(My, MS,, S,), frictional semi-diurnal (3M,85,452,
25M, and 38,M),) and frictional sixth-diurnal (4MS,.
M,. 2MS,. 2SM,, 5,. and 48M,) shallow water
constituents (Pugh 1987). Terdiurnal (MO,, SO,, and
SK,) shallow water Constituents are also generated
through inlerachions between the four mayor primary
constituents. The MK, constituent, however, is not
resolvable from the SO, constituent owing to the short
length of our records, The tidal energy spectrum al
the Central Australian Railway bridge (Fig, 3) clearly
shows energy peaks lor cach of these bands, and also
thal this energy is resolved by the harmomic analysis,
Two prominent higher frequency bands not presented
in Table |, are also shown

Table | indicates that the primary constituents have
a maximum amplitude between the Whyalla Railway
bridge and the salt bridge and also that their phases
increase between Port Augusta and the Central
Australian Railway bridge. At Port Augusta the tidal
currents lead the tidal elevation by about 60°, indicating
a northwards propagation of energy.

Each group of the shallow water constituents appears
to behave differently, although there is large variability
between the constituents within the groups. The

J2% C_ SCHLUTER, J. A. T.

clearest signal is shown by the quarter-diurnal con-
stituents, the amplitudes of which ure far greater at the
Central Australian Railway bridge. The amplitudes are
approximately in the rauo of 1:2.) for niost stations
in agreement with theoretical prediction (Gallagher &
Munk 1971) and differ by about 180° between Port
Augusta and the salt bridge (Table 1), This is consistent
with a node occurring near the Whyalla Railway bridge
where the quarter-diurnal amplitudes are a maximum.
The phase of the terdiurnal constituents also differs
by about 180" between Port Augusta and the Central
Australian Railway bridge where (he amplitudes are
about half of the quarter-diurnal amplitudes,

The frictional sixth-diurnal constituents, on the other
hand, tend to have inaximum amplitudes at the salt
bndye und very variable phases. Finally, the fricuional
semi-diurnal constituents show maximum amplitudes
al Port Augusta, with the suggestion of secondary
maxima at the salt bridge.

Discussion

The propagation of the primary tidal constituents
(and also the frictional semi-diurnal shallow water
constituents) into upper Spencer Gulf gives rise to the
most vigorous Gide in South Australia which oceurs
between the Whyalla Railway bridge and the sall bridge
where there is a generation of frictional shallow water

0.01 »

0.001

Spectral Density (m*/cpd)

0.0001

Je-05
0 1 2 3 4 5

. BYE & P, HARBISON

tidal encrgy. The position of this maximum tide
coincides approximately with the node of the quarter
diurnal tide. The maximum tidal range is detined us
the suininiawion of the nieun spring semi-diurnal range
(MSR) and the mean spring diurnal range (MDR)
where MSR = 2(M, + 8,) and MDR = 2 (K, 4
Q,) (Easton 1978). Following the above definition the
maximum recorded tidal range for South Australian
Waters occurs at the Whyalla Ratlway bridge and
corresponds to an clevation of 4.1m, compared to the
range at Port Augusta of 3.9 mn. This tidal range
identifies the tide as being al the very upper end of
the mesotidal range (2.1 - 4.2 m)

Friedrichs and Aubrey (1988) have classified
estuaries in which the lunar semii-diurnal tide (M,) is
dominant over the solar semi-diurnal ide ($2) int)
ebb dominant and food dominant, In an ebb dorninant
estuary, much wreater idal currents oceur during the
ebb following the exposure of mudflats and the release
of intertidal storage. To overcame these effects the
duration of the ebb lide is far less than the food lide.
The consequence of ebb dominance is to provide ii
mechanism for the long-term outwelliny of sediments
and pollution, In the reverse situation of Mood
dominance, the estuary usually does not contain
intertidal mudflats and thus the duration of the flood
tide is less than the ebb and as a result the currents
are greater on the flood ride. Flood dominant estuaries

Residual Spectrum

Observed Spectrum

6 7 8 9 10 11 12

Frequency (cycles/day)

Fig, 3. Power spectrum of tidal energy at the Central Australian Railway bridge.

MOST VIEOROLIS SOUTH AUSTRALIAN TIDE eau

usually consist al more unstable zeometnes and wm (he
lig teen become fled with fine sediments which
arise from fhe net transport mtr dhe estuary. Chis
Classification is based on the phase difference hetween
the prinvary lunar tides (M5) and the major shallow
water tide (My). In the ebb donwnant situation, (he
mimiium M, current approximately cemeides wilh
the maxiniunt of the ML, current, and ut the flood
dowunant situaon (he maximum of (he My current
approximulely coincides wilt the Maxirouny of the My
current,

We find thal this clissifeabon iy ot appropriate fy
the brine estuary of upper Spenver Gull, which behaves
either as hgh water oe low water dominant. High walter
dominant conditions occur when the difflerenees
between the High water level and mesn sea level are
much grewler thar) those between the low water level
and mean sea level: (he opposite oecurs for low water
dominant conditions. fi high water dommant situations
the maximum of the dominant shallow water
conabluent approxiinately coineides. with the maxim
of the primary Uidal amplitude and vice versa, For
crample, arthe Central Australian Railway bridpe. the
phase difference benween the MS, constituem and its
corresponding astronumicul generating tide 1s
(M,+S.)-MS8,=17~

The gh water danniuat enyironuient cansists of
a well detined channel which 1s very shallow at low
water, but Which can accommodate the propagation at
the meoming-and oulyomy tides except yery near high
water when overbank Now on the samptire hanks
occurs. However, with the law waiter dorninunt
environment, the channel is deep enough to have a
negligible effect on low water levels, but.as high tide
approaches, Turge yolumes ol] water spill mto the
adjacent mangrove areas filling up the interfidal
depressions und truncating the high water level. The
node between these two environments where the most
Vigorous tde ovcurs (iarks the position Where these
tw opposite effects are in balance. These properties
appear to be due to the almost equal amplitudes of M,
and S$, (see below!

High water Joniinant conditions are well developed
atthe Central Australian Railway bridge such that the
low water levels appeal to be “euler (Fig. db). There
ure four interesting teatures of this pecerd. First, the
low waler levels al /he peap lide are lower than ar the
spring tide. It is believed that this is the result.at the
formation of intertidal pools of gulf water trapped
berween the Central Australian Railway bridge and
Yorkey Crossing, Water is held in this region on the
ebbing tide which ws slowly diridined until the ude tums
bul during the neap pyele, less walter is able (o be stored
io the upper reaches and subsequently the low weter
level is less than the spring tidal level.

Second, the ditrnul inequaliy of the ide produced
hy the bewing of the diurnal dnd semi-ditrnal eon

stituents 1S Signiticanily madulated ac the Central
Australian Railway bridge relutive wo the Whyalla
Railway bridge due to the generation of the ferdiurnal
Udes.

Third, the residual records show thal stornr sunes
are usually greatly attenuated bewween Whyalla and
Central Austratian Railway brilges,

Fourth, the drainage (rom the sumphire flats retards
the low tide much tore than the high tee ‘The
approximate lags between Yorkey Crossing and Pant
Augusta, determined direedy trom the short tidal
records of the pilot study, were high water 35 min,
low water 180 min, Similar results can be obtained
from. Fig. 4a and b. The difference between the lags
is promarily due ter the major quarter diurnal shallow
water (We (MSy), 45 can be seen from Fig, Sa in
which the three tidal constiluenls M5. S, and MS, are
represented as well us the combinauon of the uirec.
An interesting feature of Fig, 46 1s the form of the tidal
Tiige curve during the ebb, which resembles an
exponential drainage curve, and is clearly reproduced
in Bip. Sa. ‘Thus the drainage from the samphire flats
is explained harmonically by the existence al MSy,.
This appears to be & unique property of the system in
which M, and S, have approximately the same
amplitude, The corresponding current record (Fig, 5th),
which is consteucted hy differentiaing the tdal
clevations with respect ta time. and scaling lo pave
niaximum tidal velocities similar to the observatians
(~U.A5 nis!) shows characteristic current spikes at
(he beginning of both the ebb and flood tides which
are of similar amplitude and also a slaw ehh (3
ems ') during low tide. This structure was observed
in the Short period current meter deployments just
south of Yorkey Crossing and at the salt bridge (Bye
& Harbison 1987, 1994).

Flattening of high water ain be seen inthe Whyalla
Railway bridge reeurd (Pig. 4a) but the impurtanee of
MS, (Table }) 0s much smaller here thar in the high
lide demninant conditions at the Central Australian
Railway bridge

Conclusigns

Following: at extensive field prograinine of (dal
elevauions and currents, the laraest ude in South
Australia § believed to exist in upper Spencer Gulf.
north of Port Augusts. This tide is the peak of the three-
quarter resonance in Spencer Gull which vives rise
to arapid increase in amplitude northwards trom near
Wallaroo, The jocalion of the Whyalla Railway bridge
was observed 19 hitve the largest amplitudes of (he
major astronomical tidal constituents (namely the M5
S.. K, and O, tides) giving acidal range whieh Ses
within 4 to 4.5 m where typically the tidal range in
both gulls 1 claser to 2.5 to 4a, Just heyond the bridee

130 C, SCHLUTER. J, A. T. BYE & P. HARBISON

10 T —T T —_ rt °° °»°»~=2~«' T
€ (a)
ey 8
Qo
=
2 6
2
o)
4 1 4 4 i

Predicted (m)
lop)

Residual (m)

= ! !
4/10/1993 6/10/1993 11/10/1993 16/10/1993 21/10/1993 26/10/1993

date

3 T T a > o———— eo
< (b)
He
Oo
2
faa
ne}
(oe)

3 1 T
£
o2
2
3
eit Ai
o

0 = a ee La =

é
w
=!
2
WwW)
@
c
=I 4 ——— ai ———|—
1/10/1993 6/10/1993 11/10/1993 16/10/1993 21/10/1993 26/10/1993
date

Fig. 4, Tidal observations tor October 1993, (a). The Whyalla Railway bridge. (b) The Central Australian Railway bridge,
The residual (Res) = the observed (Obs) - the predicted (Pred) tidal elevation.

MOST VIGOROUS SOUTH AUSTRALIAN TIDE 131

Amplitude (m)

- M,
06 + 3
Ms,

Time (hours)

Speed (m/s)

Time (hours)

Fig. 5. (a) Tidal elevations and (b) Tidal currents at the Central Australian Railway bridge, constructed using the tidal constituents
M,, S5, and MS,.

132 C. SCHLUTER, EA

a shallow water dal node occurs and we idennty Us
location as the probable position of the largest tidal
vurige. As in all tidal stadies, however, the lonver the
length of tidal recards, the betler is the harmonic
analysis, In this study we have relied on one-two month
deployments ut several locations. Longer reeords would
be necessary lo improve the decuracy of the Udal
constants.

The nonlinewr interaction between the major
astronomical constituents and the surrounding
bathymetric features leads (to the veneration of
significant shallow water tides, especially the quarter
diurnal MS, constituent which dominates because of
the similar amplitudes of the M, and S, tides,

These observations prompt the speculation that the
samphire Mats and mangrove forest environment have
evolved asa positive feedback t0 the shallow water tidal
interactions. In other words, In the absence of the
unusual South Australian tidal regime in which the
major semi-diumnal tidal constituents (M. and §.)

BYE & BP HARBISON

have a similar amplitude, the intertidal environment
would be quite different.

tis also likely that the changes: in the intertidal
envirooment of upper Spencer Gulf and its northward
extension into the Pirie-Toprens plains hat have
oecurred due tu sea level changes have been decisively
influenced by shallow water ddal unteractinns.

Acknowledgments

The wurhors would like to Unk ihe seat of the
National ‘Vidal Facility for the preparation of all the
udul equipment used in this study und also for (he
invaluable assistince in the lidal analyses. The
invesigation was supported by a Marine Sciences and
Technologies (MST) Grant 85/I0IS and a Flinders
University Research Budget (URB) Grant, Helptul
comments by the referees are alsy aeknowledged,

References

Biis, Po & Nove, Bod. (986) Tides of Spencer Gult,
South Australia pp. 519-530 Jn Noye, BJ & May. Po
(ids) “Computational Techniques & Applications
CTAC-85" (Elsevier, Anisterdar).

Bowers. DOG. & Lesson, GW. (1990) Tidal progression
ig Tear-resonunt system a cuse study [rom Sourh
Australia, Esteearine Coastal and Shelf Serenee 30, 17-34

Bri. J. A, T) 1976) Physicul oceanography of Gull St
Vineent and Investigator Stra, Trans, KR. See. 5. Aust. 100,
143-160,

(1990) Richardson Number profiles in laboratory

experiments applicd to shallow seas. Geaplys, Alstropltys,

Fluid Dyn. 52, 145-166,

—. & Haruison, P (1987) “Hydrological Observations

in Spencer Gulf and the Pirie Torrens Plains, South

Australia during (986: Cruise report 14° (The Flinders

Institute for Atmosphere and Marine Seienees, Adelaide),

& (891) Transfer of inland salts to the marine
enviromental the head_of Spencer Gull, South Austraha
Palacoseour, Palacnclimatol, Palienecoul 84, AST-368,

il BH (1994) “Hydrological Observahons in
Spencer Gulf and the Pirie- Torrens Plains, South Australia
during 1987 and 198%: Cruise report 15" (The Flinders
Institute for Atmospheric and Marine Sciences, Adelaide),

_ & Werrevieao. JR. Ine. (1975) A theoretical model
ofthe flow in the mouth of Spencer Gull, Sourh Australia.
Esinarine and Coastal Marine Science 3. 477-481,

CHAPMAN, BR. Wo (1892) The tides of the coast of South
Australia, duit Assen. ddvanee, Sci, Rep. Committee 2

baston, AK, (1970) The lides of the continent oF Australia
Horace Lamb Centre for Oceanographival Research Res;
Pap. 47,

——— (1978) A reappraisal of the lides i Spencer Gull
Soutle Australian, Adsl J Mar, Freshy, Res, 3, 467-477.

PRibbRICHS. CT & AnaREY, DG, (1988) Nonlinenr tical
distortion in shatlow well-mixed estuaries a synthesis
Faraanne Coastal and Shel} Seience 27 521-945.

GALLAGHER. BOR Munk. W. CLO7L) Tides in Shallow Water
Spectroscopy, JeHus 23, 446-463.

Lennon. G. W., Bowers, DOG. NUKES, RL A., Sear, B.
BD, AU, M., Boyre, 3, Wes, C, Weeeonip, Mo,
Joransson, G,, Nine, S. Perrosevies, 2. StepHpson.
P, Susmin, Av Ag dk Winkie. SB A, (887) Gravity
currents and the polease of Sall from an inverse estuary,
Nature 327, 695-697,

Nove, Bo J. (984) Physical processes and pollution in (he
waters of Spencer Gull, Mur. Geol. Gt, 197-220,

—. May, R. 1, & Trunser M- DB. (O81) Three-
dimensional numerival of tides mn Spencer Gull) Qeeu
Management 6, TAR.

—, Bins Po & Lewis. G (1894) Predionan afar
Slick mayerment in Northert Spencer Goll pp, 320-328 tn
Gardner, Ho, Singleton, DL, Stewart, OD. (Bas)
“Computational Techniques & Applicatianss CTAC-93%
(World Scientific, Sinuapere)-

Nuwpes. Ro A, & Lenwon, G. W. (1987) Episodic strut
cation and gravity currents iA a marine environment et
modulated rurbulenee. .. Geapliys, Res. 93S, $465-S480,

Puc, DT. 41987) “Tides, surges, and mean sew level”
(Wiley, New York).

SAMAR ASINGHE. LR. de sibva (1989) Transient salt wedges
ia tidal poll’ A criterion for their formation, Esmarine
Comite! and Shelf Science 28, (W148.

Scubuper. C, CG, (1993) Tidal Modelling withia aw Mangrove
Environment, preprints af the th Australasian Conference
on Coastal and Ovoun Enymecring” (The Institution of
Engineers, Australi).

Simpson. J A & Beavers. Do 981) Morlels of stiarfcation
and fron) moverient in shell seas Decp Seu Rey. IRA,
727-738,

STUDIES ON EUTOBRILUS HEPTAPAPILLATUS (NEMATODA:
TOBRILIDAE) THE PREDOMINANT NEMATODE INHABITING
THE BOTTOMS OF LAKE ALBERT AND ALEXANDRINA,
SOUTH AUSTRALIA

By ALAN F. BIRD*

Summary

Bird, A. F. (1995) Studies on Eutobrilus heptapapillatus (Nematoda: Tobrilidae) the
predominant nematode inhabiting the bottoms of lakes Albert and Alexandrina, South
Australia. Trans. R. Soc. S. Aust. 119(3), 133-141, 30 November, 1995.

Eutobrilus heptapapillatus, a cosmopolitan fresh water nematode has been isolated
from the bottoms of Lakes Albert and Alexandrina where it comprises up to 87% and
85% of the nematode population respectively. The environment at the bottoms of the
lakes in which these nematodes live is described and measurements of males and
females from each of these environments are compared with those of a South African
population. There are significant differences in tail length between the Australian and
South African populations. Egg laying in the Australian population has been observed
and is described. The presence of crystalloid structures in these nematodes has been
noted and the possibility of their occurrence being associated with increased salinity
is discussed.

Key Words: Eutobrilus heptapapillatus, nematodes, Lake Albert, Lake Alexandrina,
sediment, eggs, morphology, measurements, crystalloids.

Hansaetiinis of te Raval Saciety af 8. Aust. (1995). W931. 19-141

STUDIES ON EUTOBRILUS HEPTAPAPILLATUS (NEMATODA:
TOBRILIDAE) THE PREDOMINANT NEMATODE (NHABITING THE
BOTTOMS OF LAKES ALBERT AND ALEXANDRINA, SOUTH AUSTRALIA

by ALAN F. Birb*

Summary

Gino. A, F (1995) Studies on Fawhrilas heptepapiliaws (Nematoda , Tobrilidac) the predominant mematade
inhabiting the bottoms of lakes Albert und Alexandrina, South Australia Jruny, Ro Soc S Aiest, 13), 133-141,

40) NoveTiber 1995.

Eyinbriluy heptupepillais 3 cosmopolitan fresh water nematode has been iselarert from the bottoms ol Lakes
Albert and Alesundrina where it comprises up lo 87% and 85% of the nematode population respectively. The
cuviropment al the bollors of the lakes in Which these nematodes live is deseribed and measurements of inales
and females from exe of these environments are compared with thase of a South Aftican population, There
are significant differences in tail length berween the Australian and South African populations, Bee laying yn
the Australian pepulalion has been observed and 7s deseribed, The presence of crystalloid structures in these
nematides has been noted and (he possibility of their occurrence being associated with mereased salinity 1 discussed.

Kiy Wanos: Buobrifus beplapapitiaius, pemulodes, lake Albert, Lake Alexandrina, sediment

Morphology, measurements, crystal los

Introduction

The nematode Autobrilay heptapepillains (Joubert
& Heyns, 1979) Tsalolikhin, O81 has a world-wide
distribution in a range of freshwater habinuts. [n South
Australia this nematode occurs at various sites on the
shores of Lake Alexandrina and Hindmarsh [sland at
the mouth of the Murray River (Nicholas eral. 1992)
ind was the mast comin species extracted from a
suiple dredged from a depth of 3 m at the southern
end of Lake Alexandrina. To date no studies haye been
published on measurements of this nematode nor of
its presence or absence at the bottom of the adjacent
Lake Albert.

Th this paper | compare measurements of males and
females oF E. heptapapillates: trom Lukes Albert and
Alexandrina with those from South African populations
(Swart & Heyns 1988). L also describe thetr habitats
dnd their proportions to ather nematode species Jound
in these habitats, as well as the percentage of F.
heplapapillarus containing crystalloid inclusions.

Materials and Methods

Collection of material

Samples were collected using a benthic grab from
the bottoms of Lakes Albert and Alexandrina al the
following localities. For Lake Alexandrina the
cOlleeting site was at thé navigation marker No. 84
(Fig, 1 site [1)), The Lake Albert collecting site was
2-3 kin off shore from the town of Meningte with
compass bearings 135" on the town’s water tnwer,
285° an trees. on the Coorong side of the lake, 205" on a

*) Playford Road, Mitcham, S, Aust, 5062,

apps,

headland on the port side and KS” on a barren hill top
on the starboard side (Fig. 1 site [2]), In each case
the contents of the benthic grab were placed in a plastic
hag and stored in u cooled msulated container,

The dry weight of the sediment was determined by
allowing it 10 grayitate from the water included in the
benthic sample in a graduated cylinder. The supernatant
was removed by suction and the sediment was then
spooned into a weighed beaker whieh was placed i
an incubator af 40°C. Dehydration was maintained
until a constant weight was reached.

Membrane (0,2 am) liltered walter sarnples from the
Jakes were taken simultaneously with the sediment
samples taken with the benthic grab. Salinity was
calculated from. electrical conductivity (Nicholas et al,
1992) and a range of elements was analysed using the
technique of Zarcinas and Cartwright (1983), Particle
size of these samples was measured using various
techniques as deseribed by Beech (Nicholas et al
1992),

A large plastic container was filled with lake water
from the sampling site and this: water was used lo dilute
the samples during the sieving procedures used to
separate the nematodes, This consisted of passing the
samples through 2 mm, 850 pm. 710 zm, 250 pm, 120
wm and 90 pm sieves. Ln samples containing much
sand, further sieving through 73 pm, 53 pm and 38
ji Sieves Was undertaken. However, in the case of
Lake Albert samples, the sediment which passed
through the 90 pm sieve would have blocked the
remaining three sieves. Accordingly, the material
obtained on the 120 zm and 90 win sieves was diluted
to facilitate microscopic observation and aliquots were
examined under the dissecting microscope. The
nematodes were picked out alive on mounted eyelashes,
their Movement indiculing their presence in the sample.

134

They were placed in a test tube in a small volume of
filtered lake water and an equal volume of boiling
double strength FA 4: | solution (20 ml 40%
formaldehyde and 2 ml glacial acetic acid in 78 ml of
distilled water) (Hooper 1986) was added to the shaken
suspension of nematodes. These specimens were
processed to pure glycerol using Seinhorst’s (1959)
method and mounted in anhydrous glycerol on slides
sealed to a coverslip by molten paraffin as described
by De Maeseneer and D'Herde (1963). Nematodes
fixed and processed into glycerol in this manner were
photographed with Ilford Pan F film. Living
nematodes, for example females laying eggs, were

HINDMARSH
ISLAND

A, F. BIRD

photographed using Uford Delta 400 film. These
nematodes were observed and photographed using i
Vanox AHBT research microscope equipped with
bnght field and interference contrast (Nomarski)
optics.

Results

The water enviranment

Lake Albert is a relatively large body of water about
16 km x 10 km connected to Lake Alexandrina, which
is approximately 30 km x 15 km, by 4 narrow channel
of water (Fig. 1).

LAKE ALEXANDRINA

~~ \ site (1)

GOOLWA
e <<

MURRAY MOUTH

SOUTHERN
OCEAN

LAKE
ALBERT

site (2)

MENINGIE

Coorong

Fig. 1 Map showing the location of collecting sites (1) and (2) in Lakes Alexandrina and Albert respectively.

Taste |. Analyses of major seluble ions in water fram the shores of Lakes Alexandrina and Albert sampled en the sume
day at a six monthly interval and from water sampled from the middle of Lake Albert ata later date.

- mel

Date Locality Na Cl Ca Mg K s B.C rss*
(Site) ds m' oe

29 April 1993 Alexandrina 48 82 15 if 5 6 0.42 0.03
(shore) Albert 214 348 8 33 12 22 1.6 O10
22 Oct 1993 Alexandrina $4 105 ¢ 8 3 7 0.4 0,03
(shore) Albert 200 375 34 31 10 22 15 0.10
20 May 1994 Albert 188 205 34 28 10 16 1.23 0.06

(mid-lake)

“EC. = electrical conductivity (deci-siemens mm)
" TSS = total soluble salts (estimated percentage)

A NEMATODE FROM LAKES ALBERL AND ALEXANDRINA li

The results of (he analyses of water collected from
Lakes Albert und Alexandrina are given in Table 1,
From these results it can be seen that there is mostly
a three-to four-fold difference in the tou soluble salts
in water samples collected from the shores of the lwo
lakes on the same day, These differences in the major
ions persisted in water samples taken six months later
(Table 1),

The sediment environment

The surface 15 cm of the soil ut the boule of Lake
Albert consists of a slimy sediment, largely Composed
of clay which made up 48-61% of samples of this
surface sediment or slime laken from various pants ob
the lake as the top component af core samples (Taylor
& Poole 1931),

{t Was estimated that only |/6th of the sediment from
Lake Albert consisted of solid material. This material
comprised 47% clay, 25% silt, 3% fine sand and less
than 1% coarse sand.

The nematade

The most common nematode in the Lake Albert
sediment, Extebrilus heptapupillatus, comprised up to
87% of the nematode population; the remainder mostly
consisied of monhysterids. Similarly the benthic sample
from Lake Alexandrina comprised up to 85% &,
heptapapillatus.

The ratios of larvae, males und females were similar
in two different collections from Lake Albert, In one
instance an aliquot containing 137 nematodes had 43%
larvae, 16% males and 41% temales, In the other
harvest the ralios were 39% larvae, 23% males and
38% females, In an aliquot cuntaming &,.
Neprapapillaris trom the Lake Alexandrina benthic
sumple, (he ratios were 51% larvae, 37% imiles and
12% lemales. An obvious difference beiween these |wo
pupulations was the presence of crystalloid inclusions
(Fig. 2) in 32% of the nematodes from Lake Albert
whereas none was observed from the Lake Alexandrina
sample.

Comparison of populavons of E heptapupillatus,

Specimens from both lakes were meusured and
compared with each other and with those from South
Africa (Swart & Heyns 1988), [rt can be seen (Table
2) thal the males of these three populauions are similar
in many respects. For example. they are of similir
length, have the same body width at the anus and have
similarly-sized copulatory spicules. Diflerences in
maximum body width and pharynx length could nol
be analysed statistically due to the absenee of certain
measurements of the South African population, There
is, However, 4 significant difference in tail length
(P.<0,001) between the Australian populations (Lake
Albert with a mean of 179 yim and Lake Alexandrina
with a mean of 173 ym) and the South African
population (mean of 244 ym). This significant

difference tetail lenge ys mot se pronounced (PF <O01)
in the females of these populations (Table 4), The males
ot EB. heptapapillatus (Pigs 3. 4) have w dioretne
reproductive system consisting Of a pall ol testes, & vas
deferens and ejaculatory duet connecting with the
copulatory spicules. The thost obvious components of
the males acvessory structures are the seven
supplementary organs (Pig, 4) from which tts specific
name is derived,

The distances between these supplementary organs
in the South African population have been measured
(Swart & Heyns |988) and so coun be counpared with
the Australia populitions, Measurements ob the
Australian populations are eapressed as percentapes
of the sum of these distanees rather than as direct
measurements. This is because direct ni¢usurements
using colled and uneoiled nematodes revealed a
significant difference in (he mean value of rhe distance
between supplements in the coiled (27.9 xm) and the
unvoiled (414 pm) (P<0.00)), However. when these
distances Were expressed 4s percentayes of the sum ab
the distineces between supplements. there was nu

Fig, 2. Photogriph of part of an adult female Kucabrilus
heprapapillans that had been maintained in shallow distilled
water ina Petre dish for over a month prior to being
pllowaraphed. Bright field optics showing the presenee of
nufyerous erystalloit bodies (silall arrows). Seale bar =
20 pm

144

AF BIR

TABLED. Comparison ef measurements of eifferent papulatiany af males of Eutobrilis heptapapillatis.

Taitsikama Foresr
Cape Prowince, South Atria
tSwart & Hevns |RKy

All measurements jin

Lake Alben
South Austatha
(present study

Lake Alexandrina
South Australia
(present ostuity }

n= n=o hed
Parts measured Range Meun Range Mew = SD Range Mein = SD
Body length (Ly 1550-2120 an TR73- 2000 U1 +57 1H). LOOG) 1896, +h
Max, body width 53° 77 7} +5 M77 646 +4.)
Pharynx length 460" 45-397 aul 4 THA 29) +18
Tail Jengih 211-400 244 IHR-11) Vu 419 4-192 172 +20
Body wideh ar anus ay 30-41 38 +27 42-40 3K 6
Spiculo 4-57 541 Sd-35 34 +2.2 52.56 54 +17
Gubermaculuin 38-39 370 13% atl +57 3-36 ER) +27
De Man's medices a 32.1-4).3 36,2 15,630 27 +18 2K 29 +13
De Man's indices fh S153 $2 5.7-6.6 62 HS 6.170 0.6 H)4
De Man's indices c f.2-8.8 7 Wh4-ll WLR HOS 99-124 Wal tit
De Man's indices c S.8-8.0 65 445.4 47 H)4 444.8 6 +2
* valoulated From data of Swart & Heyns 1/988)
Taset 3. Comprison of measurement of citferent papadations of feiales af Katobrilus heptapapillatis
All Measurements jum “Thitsakuima Forest Lake Albers Lake Alexandrina
Cape Province, Sout Altea South Australie South Australia
{ Swart & Heyns 198) (presen! study) (present study)
w=! neS nes
Haris meaxured Range Mew Runge Mean = 5D Range Mean = 3
Wody fengeth (1) 172-2280) 2040 2)82-2054 2200 £3) 1940- 22K) 204 128
Max. brody silts uT! RI-4S tata} th H5-N2 74 +6.2
Mharyes length AXS* 3M IKI ASK +18 245.445 320 £22
Tail length 224-337 2 21R-2o8 242 #20 224-246 261 +27
Bedly width alanuy 344" Has 2 22 36-40) aK pa
De Man's indices ¥ Wi). 42 4)-A47 45 +| 33-44 4) 48 .()
De Man's indices a ThH-4| 4 23.27 Ka] #2 ii-34 24 132
De Man's indices b ARSY 53 5O-n6 f2 +,3 6.1-6.9 fn Sale)
De Mun'y. indices e 61 -O 7,1 R212 Ya LK 74-87 Kd HOS
_De Man's indices G80 a7 S365 $.8 45 62-74 68 +04

‘calculated from data of Swart & Heyns (1988)

significant difference (P >0,05) between [he means ot
coiled (14.3%) and uncoiled (also 14.3%) nematodes,
Thus expressing distances between supplements as
percentages rather than actual measurements when
making comparisons between nematodes that are coiled
into various shapes provides a standardised measure
for differently-coiled nematodes.

Measurements of the Australian populations were
combined to obtain a pooled estimate of the means and
standard deviations, Because these values uppeared
normally distributed, standard deviations of the South
Affican values were calculated assuming a normal
distribution (Table 4).

The positions of the Australian population male
supplementary organs differ in some respects. from
those of the South African population (Table 4)
ullhough (hese differences are not significant except
forthe $. and S,(P<001), These differences appear
minor compared with the similarities that exist between
the populations. Thus the females (Table 3) are of
similar Jength and have a similar vulval position
although the South Affican population appears
narrower with a longer pharynx and a significantly
longer Lail, The females of this species (Figs 5, 6) are
didelphic and amphidelphic, The genital tract (Fig. 6)
consists of ovary, short oviduct, pars dilulala and uterus
that may contain oval-shaped sperm.

Ege laying

The laying process was observed in a specimen
callected the previous day from Lake Alexandrina, Tt
wis iia sitting drop shde in fillered Lake Alexandrina
water (0.2 am membrane). Egg laying took place at
23'C and was very rapid, the actual emergence of the
egg being completed in several seconds. The whole
process was filmed (Fig. 7) (Mord XP L400 film),
The egg is shown moving from the pars dilarata into
the uterus (Fig, 7A, B) and from there into the vagina
(Fig. 7@, DP). During the final stages of laying, the eg
irioves Fron the vagina lo the exterior through the vulva
(Fig. 7E, FG, H). The eps which is oval (ellipsoidal)
within the nematode assumes a spherical shape soon
afer laying (Fig. &), Tt has a mean diameter of 708
ym (42.9 SD) including the shell which has a mean
thickness of &4 ym (+0.6 SD). This compares with
in wero measdrements of (ixed material of 73.5 jn
x 48.9 jun including an egg shell thickness of 5,5 un
(Swart & Heyns 1988).

Discussion
Over 60 years ago Taylor and Poole of CS&IR Division
of Soil Research (now CSIRO Division of Sails)
published the results of a soil survey ofthe bed of Lake

Albert (Taylor & Poole 1931), This work resulled from
a request by the appropriate branches of both the State

A NEMATODE FROM LAKES ALBERT AND ALEXANDRINA 137

TaBLe 4. Comparison of measurements of distances between supplementary organs, expressed as percentages of the sum
of the distances between supplements, in different populations of males of Eutobrilus heptapapillatus.

Parts measured Tsitsikama, Forest Lake Albert Lake Alexandrina

Cape Province, South Africa South Australia South Australia

(Swart & Heyns 1988) (present study) present study)

Rinse Mean SD* Range Mean SD Range Mean SD
Cloaca + ST 76-114 93 +1.6 4.7-11.8 8.5 +2.6 91-118 9.8 +1.1
Ss? + S6 7.6-9.2 3 +0,7 6.0-9.9 8.1 +1.4 8.7-12.3 10,3 +13
S6 * $5 88-113 10,2 +1.1 9,7-12.2 Wal 41.2 10.3-13.0 11.6 10.9
$5 » S4 16.2-17.8 17.4 40,7 16.5-23.1 20.1 42.8 17.0-21.7 19.1 +1.6
S4 + S3 14.4-17.6 15.6 +14 {3,2-15.3 14.3 +1.0 12,3-15,3 13.6 41.1
S3 + §2 19.9-21.6 20.8 40.7 15.3-19.7 Ti? +1.7 15,4-18.2 16.7 +1.0
82 + $i 16.4-20.3 18.4 +1.6 18.6-25.0 20.7 12,5 15.1-20.8 18.9 $2.0

*= estimated using sample size and range and assuming normal distribution

Fig. 3. Photograph of an adult male Eutobrilus heptapapillatus from Lake Albert. Nomarski optics showing pharynx (p),
intestine (i), pharyngeal glands (g), testis (t), retracted copulatory spicules (s). Scale bar = 100 pm.

Fig. 4. Photograph of an enlarged portion of an adult male Eutobrilus heptapapillatus from Lake Albert. Nomarski optics
showing the seven supplementary organs (small arrows) and the everted copulatory spicules (s), Note relatively short
tail. Seale bar = 50 ym.

138 A, TDD

Fig. 5, Photograph of un adult emule Euzobriles heptapapillatas Tran Lake Atbert. Norrshi optics showing pharynx (p),
intestine (i) and vulva (¥). Scale bar = 100 pin

Fig. 6 Photograph ofan entarged portivn ofan adult fernale Eutobrifics hemmapapiiawus tram Lake Albert. Nomarski opries
showine (he didelphic reproductive syste voryisting on etch side of ovary Co} shore oviduct pod), pas dibatata (pal), HePus
tur und vulva ty), Scale bar = 50 pm.

A NEMATODE FROM LAKES ALBERT AND ALEXANDRINA 139

Fig, 7. Photographic sequence of egg laying in Eutobrilus heptapapillatus. Nomarski optics. Arrows indicate. vulval opening.
A.. B. Movement of the egg from the pars dilatata to the uterus. C., D, Movement of the egg from uterus to vagina. E.,
F,, G, The process of laying as the egg passes from the vagina to the exterior via the vulva. H. The newly laid egg. Scale

bar = 50 pm.

140 S & BIRD

und Commonwealth Governments regarding the
leasibilily of using the sail at the buttonr of the lake
lor agriculture after it had been drained. Taylor and
Poole (1931) showed that the drained lake would be
unsuitable for agricullural purposes. Thus. at a time
when the clearing of land was in full swing, these
workers. were able to show, as a result of their snil
survey, condueted under difficult conditions, that
drainage of this lake would have been a costly mistake.
Furthermore, their detailed results (Taylor & Poole
1931) provided valuable information on which to base
further studies of the lake's benthos

Taylor & Poole (1931) reported that lake Albert was
once connected to the saline waters of the Coarang
by an ancient river channel which persists today us 4
lagoon extending from the southern side of the lake
(Fig. J). Thus, although Lake Albert is pot now flushed
through by waters of the River Murray, as is Lake
Alexandrina, it my once have been when the rivet was
in flood

The barrages at the mouth of the River Murray were
not built until 1940 so that at the time of Taylor's and
Poole’s survey ducing Match to April 1930 when the
river was running law, Lake Alexandrina had hecome
very saline as a result of incoming sea water, Thus
water from Lake Alexandrina Was thaking (he water
fron Lake Albert more saline and “was not potable
for humans and taken uowillingly by stock until
accustomed to 1” (Taylor & Poole 1931). Today, due
to the presence of the barrages, Lake Alexandrina is
much less saline than it was and although the quantities
of soluble salts contained in its waler can and do vary
from lime to time depending on river fushings, iL is
clear from the samples collected on the same day fron
both Jukes ata six-monthly interval (Table 1) that Lake
Albert has a higher concentration of soluble salis than
Lake Alexandrina during normal river flow. Nicholas

vt ul (1992) have shown that the concentration of

soluble salts in water collected from sites on the share
of Luke Alexandrina varies from month to (ionth and
hence iL was necessary. to colleet water from (he Iwo
lakes on the same day $o that valid comparisons could
be rhade,

IC is interesting to speculate whether or net the
presence of crystalloid bodies observed jin &,
heplapapillates trom Lake Albert but not in specimens
of the nematode collected on the same day from Lake
Alexandrina when its soluble salt values were low.
might he associated with increased salinity in these
lakes. Crystalloid bodies were [ound in &
heprapapitlatus collected (rom Lake Alexandrina both
ut the water's edge and From the bottom of the lake
in 97% of the nematodes examined (Bird ef al, 1991)
but no correlation Was tade with the salinity of the
lake at that lime. However, examination of these dara
shows that the concentrations of sodium and chloride
ions (Table 2 - Nicholas et ul. 1992) were greater than

those Obtained Troi this lake during the present study
(Table |), Furthermore, contrary te our bindings that
nematodes nwintained in uquaria over ud pericd OF hwo
months “appeared to be Free of erystalloids? b have
found that there was un almost (hree- fold anerevse in
both numbers of £, heptapepittalus and (heir erystalloid
content when mud From Lake Albert was plaved inant
aquarium tank, covered with Luke Albert water and
left for four tnoaths with ovcayional aeration, Under
these vonditions, the ratio of larvae (o lemales lo males
was 85 512) 3 of which 93% contained erystalloids.
The large number of larvae present would probubly
be due lo relatively recent hatching from eps und the
low number of udults to luck of food. The increase in
the percentage of erystalloids present from 32% to 93%
in nematodes kept iy a aquariaum tank for Jour months
would probably be due lo an inerease in the
cuncentration of soluble salts due ta evaporation From
the lank. This partieular butch of Luke Albert water
(eollected on 20 May 1994) had initial sodium and
vhloride ion readings of IK8 and 265 mg 1!
respectively (Table |) which are apparently high
enough to induce the development of crystalloids.
The vechrrenve and possible functions of erystalloids
ina moinber of penera of aquatic free-living nematodes
uve been mécanded hy various Workers (Bird ef ef.

——

Fip 8. Phouaraph fy Of! treshhy laid cas of Enrabriles
the yapapilaius Brighe held opoes showing vulva Cy) and
Sphetical eves (ie) sett (herr refalively thick shelly (esp
Scule bur = 20 pm

a NEMATODE FROM LAKES ALBERT AND ALEXANDRINA ‘4

1941) hur as yet io clear-cut evidence for thetr funetion
has been oblitined,

Clearly fumher reseanch is required (9 lest whether
these corystalloid struwtures ure produeed in’ the
nematodes yp response to changes in thei enyironment
and whether or fut they are a manifestation ala
diseased! slate, since they appear ty be associated with
small regular particles thal resemble idosahed ral
viruses in some respeets (Aird er al. 199),

Tthink that his nematode which predominates 1 the
sediment at the bottom: of these Jukes is E
heplapapiliaias altnough, ax will be discussed below,
there are-some diflerences hebween Lie Australian anc
South African populations. {has ull the characteristies
of its Subfamily (Rutobrilingeé) deseribed by Tsalolikhin
(1983) namely, hedgehow- like supplementary organs
(Fig. 3), a muscular yagina and Well-differentiated
female genital system (Fig. 5) and well-developed aod
rounded pharyngeal whinds. Sintilarly, tis less than
3.5 mm in length ind it has the described species
characteristics of a cuticle withant pronounced
annulations, head bristles that do not exceed 14 pny
and, most obvious of alf, mules with seven
supplementary organs. Tt does nol quile Ail Tsalolikhin’s
key in having females whose tails are not len lines
greater than the body width at the anus (‘Table 3) but
(his also upplics tothe South African spectiens (Swart
& Heyns 1988). However. these various morphotogical
Jiflerences ure likely to be reflections of variability

hetween different populations of this species of

nerhatode rather than suggesting thar the Australian and
South African nematodes are different species.
Another apparent difference is thal the egg. although
oval-shaped when within the female. becomes round
wher laid (Figs 7, &). However, it Seems likely that
cue measurements in the past may have all

been made while the eggs were within the female in
fixed material. The transition from oval within the
female to spherical an laying is, however, illustrated
by Tsulolikhin (1983) in his book, Since eggs of these
species can not be identified unless laid from an
identified female, their shape for laxonomie purposes
is fisted in the deseription of the genus Futobrilus as
oval (Tsalolikhin 1983), Furthermore, since fixation
leads to shrinkage, Measurements from fixed material
will always be lower than those for unfixed, freshly
laid eges.

The combination of characters described above
places (hese womis firmly in the Tobrilidae, Minor
differences between the populations in South Australia
dnd South Afnea ure nor considered significant and
the worms are confidently referred to as &.
heprapapillatas.

Much remains to be learnt about this cosmopolitan
nematode. Its feeding habils, rates of growth and
longevity im the lakes wre unknown. Such information
is needed af its value as-dn indicator of envirunmentil
pollution jn these cnvjronments is te be determined.

Acknowledgments

] wish to thank Angela Reid, CSIRO Biometrics Umit
for the statistical analyses, the Soils Division, CSIRO
for accommodation, facilities and expertise, including
that of Adrian Beech, and the Zoology Department.
University of Adeluide for facilities and equipment and
especially for the help of lan MeGruith. | thunk Jean
Bind for help with collecting and Jean Bird, Lester
Cannon, Mary McHugh and Warwick Nicholas for
constructive criticism of |he manuscript, This work
Was made possible by a grant trom the Australian
Biologival Resources Study for which L am most
grateful.

References

Rinn, A, FE, MeCiiure, S G & Nicuor as, WoL (I99T)
Observalions on crystalloid bodies in (he pseudocoelom
OL kufobrilas heptapapillatus, J. Nemetal. 23, 39-47,

Di MA sSeneeR, 1, & D'Herpe, J, (1963) Méthodes utilisees
pour VPetude Wea anguillules libres duo sal. Revur de
Cagrieuliure, Bruxelles We. 44-4 47_

Hooree, Do 4 O86) Handling. fring, staining und
mounting nematodes pp. SY-80 i Southey. Jk (Ed)
“Luboratory Methods tar Work wih Phint and Soil
Nematodes” (HMSO, Londont,

Nicnonas. W, L, Bian, A, Ry Bercy T, A, & Siewari,
A.C. (1992) The nematode fauna of the Murry River
egtuary, South Australias the etfeets of the barrages across
iis mouth, Midroblelevie 234, 47-101,

Spinniorst, J. W. (S90) 4 rapid methad for fhe transfer of

nemitodes from fixative to anhydrous glycerin.
Nematolovica & 67:69.

Swart, A, & Hbyns, J, (988) Redescription of Eutebrilus
heprapapillaris (Joubert & Heyns, 1979) Tsalolikhin, 1981
With notes on is morpholagy and possible exeretary system
(Nematoda » Tobrilidae). Phytophylacica 20, lol-lok.

TVavior, . K. & Poore HG. (1931) Report on the soils
ofthe bed of Lake Albert, South Australia. Coun. setent
inet, Res. Aust. 4, 83-95,

TSALOLIEHIN, S.J, (983) “Nematode families Tobrilidae and
Tripylidae world fiuna” (Nauka, Leningrad),

Zarcinas, BA, & Cartwrioni, B. (1983) Analysis of soil
und pliant material hy inductively coupled plasma-opticat!
emission spectrometry, Division of Soils Tech, Paper No.
43. CSIRO, Ausinalia

DISTRIBUTION OF SPECIES OF TRICHOSTRONGYLOID
NEMATODE PARASITES IN THE SMALL INTESTINE
OF THE BUSH RAT, RATTUS FUSCIPES

By L. F. SKERRATT*, I. BEVERIDGE* & M.-C. DURETTE-DESSETT

Summary

Skerratt, L. F., Beveridge, I. & Durette-Desset, M.-C. (1995) Distribution of species
of trichostrongyloid nematode parasites in the small intestine of the bush rat, Rattus
fuscipes. Trans. R. Soc. S$. Aust. 119(3), 143-148, 30 November, 1995.

The distribution of three trichostrongyloid nematodes, Nippostrongylus magnus
(Mawson, 1961), Odilia bainae Beveridge & Durette-Desset, 1992 and
Paraustrostrongylus ratti Obendorf, 1979, in the small intestine of bush rats, Rattus
fuscipes, was investigated. Each of these species exhibited a significantly different
longitudinal distribution within the small intestine. Interactions between the three
species, identified by comparisons of the fundamental and realised overlaps in
nematode distributions, were the probable cause of the differences in distribution
between species. The different distributions, which are here interpreted as niches,
occupied by the nematode species are consistent with the hypotheses that O. bainae
was probably a parasite of hydromyine rodents which filled a vacant niche when it
switched to R. fuscipes as a host, while P. ratti probably occupied another vacant
niche when it switched to R. fuscipes from an original marsupial host.

Key Words: Parasite, ecology, niche, Trichostrongyloidea, Rattus, interaction.

Transactions if tie Reval Socters of 8 aus (1985)

W193), a 148,

DISTRIBUTION OF SPECIES OF TRICHOSTRONGYLOTD NEMATODE PARASITES IN
THE SMALL INTESTINE OF THE BUSH KAT, RATTUS PUSCIPES

by L. F Skerreary* |. BEVERIDGE* & M.-C. DuRWITh-DESSETT

Suinuniary

Skearar), Lh, BEYERIbeb 1, & DURE Dressel, MC (1995) Distribution of species of trighostrongyloid
Hermulode parasites in the small antestine of the bush rat, Rats fascipes, lramy. Ro Sac. 8. Aust. W949). 45-148,

AQ Novyerber (995

‘The diamburion of three michostrngyloid nemalodes, Mippeytronuviay meinies (Mawson, 961), Odilies battae
Beyerdwe & Durctlo Dosse( OZ und FRaraustrostrniies haiti Obendert, 979 in the small intestine of bush
rals, Rares fieseipes, was investigated Buell of theso species exhibited a signifivantly difleren) fongiwdinal distribution
Wilh the small iitesmme lntertictinnas berween the three species, identified by compansens of the fundimental
and realised Ovceluprs je nematode dishibulions, were the probable cause of the differences in distribution between
species. The different distributions. whieh are bere interpreted as niches, oceupied hy the nemutnde species ure
consistent with the hypotheses (hal GO bamtee wis probably a parasite ol hydromyine rodents whieh filled a vacant
Hiche when U swatehed te dk fseipesas a ost, while PB rane probably oceupied andther vacunt niche when
W switened to Ro faserpes fron an urigenal inarsupnil host.

Rey Wokos, Panuile, ceolawy, quohe, Trichostraaeyloideu, Ralls. iferactan

Introduction

One mode by which parasile evolution may occur
is “host switvhing” (Chabaud 1965). This involves a
break-down in host speciticity allowing the transfer
ata parasite from its asaal host to an unrelated Host
species necupying the same environment. The new host
may be infected trough dre skin by free-living suiges
af the parasie or may ingest the infective form of the
parasitewith its food (Chabaud !965), The mechanism
of host switching appears to be common among
parasiie neniatodes (Chabaud 1982) and (§ based on
the ussurmption that the invading nematode parasite is
veeupying a previously vaedut niehe within the new
host, In the case of intestinal parusites this is. usually
defined as a restricted longitudinal of radial distribution
within the gulof the host (Schad 1963), Host switching
within the nematode superfamily Trichostrongyloidea
is well documented (Durette-Desset 1985), yet few
studies huve examined whether the invading nematode
acrially occupies a separate or previously -vaeant niche.

The {richostrongyloid nematode pitrasiles of the
native bush rat, artes fieselpes, olfer the opportunity
ro examine such an hypothesis. At one locality in
Victoria (Blackwood) Obendart (1979) found that A
fuyeipes wis parasitised by three species of
trichostrongyloids, the heligmasomes Nippasrrangylas
rhavnus (Mawson, 1961) and Crilia bainde Beveridge
& Duretle-Desser. [942 and (he herpetostronpylid
Paraustrosrrangyius rai Obendorl, 1979 (Obendart
1979; Beveridge & Durette-Desset 1992 a,b, 1993). P

University of

* Departmen) ot Vetenmary sciences
Melbourne, Parkville, Vie. 3052.

tT Laboratoire de Biologie purwvidure, Muscun nahanal
U'Historre nitluretic, Pures, 75005, France

rat! belongs to a genus which otherwise occurs
exclusively in marsupials and which presumably has
switched to its-current eutherian host (Obendorf 1979),
Iris considered (Obendort 1979) 10 be one of only Wwe
wuumples of trichostrongyloid nematodes switching
from marsupials to native rodents, the other being
Roolleva hydromyos in the water rat, Aydromv chryse-
waster (see Mawson 1961. 1973). O. bainae belangs
tow venus whieh is paraside primarily m hydromyine
rodents and species of the xenus were considered by
Durette-Desset (1985) to have switched secondarily toy
murine rodents such as Ro faveipes. Only Nippo-
strongylis magnus can be cunsidered an original
parasite of this murine rodent (Beveridge & Duretie-
Desset 1992).

The current study was therefore undertaken to
determine the ecological niches occupied by N,
magnus, O. bainae and P raiti within the small
intestine of R, fuseipes and to examine the extent of
overlap between them to establish whether or not each
mecupies a distinctive intestinal niche.

Materials and Methods

Ten bush rats, Rartus fiuscipes, were trapped front
along the bunks of the Lerderderg River, Blackwood,
Victoria, Australia (37° 29° 5, 144° (9° B) using
collapsible aluminiuny traps baited with peanut butter,
Immediately following euthanasia wilh chloroform, the
small] intestine was removed and divided into sixteen
equal parts. The rotal length of the snail intestine was
measured. Gul segments were incubated in saline at
37°C for at least 2 hours and all nematodes which
emerged fram the mucosa were fixed in hot 70%
ethanol betare heing counted. P rat! was distinguished

144 L. F SKERRATY, |. BEVERIDGE & MO DURETTE-DESSET

from other Species based on the deseriptinn of
Obendorf (1979) using a stereomicroscope, Whereas NV
Magnus and OL baihae were differentiated Using the
Uescriphions of Beveridge & Durette-Desnet (1992 a,b)
and a compound microscope following clearing i
lactophenol, The rats were also routinely examined for
other helminth parasites,

The number of cach species of neniatodes. in
individual sections of the small intestine was converted
to a percentage of the total number of each species
present. For cach species, the positions of the anterior,
median and posterior nematodes were deterinined
using the method of Bush & Holmes (1986), such (hat
the section number in which nematodes oucurred was
converted to a percentage of the total length of the smiul!
intestine. It was assumed that nematodes were
uniformly distributed within each seetion.

Differences m distribution between species were
tested Statistically using values calculated [rony a 2 %
5 contingency luble (species x sector of intestine) lor
each species pair. Sections 5 to 16 of the intestine,
where very few nematodes were found, were cojnb|ned
to form av single cell in the table,

The extent of niche overlap between the three species
wus determined using the equation

Cay = i- 1 CHIP — Pyjl)

where Pri Py (Hurlherl t97s)
lex , i= Yi
gy

such that Pxi and Pyi are the proportions of twa
species, x and y, in different segments of the intestine

This equation was used to calculate the *fundaniental
overlap” belween species pairs, which 1s the extent of
overlap (Cxy) io the mean disoributions af the wo
nematode species and the “realised overlap” which ts
obtained by determining the extent of overlap (Cxy)
between two nematode species in individual rats and
then computing the average of the individual overlaps

Differences between fundamental and realised
overlaps. that is when the realised overlap was substan-
Hally Jess than the fundamental overlap, were used hy
infer the presence Of competition between nematode
species,

Seven laboratory rats, RL norvegicus, were infected
either subcutaneously or orally with 200 - 900 infecuve

Tabre 1. Positiore af mean anterior median and postin
individuals (+ slandard errr of mean) of Sippoxtrongy lis
magnus, Odilia bainae and Paraustrostrongylus rally os
percentage distances along the small intestine in ten narurally
infected bush rats, Rattus fuscipes.

Anterior Median Postenor _
No magnus 1.01+0.64 1[5,14+4.84 5663+ 10,17
O.bqinae = 2.1740.84 182349.07 4840+ 6.47
Porat 3,22 + 1.62 19.064 2,25 49.40 + 4.47

larvae Ol No srasitus ar (2 hartge (Table 3), The
infective lame were oftained by culling a mixture
ol faeces Ip naturally iniected rals with wolivalel
Chareual on morsi filter paper and recovering developed
larvae by sedimenkifion in waler Larvar wen
Separated On the buss af morphological differenaes
(shape of the tail) identifiable using a Sterermicraxcopa:
(unpublished observations), Infected raty were killeal
with chloroforn) 14 days after infeetion and the
distribunon of nematodes in the sunull inrescine
determined ina similar fashion to thal described dbove-

Expertitental auteetns wilh Poreifh via oral.
subcutaneous and perculaneaus, Toutes were
uinsuceesshil

Results

Nemataite disrriherions in wile rts

No magniity occurred in 100% nf rhe wild bush. rats
examined), with a metin intensity of 67 uind.a range of
L-+183, ©) fara was present in 00% of wild rats
examined. with a mean tatensitv of 64 and a range of
LTS. nat was present in 0% ot wild rats examined
with a medn intenwity af I7L and a range of 11094
(Tadle 4),

Oder parses found in the (en qaturally intbetee
bash rats were very small numbers. of Capillarica sp,
(Nematoda) in the first segment of the sanall intestine
and Cupelfortid gusrrica (Baylis, 1926) within the
squamous epithelium al the slomach in five rats. There
was no patholowical reaction fo wlll Cepillaria or eee
wiltiin the squamous epiihelium, Aerenikix spumoast
(Schneider 1866) was found us the lange intestine of
eight rats. The cestodes Chuaneaueniad ritticate
(Sandurs, 1957), Hymenclepw cdininura (Rudalpby,
(819) and Berfiella anapolvtine Baylis, 34 were found
inthe small intestine ©. raifieela inhabtied the opening:
of the bile duet. causing hyperplasia of the tile duet
epubelum 4, diminutd and B anupolyned were found
in segments 412 und SIG respectively. However,
ditilhuia was distributed anteriorly te B. anapialyrieg
when bath species oceurped jn rats.

N, rhayruty occurred mainly (64%) in the anterior

two segments of the ered! iresting in naturally infected

rats. WIT Maximum intensity in the first segmenr, and
its Numbers declined progressively in the rerwiming
sections (Fig). The majority of O, buinae (82%)
inhabited the five most alteriod segments in naturally
jatected bush cats with a maximum intensity (30% |
in the second seement (Fig. 1), The majority of PB raili
(87%) were distributed throughout the five anterior
segments af the sonal aitestine in muturally infected
tats (Pig. 1) with maximum intensity deeurring it
segments | ro 4. Simall intestine lengths ranged from
56-97 em (mewn 71 em)and hence the average length
of cach segment Was 4.5 ei,

TRICHOSTRONGYLOID NEMATODES OF THE BUSH RAT

Nippostrongylius mugnius

bainae

Odilia

ratlt

Paraustrostrangylis

‘Mean percentage of nematodes in each section

Section of small intestine

Fig 1 Distribution (mean percentage + standard error [error
bars|} of Nippostrenzylus maynus, Odilia bainae and
Paraustrostrongvius radi in each segment (sixteenths) of
the small intestine of ten naturally infected bush rats, Reis
fuscipes

N. magnus, O. bainae and P. rary were sequentially
distributed along the small intestine of naturally
infected bush rats but differed in the positions of [heir
Inegn anterior, posterior and median individuals (‘Table
1). The mean anterior and median individuals of N,
magnus Were anterior to those of OQ. hainae which were

TABLE

Fundamental and realised overlap beiween niches

145

more untenor to those Po rari. However, the meun
posterior individual of NV. mayniy was posterior to both
O. bainae and Prati. This was due to Ny magnus
inhabiting the posterior quarter of the sniall intestine
in two rats, Whereas O. barnae and Po ratti were not
found in this segment. The distributions of the three
nematode species were significantly different. Chi-
squared values obtained for pair-wise comparisons
were N. magnus - O. bainae, y= 77.5 (p< 0.001).
O. bainae - Prati, x 90.9 (p<0,001), N.
magnus - Po raiti, %~ = 18.5 (p<0001), There was
no correlation between mean posilions of nematodes
with intensity of infechon, except in the case of the
posterior position of QO. bainae (vr 0.68, p =
0,004).

The varialion in numbers of trichostrongyloid
nematodes in each segmentof the intestine of naturally
infectéd bush rats was large (Table 4), with the standard
deviation equal to or greater than the mean (sce
standard error burs in Fig. 1). Despite this variation,
the difference in distribution was greater in indryidual
rats (1- realised overlap) than the difference in their
mean distributions (I fundamental overlap). The
realised overlap was lower than the fundamental
overlap in 25 of 28 species interactions, The mean
realised overlap between N. magnus and Po ratti wis
lower thin the overlap between N. magnus and O.
bainae and between O. bainae and P. ratti (Table 2).
The total numbers of each species of nematode in
individual rats and the realised overlaps in. natural
infections varied between individuals but the two
purameters were independent of one another (r<
0.3). Capillaria sp. occurred in such low numbers that

of Nippostrongylus magnus, Odilia bainae cre

Paraustrostrongylus ratti in the small inrestine af ten naturally infected bush raty, Rattus tuscipes.

No. of rats infected

Nematode species } C
with both species

pair

Fundamental overlap Realised overlap

(+ standard deviation)

N. magnus — OQ. hainae 10
N. magnus — P. ratti 9
O. bainae — Prati 9

0.68 0.55 £0.14
0,61 0.40 + 0.27
0.82 0.55 + 0.23

TABLE 3% Percentage of Nippostrongylus magnus aad Odilia bainae vecurring in /6 segments of the small intestine of
lahordtory reared Rattus norvegicus following oral or percutaneous infection with third stage larvae

Segment no. of intestine

Mean percentage of nematodes in segment
(+ standard error)

Nippostrongylus magnus

Odilia bainae

No. of rats 4 3

T Bt T3 OTOF3.8

2 2342.2 7.0 +6.7

3 25425 2.342.3
4 10.0 + 6.8 Q
5 16 0 U

L. fh SKERRATT, 1. BEVERIDGE & M.C. DURETTE-DESSET

Pat d. Numbers of nematodes, Sippostrongylus magus. Odilia baynse ad Paraustrosicongylus ralli in aectiony (EG) uf
thre sinalt testing ef ten materally pifected husk rats, Rattus Tuscipes,

— Rat number
5

Secyon I 2 3 4 t 7 4 i) i)

)

MWS ——s vs
} 14.5.0 0.0, 21 5.01 wa 5.37 2.221000 43.2,2 37,31 49,4238.
2 (0,0 HOS 6140 14,3,5 16,0 8,310 A3136 21h) BATT AN, TG Fhe
} 2.0,0 26,2 19.0 A432 336,27 O06 49,25 ALO 13.7.4) 11,2.228
4 230 11,2 w70 KIS 12,0 1.0.12 a7,56 FAB 312.26 9,5.133
4 U,2,0 0.0.0 61.0 35,18 (13.1 0.9.0 4216 10 31.34 9278
b 0.1.0 0.0.1 0.0.0) 144 24,0 0,6,1 0,1.22 O14 1,17,10 43,34
7 0.0.0 00,0 0,00 10,4,18, O10 07.0 O07 O54] 030,11 2.015
bs 0,0,0 0,0,0 0,0,0 214 25,0 0.12 oO1s 0.11 6.25.2 1.2
y) 0,0,0 (0.0 0,00 21.3 2.2.0 0.00 0.0L O01 aval 0,04)
Tt) 0.0.0 0,0,0 0.0.0 int ina 0.04) 0.00) 00,0 (231 0,00
| 0,0,0 40,0 0.0.) 1a! Lhd 0,00 0,00 0,0,0 (2131 0,00
12 ),0,0 (0.0 (,0,0 10,1 1,10 0.00 0,00 (00 12.130 0,00
8 1,0,0 0.0,0 (1,00 10,0,0 00,0 0.00 0.00 0,0 10,0,0 0.0.0
\4 0,0,0 ),0,0 0,0,0 9.0.0 00,0 O00 0.0.0 0.00 10.0.0 0,00
5 U.00 0.0.0 6.0.0 90.0 0.0.0 0.0.4) O00 0.00) 9,0,0 0,00
if 0.0.0 0.0.0 00,0 90,0 0,00 00,0 0,00 (0,0 9.0.0 00,0

Total 20000 Uh1614 16.361 96.22.70 491033 147744 577378 99-6002 KITA 46 125,92, 1044

its passible interactions wath the trichostrongyloid
pemmlodes Were not considered,

Nemeterle distributions in experimentally infected reais

No magnis occurred primarily in jhe anterior
seemient of experimentally infected luboratory Fat, with
a relatively small population of nematodes established
in segments 2 to 4 (Table 3). Similarly, O. huinue
became established primarily in the first segment of
ihe miestine, wath small numbers of nematodes present
in segments 2 to 4. The mean intensity of infection was
WW lor NM meenus and i for QO. bainie.

Discussion

The significantly distinct sequential distabutions of
(he three species of trichostrongyloid nematode along
the small intestine i natural infections and
experimental micetions suggest that each species
oceupies a distinct niche. Furthermore, the
fundamental ovetiaps in natural infechors between the
species pairs \. negnus. - QO. bainae (68%) and N,
magnus - Prati (6l%) were lower than the 70% yaluc
suggested by several authors (Pianka er al, 1979:
Holmes & Price 1980: Bull era/. (989) to indivate the
eustence Of ecologically relevant differences. Only the
fundamental overlap between the species pair. O.
bainee - Po rani (82%) was greater than 70%.
jlowever. the mean realised overlup (55%) between
jhese two species was substantially less than 70%,
imficating that these nwo species interact to separate
their niches in individual rats, The distribunion of C,
bainae in experimental infections (Table 3) differed
when compared with natural infections (Pig. 1) in thur
in MOnospecific infections i occurred in the mist
walerior segment of the duodenuin and (his may be due
ti Ihe absence of competition from NM) meninuays ort

nati. However, other contributing factors may have
been the different species of host, te smaller sample
size in experimental infections or the smaller numbers
of O; bainae in experimental infections.

These observations sugwest that the nematode
community in A. fiiseipes is an interactive one. Holmes
and Price (1986) separated communities pf parasites
into two categories, isolationist and interactive, based
on their infrapopulations, that is, populations in
individual hosts. They suggested that un interactive
community has no vacant niches, purasiles are mot
distributed independently and realised distributions of
parasites are dependent on other guild members. Sotne
of these features are present in the cause ol the
(richostrongylou nematode parasites of the bush rar,
since the realised overlap was less than the fundamental
overlap in most interactions in natural infechons.

However, (he small intestine of the bush pat does
appear to have vacant niches in natural infections
despite (he above evidence fur an interactive
community. Although host immunity may reduce the
size of apparent yvacunt niches (Noble e¢ af 1989) and
low (riosmisyion milessniay prevent parasites filing all
available niches (Price 1980), the distribution of the
trichostrongyloids in individual rats wis independent

of nematode numbers. Thus, assuming vacant niches

over. the trichostrongyloids of the bush rat alse
demonstrate one feature of an isolationist conmmunily
(Holmes & Price I986). The “populatian
concentration” and “individual response” hypotheses
both explain why the species NV. magnus, O. hainee
and P. rari should occupy distinct niches even when
additional yvacunt niches are available (Holmes & Price
1986). The “population concentration” hypothesis bas
two components, that narrow miche oeeupabon. ts
exsentisl for the maintenance Of intraspecihe contact
for mating purposes (Rohde 1979, 1982).and that the
Qecupalion Of diserele miches is important as a

TRICHOSTRONGYLOID NEMATODES OF THE BUSH RAL lAT

reproductive isolating mechanism preventing
hybridization (Sogandares-Bernal 950° Martin 1969),
The “indiwidual response” hypothesis (Price 1984)
argues that parasites adapt wo the environment they
inhabit and consequently Mil nurrow niches.

The distribution and overlap of trichostrongyloid
nematodes jn individual bush rats Vary greatly from
the means but are independent of nematode oumbers
exceptin the case of the posterior extent of Ch byinue
in the imestine. The varrability in these
intracommiunities may be due (o the biological features
of the nematodes and their interactions with one
another but may also be caused by variability in the
characleristioes of the host which influence parasite
infraenmmunities such as host diet (Croll 1976) and
blood supply (Croll & Ma 1977), The non-specific host
response, pathological responses and acquired immune
responses of the hast may alse influence the distribunen
ol parasites.

The three tichostrongyluid nematode specics found
in RK. fiiscipes have different biogeographical origins.
The genus Nippestrongvluy occurs primarily in Rarfees
spp. in south-east Asia and in other rodents in Asia
and the Middle-East. with a single species ul
dermopterdns (Beveridge & Durette-Desset 1992).
Since the endemic species of Ratrys in Australie
probably reached the continent from south-east Asia
(Waos & Astin 1981). it is likely that Nipposrrne vies
reavhed Australia with them and that subsequent co-
specuition led ta the evolution of N, magiis in R.
fuseipes (see Beveridge & Duretic-Desset 9924). The
genus Odilie occurs pomiurily in hydrogyine or “ole
endemuc” rodents in Australia, principally in the genera
Melumys and Uromys. Species occurring in Rares spp-
have been interpreted as transfers From “old endemic"
rodents, which probably evolved between S and 15
Million years ago, to the “new endemic’ Rariys spp.
which have been presen! on the continent for about
Loanihon years (Watts & Asin 1981). Species of
Paraustrastrongylas occur in possums (Phalangeridiac.
Petuuridae, Burramyidae) and rat kangaroos
{Polorgidae) (Spratt ef a/, 199)), wilh single species,
P rantina rodent. The transfer of Puraustrastrongyius,

und probably Odilia, ss therefore presumed to be of

revent origin (ess than | million years),
Holmes (1973) supgested thar stable communities are

older than imterachive ones and since most parasite
comraunities dre stable, he coneluded thal they ure
relatively old, The interietive component of the
tnchostrongylod intracommunity in bush rats suupests
that iC is a Comparatively young community: This 1
consistent with the hypothesis that 2 rans und possibly
also O. haihae dre réevent invaders. Following their
invasion of R. fiuscipes. PB raw and O bajnar have
occupied distinctive niches within the new bost
possibly duce te isolationist lorees and/or their
interactions with the other trichostrongyloid nematode
species present in the small intestine. Tt 1s possible that
P raiti colonised & previously vacant mehe at the Lime
of switching since No meagaas and ©. bainae do nol
occupy this niche even when A rai is ubsent

The wther parasites found in the small intestines ol
bush rats. appear to occupy completely different niches
from those inhabited by the (richostrongytoid
nematodes or ta occur at a very low Tatensily and.
prevalence (Cupillaria sp.) and therefore were pot
vonsidered in the interactions of the trichastrongyloid
com@munily in the present study, The distributions of
A. dinijniuta and B. avapolytica overlapped, However.
the Ko species were never found inthe same segment
of the small intestine although only two fats were
iMfected with both species, Beewuse of interactions
which tay occur between ther, they may occupy
sepurale niches (Holmes 1973), A larger sample ov rats
would be needed to examine the extent of interaction
between their cestode parasites.

Acknowledgments

Christine Anderson and Lee Bergerare thanked for
thei help in tapping rats and collecting the nematodes.
Robin Gasser is thanked for his adviec and help with
trapping and Neil Chillon tor comments on drafts ol
the paper, Rats were collected under permit no
RP9I-095 from the Victorian Department of
Conservation and Natural Resources and experimental
procedures carried out under The University of
Melbourne, Animal Experimentation Ethics
Committee no, 270-056-0-89-077) The work was
supported financially by the Australian Reseureh
Council

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Spratt, D, M., Beveripor, I. & Warrier, E. L. (1991) A
catalogue of Australian monotremes and marsupials and
their helminth parasites, Rec. §, Aust. Mus., Monogr. Ser.
1, 1-105,

Warts. C. HS. & Asus, H. J. (981) “° The Rodents of
Australia” (Angus & Robertson, Australia).

TRANSACTIONS OF THE

ROYAL SOCIETY
OF SOUTH AUSTRALIA

INCORPORATED

VOL. 19, PART 4

EIGHT NEW SPECIES OF AUSTRALIAN BUPRESTIDAE
(INSECTA: COLEOPTERA)

By §. BARKER*

Summary

Barker, S. (1995) Eight new species of Australian Buprestidae (Insecta: Coleoptera).
Trans. R. Soc. S. Aust. 119(4), 149-156, 30 November, 1995.

Six new species of Castiarina namely C. corallina sp. nov., C. ernestadamsi sp. nov.,
C. euknema sp. noy., C. octopunctata sp. nov., C. oedemerida, sp. nov. and C. prolata
sp. nov., one new species of Themognatha, T. viridescens sp. nov., and one species of
Astraeus, A, powelli sp. nov. are described.

Key Words: Coleoptera, Buprestidae, New species, Castiarina, Themognatha,
Astraeus.

Traicdertons af the Reval Soerety of So Ast (99S). WY 149-456

EIGHT NEW SPECIES OF AUSTRALIAN BUPRESTIDAE (INSECTA: COLEOPTERA)

by §. BARKER*

Summary

BaAkkeR, 5. (1995) Fight new speeies of Aostrahan Buprestidae (Insects, Coleapten) Trans. R. Sav, S. Aus,

119(4), 149-156, 30 November, 1995,

Sik new apecies of Casriarina namely C. cerallina sp. nov, C. ernestadamsi sp. noy,, Co enkriertia sp tov,
Co sctupunctaa sp. poy,, C eedemerida, sp. nov. and C. prelara sp. noy,, one new species of Themognailia,
L viridescrns sp. nov,, and one species of Asiraews, dy powell sp, nov. are deseribed-

Key Worps: Coleuptera, Buprestidae, New spevies, Confaeiia, Memoynatha, Astraeus

Tatroduction

Despite the increase in specialist collecting of
Australian Buprestidae undertaken by amateur
entomologists over the lust twenty years, new species
are still being found. This reflects the vastness of the
conlinen( as well as the cyclic nature of the lle histories
of many of the arid area species, OF the species
deseribed herein, specimens of Astrecus powelli have
been available since 1970 but the specimens im the
South Australian Museum collection were
madvertemly sorted into a tray full of A, aberrany ¥,
de Poll, the closes! species, und it was only through
the alermess of Me M_ Powell thal this species has heen
recognised. Jemounathe virtdescens was collected
many years ago in inaccessible country ut Jron Range,
Cape York Peninsula, A serics bas now become
available from the Jack Macqueen collection, lodged
with the Australian National Inseet Collection (ANIC),
Three species. have only reeently been collected.
Castiavina pedemerida and C. pralata in Queensland
und C. corallina in Western Australia, Castiarind
euknema was known lor some years from two female
specimens and a male has only been located recently
in the collection of ANIC. Castrarina vrtapuncrstia
specimens have been collected frequently but until now
have not been distinguished from C. paralleta. Until
recently C, ernestadamsi has been confused wilh C.
srramined Macleay,

Materials and Methods

Male genitalia were prepared and displayed by the
method described by Barker (1987), The holotype ts
ilustrated in all species. Measurements given are mean
total body length and width with standard error, except
Where there are insufficient speeiniens availible to
niake the last calculation, Codens used in the texr for
muscuntand private collections following the four letter

* Departinent of Zoology, CU niverstry of Adelaide 5, Aust
SOS.

system of Watt (1979) and Arnett et al, (1993) ure:
AIMS: Australian Tnstitate of Marine Science,

Townsville; ANIC: Australian National Insect
Cullection. CSIRO. Canberra: SAMA: South
Australian Museum, Adelaide: WAMA: Western

Australian Museum, Perth: SHIQ: Mr J, Hasenpusch,
Innisiail; MASA: Mr T. M, S, Hanlon, Sydney;
MPWA: Mr M. Powell. Meiville: GWQA: Mr G.
Wood, Athertan.

Castiarina corallina sp. cw.
(FIGS 1D, 2C)

Holawpe> oro \44 kin NW Wittenoom, W.A..
25.11.1994, M. Golding, M. Puwell, WAMA.
Allorype: 2, IO km NW Wiltenoom. W,A,,

25.11).1994, M_. Golding, M. Powell. WAMA.

Faratypes; 2-2 9, Millstream Stn, W.A., 25,171.1994,
M. Golding, M, Powell. MPWA: 1 co’. same data as
allotype, MPWA: 4 cr cr. 145 kin NW Wittenoom.
MPWA, SAMA,

Colour

Head. antennae dark blue Pronotum red-brown, in
some specimens with medial smudge of dark blue
along basal margin, Scutellum dark blue. Elytra red-
brown with the following markings: dark blue along
busal margin, in some Specimens with mark extended
fosurround scutellum) large post-medial spot on each
elytron, apical mark. Ventral supface: pre-sternum red-
brown, meso- and meti-sternum dark blue: abdomen
red-brown except laterally at base variably dark Blue,
legs dark blue.

Shape and sculpture

Head closely punctured, broad median sulcus, short
muzzle. Antennae. antennameres |-3 obconic, 4-11
wwothed, Pronotum closely punctured. narrow basal
fovea, apical margin projecting medially, basal margin
bisinuate; laterally pardllel-sided at base, ungled
outwards, rounded to widest pre-medially, tapered to
apex. Scutellum cordiform, glabrous, tat. Elytra
punctale-sirtale, intervals conver, punctured: laterally

150 5. BARKER

paralfel-sided al base, rounded ut humeral callus.
coneuve rounded post-mediully. narrowed to spineless
apex, apices hardly diverging, apical margin subser-
rate, Ventral surface, shallowly punctured, edges of
abdominal segments glabrous, withoul hairs, $,:
upically rounded in both sexes,
Stee

Males. 4.3 + O48 4 5.4 + 0.14 mm (6). Females,
16.8 x 6.5 mim (3),

Aedeavus (Fig. ID)

Parameres angled outwards from basal piece,
rounded apically, Penis pointed. sides acutely angled
away. Apophysis of basal piece medium width, tapered,
rounded apically,

Remerrks

The dorsal tnarkings of this species resemble those
of Cusdarina quadriplagiata (Carter) which has only
heen collected in Queensland. However, C. cerallina
in 4 narrower species, it is a paler red than C.
yuudriplagiata and the male genitalia differ (Pig. IC).

Epymolagy
Derived from L. eeraflinas, coral coloured.

Castiarina vedemerida sp. nov
(FIGS 1H, 2F)

Holonpe- oo, Georgetown, Old, 284).1993, |
Hasenpusch, SAMA I 21285,

Paratype: Qld: lor. Georgetown, 10.1993, J
Husenpusch, JHIJA.

Colour

Head black. Antennae dark blac. Pronotum brow
with the following black markings: medial spot,
smaller spot on cach side. narrow basal border,
expanded anteriorly on each side. Seuteluin black,
Elyva yellow-brown with the following black
markings; M-shuped mark along anterior margin, ats
covering humeral callus; post-medial fascia reaching
margin. expanded anteriorly and posteriorly along
suture, apical) mark. Ventrul surface: prostermum
yellow-brown) meso-sternum and imeta-sternum and
coxaue dark blue; abdemen yellow-brown, 8. with
lateral black spol, §, with lateral black bar, 8, black
apical edge, edges of S.. S,, S., testaceous: legs dirk
blue. Hairs silver.

Fig. 1. Photmicrographs of tnale acdeagi ot the following Casriarina, Astraeus and Themognarha species. A. Themognathe
viridescens, B. Castturina acuminata Kerrenians, C. Custiarina yuadriplagiahs Carter. D. Casdurind corallina sp. nov.
F. Castiarina prolata sp. noy. F Castiaring eciopunctaia sp pov, G. Astraeus powelli sp. nov. HL Castiarina cedemerida
sp. nov, Lb Casdarina ernestedamsl sp. nov, J. Casnerina srraminca Saunders. K, Castiering eukneme sp, nov, |. Castaring
rufipes MacLeay. Photomicrographs of procigers of types of Castiarina ectapanetata sp, poy, M. Holotype mile, N.

Allotype temale.

EIGHT NEW SPECIES OF AUSTRALIAN BUPRESTIDAE 151

Shape und sculpture

Punctured over entire dorsal surfiee, Ted closely
punctured, median suleus, short muzzle, Antennae.
intennomeres +3 obconic. 4 Vs toothed, Sl toothed.
Pronotum closely punctured, glabrous urea
surrounding basal fovea, fovea at each angle; apical
margin stright, basal margin barely bisinuate: laterally

rounded from base to apex, widest pre-
~~ ~ i
. f i
{2
a " A ‘a
| {
i- . x

Fig. 2.
species, A.C ernesiadamsi sp. nov holoiype. B.C. yirigeta

Habitus ilustrations of the following Crsiarinia

Macleay, C. C caralling sp. nov, holotype. BO &
Herp tit Sp, nov, holotype. b. ©. enknen sp. nov,
holotype Fe) wedemerida sp. now holviype CC prolate
sp. tov. holotype

medially. Seutellum scutiform, punctured, excavate.
Elytra punctate-striate, intervals convex, heavily
punctured; laterally angled out from base, rounded at
humeral callus, concave. rounded post-medially at
widest part to bispinose apex; small marginal spine,
sinall sutural spine, margin rounded and indented
belween spines, apices slightly diverging. Ventral
surface closely punctured, edges of abdominal
sezments glabrous, few short hairs, S,: males
truncate, indented medially; females unknown. Legs:
in male tarsomeres 1-3 without pulvilli, replaced by
median longitudinal ridge.
Sice

Males, 10.0 x 3.5 mm (2),
Aededazus (Fig, |)

Parameres angled outwards [rom basal piece.
rounded at apex. Pems blunt, sides acutely angled
away. Apophysis of basal piece medium width, tapered.
rounded apically,

Remarks

This species appears to be an oedemerid mimic as
its colour and pattern are similar to known oedemerid
Species; the model is unknown. It is not close to any
other known species.

Etymology

The name is derived from that of the beetle family

Oedemeridae.

Castiarina euknema sp. nov.
(FIGS IK, 2B)

Allotype: 9, 2 kin N Jabiluka, N.TL. 21111981, M,
Cappo. SAMA I 21286,

Furatype. WA.: 9, 34 kin 5S Roebuck, W.A,,
Bwii. 984. M. Powell, M, Golding, MPWA,

Colour

Head black wilh yellow reflections, elongate yellow
frontal spot. Antennae blue-green. Pronotum: anterior
and basal margins black with yellow reflections: yellow
laterally: medial black mark with yellow reflections
in the shape of sleeveless T-shirt. the neck enclosing
avery small yellow spot. Scutellum black with yellow
reflections. Elytra yellow with the following black
markings: narrow busal margin; pre-medial fascia with
ends expanded anteriorly over humeral callus reaching
anterior margin and enclosing yellow basal spot,
posteriorly reaching margin and enclosing yellow spot
on margin; post-medial fascia reaching margin and
Necting posterior extension of pre-medial fascia and
with iL enclosing yellow medial spot; mark covering
whole apex, clongate yellow mark reaching margin,
but nol suture, lying between this and post-medial

152 & HARKER

fuscia, In allutype black marks less prominent and
upial tnark encloses a small yellow apical spot., Ventrul
surlace yellow, edges of sutures black with green
reflections, 5,, 5. 8, with lateral spots coalesced with
marks along edges, Legs: femora and tibia blue;
larsomeres durk blue, Hairs silver,
Shupe and seulplire

Head shallowly punctured, shallow median sulcus,
medi Jength muzzle. Antennae compressed,
antennomeres |-4 obconic, S-Il toothed, Pronotum
Shullowly punctured, basal fovea extending forwards
to middle as glabrous line, basal notches on each side
closer to margin than middle; apical margin straight,
basil margin barely hisinuates laterally rounded from
base. widest piv-medially, rounded and natrowed tu
apes. Seutelleny seutiform, without punctures, excavate
along anterior margin, BLyWa pufctale-striate, intervals
flat anteriorly, convex apically; laterally angled oul
from base, rounded wt humeral callus, concave,

rounded post-mediully and narrowed to Irispinose

apex; marginal and suturul spines equal, medial spine
Slightly larger, margin rounded und indented between
spines, apices shightly diverging, Ventral surlace with
shallow punctures, edges of abdominal segments
wlubrous, few short hairs. 8, truncate both sexes.
Size

Male. 10.9 x 4.2 mn (1). Female, 1.9 x 425 mm (2).
Avdvuyus (Fig, WK)

Parameres parallel-sided from basal piece, rounded
medially then purallel-sided, rounded at apex. Penis
pointed, sides angled away. Apophysis of busal piece
medium width, lapered, rounded apically.

Remarks

This species has similar markings to those of C.
rifipes Macleay, except that the legs are blie whereas
they are ted in the other species, The male penttala
ure quite different (Pig. 1L),

Eryvinology
Derived from Gr. enknemes, beautiful legs.

Castiarina prolata sp. nov.
(FIGS IE, 2G)

Holotype, Of, Cardwell Ra, Qld. 23, 9,1993, J,
Husenpusch, SAMA 1 21287.

Alloipe: &. sume data as holotype, SAMA | 21288,
Pararypes: | or, | 9 same data as holotype, JHOA,

Colour

Head and antennae black with green and gold
reflections. Pronuotum black with green reflections ut
hase. Scutellum) black with blue reflections. Flytra
yellow with black inarkings coalesced forming ao
elongate yellow basal mark. a round yellow pre-medial
mark and a preduminantly red subeapical mark,

louching Margin not reaching suture and merginy ute
yellow closest to it, Ventral surface green with yellow
reflections, Legs; femora, dorsal surface deep blue,
ventral surlace green wilh old reflections; tibia and
tarsi dark blue. Hairs silver.

Shape and sculpiure

Head shallowly punetured.. broad median suleus,
shor muzzle, Antennae, antennomeres 1-3 obconic,

4-11 toothed. Pronotum. shallowly punctured, glabrous,

small basal fovea, larger fovea at each basal angle;
apical margin projecting slightly medially, basal margin
bisinuule; rounded from base, narrowed to apex.
Seutellum tulipiform, few punctures, glabrous. fat.
Elytra anteriorly punctate-striate, intervals raised
postenorly, intervals convex; laterally parallel-sided
at base, rounded at humeral callus, rounded and tapered
posteriorly, then attenuated, rounded at apex (o pointed
marginal spine, deeply indented and rounded to minute
sharp sutural spine. apiwes diverging. Ventral surface
with shallow punctures, edges of abdontinal segments
glabrous, Sparse medium length hairs. Mesosternal
process inflated, Elytra. cuntlevered oyer last visible
abdominal sternite, Legs: femora and tibia long and
thin: tarsomeres with enlarged pulvilli. $5: truncate
both sexes.

Sine

Males, 10-9 x 3.4 mm (2). Females, 12.4 x 3.6.mm
(2)

Aedeugus (Fig. 1E)

Purameres elongate, angled outwards front basal
piece, apically rounded, Penis pointed, sides obtusely
angled away, Apophysis of basal medium width,
elongate. tapered. rounded apically.

Remarks

This species 6 a 'C. produvta’ group mume. lt
resembles C. acuminata Saunders which also has
urdatly attenuated apical spines but is heavily costate
Che apical spines of C prelata are both obvious
wherews in C. geuninara the marginal spine is absent.
Male venilalia differ (Pig 1B)-

Erymalogy
Derived foun L. predatus, elongate.

Castiarina ernestadamsi sp. nov
(PIGS (1, 2A)
Halatype: oO. Mourangee. Edungalba, Qld,
26.41.1979, E. E. Adams, SAMA T 21289.

Pararypes: Qld: 3 o, Mourangee. Edungalba. 8. A
Adanis & BF. b Adan, SAMA.
Colour

Heud maroon at base blending into blue-green, blue
muzzle. Antennac, antennomeres [-2 blue-green. 3-1

BIGHT NEW SPECIES OF AUSTRALIAN BUPRESTIDAE, In

bronve green. Pranotum maroon, Scutellum blue or
blug-gecen, Elyira pale yellow with lollawmy dark blue
nuirkings: darrow basal margin; small mark over cach
humeral callus; remnant post-medial fascia touching
jatcral margin reaching hall way to suture, small
remnant murk ofsuture, small apical mark. Ventral
surhice: Slernum muroon, abdarien teskiceous, lees
blue. Nias silver

Shapy and sculplure

Head shallowly punglured, medium sulcus, short
mazele. Anteunac, antennomeres LF obeonic, 4-11
toothed. Pronotum shallowly punctured, basal foves
extending forwards lo middle us globrous line, basal
notetes represented by glabrous area on each side
closer to mhargin (han quddle; apical margin projecung
mediilly, basal margin barely bisinuate: Taterally
parillel-sided at base. angled outwards and rounded
ta widest pre medially, rounded and narrowed to apes,
Sculetlum scutiform, glabrous, flat, Flytra punctate-
striate. intervals convex. lightly punctured; laterally
angled outwards from buase, rounded at humeral cullus,
voneave, rounded post-medially and narrowed to
bispinose apex; both spines small and sharp. margin
rounded and indented between spines, apices hardly
Jiverging, apical margin subserfate, Ventral surface
with shallow punctures, edges of abdormmal seetnents
glabrems, stermum with few medium length hairs,
abdomen almost hajrless. S,: males truncate, female
unknown,

Site
Males, 24 + 0.09 x 4.8 + 0.04 mm (4),

Aedeagus (Fig. 1)

Lightly tanned. Purameres parallel-sided [rom busi!
piece. pre-medially rounded, parallel-sided then
rounded lo apex. Penis pointed, sides obtusely angled
away. Apophysis of basal piece narrow, rounded
ayneatly

Renuirks

This species was previously confused with ©.
siraiminea MacLeay. However it ig smaller, it has i
smooth pronotum with small punctures whereas C
straminea bas heavy punctuation. I has darker elytral
markinys than C. straniinea and the male genitalia
differ (Fig. Wi. A form of C. strigara MacLeay (Pig.
2B) occurs mm the type locality, sinnilar in size and
pattern but not in colour tr has very light elyeral
inarkings, the post-medial fiscia being considerably
reduced. The head and pronotum are green with
reddish reflections in some specimens:

Ltymolouy

Castiarina oclopunectata sp. nov.
(FLGS IF, 2D)

Holonpe: o 9 km NNE Zuothus, WA. 21x 1986,
M. Powell, WAMA

Allorpe: G. Wialka, W.A., 2L.1%.1970, S. Barker,
SAMA J 21290.

Paratypes: W.A.. | co SAMA: | 9. summit Mt
Cooke. 10.%7,1956, S. Barker, SAMA, 1 9. Wialkiv
1k.ix.1957, 8. Barker, SAMA; 2 cor, Kalbarri N.P.-
23.4x.)969, FH, Uther Baker, SAMA, 4 orot, 2 9 @,
Beverly T. 0.. Brookton Hwy, 19,ix.1970, 8, Barker.
SAMA; 2 Oc, sume data as allmype, SAMA, 1,
6 kin S.Tammin, § x1.1970, §. Barker. SAMA: 3 oo,
3.0 9, 1/2 way between Glen Eagles and Brooktou
Hway. 13.x 1980. 8, Barker. SAMA: 1 9, & kin 6
Woolgangie. 22.%.1980, S, Barker, P. G. Kempster,
SAMA: | oy. 10 kan E Merredin. 12.4, 1990, §, Barker.
SAMA? | ot, 4 km W Zanthus, 2!.4.19%6, M- Powell,
MPWA, I of, same data as holotype, MPWA; 1 4).
Moora, WAMA: | ©, MeDermd Rock
27.1%/3.%.)978. 1. F Houston, WAMA; 2 oor, 1 @.
Lake Ningham, WAMA; | 9, Merredin. WAMA; |
o, Karlgarin, W. Duboulay, WAMA: | 9, Dedari,
7.x.1978, T. M.S. Hanlon. WAMA; I @. 21 km W
York. 4.xi.1978, T, M.S. Hanlon, WAMA; 1 oF, 1.5
kin S Mt Jackson, 5/11.34.1979, TF. Houston ef al. ,
WAMA; |oc, | 9, 12 km NNE Bungalbin Hill,
H/tB.ax1979, T. B Holston eral, WAMA: 1 oo, |
@, 14km NNE Bungalbin Hill. WiS in 979, T. F,
Lluuston ec: al. WAMA: 3 oo, | oD, 18 kin NNE
Bungalbin Hill, W/18.1%.1979. Tl) Fo Houston ef al),
WAMA; 2 © Q, Dedari, 21.in.1979. 1. M.S, Hunton
WAMA; | c. Mt Dale, 20x, 1980, 'T. M.S. Hanlon.
MHSA; | o, Muckinbuddin, 10,x,1978, RB. P
MeMillan. WAMA: | ot, Mt Walker. 34 kin b
Narembeen, 23.«, 1980, R. PB. MeMillan. WAMA: 3
co. | @, Mt Observation, 2bx.1987. Ro P
McMillan, WAMA: | 9. Beacon, 20.%.1981, R. PL
MeMillan, WAMA; L Q, 20.1%, 1990, Bonnie Rock,
S. Barker. SAMA: | o&, 10 km E Mervedin, 12, 7.1990,
S. Barker, SAMA: 3 cr ol 2 9 @. Dedari, 22.«.1991,
TM. S. Hanlon. MHSA.

Colour

Head. antennae. pronotum dark coppery-purple.
Seutellum dark blue with coppery-purple teflections.
Elytra yellow with coalesced dark blue markings with
coppery-blue reflections forming the following yellow
marks: 4 large medial spots in a row on each elytron,
the basal, pre-medial and post-medial roundish, the
pre-upical clongate: narrow margin from base. not
reaching apex. Ventral surface and legs brown with
coppery-purple reflections. Hairs silver.
Shape and sculpture

Head closely punctured, broad median sulcus, short
muzzle. Antennae, antennomeres 1-3 obconic, 4-Il

ist S BARKER

louthed. Pronofum) closely punctured, basal levea
estending, forwards to middle as glubrous impressed
Hiessupleal margin projceting broadly medially, basal
Htrgin almost straight, laterally parallelsided al base,
slightly rounded, widest medially. slightly rounded to
apes, Sculellum scutiform. wabrous, excavate. elytra
puneRde-striate, intervals Convex, smooth medially,
punclured and rough laterally: luterally angled out from
base. rounded at humeral callus, concave, rounded
Post-medialhy ond arrowed t spineless apex, last
iMerval indented and straight, apices slightly diverging,
Eniire ventral surface covered in dense. flat hairs, alse
present uround Jaterul margins of pronotum and in
SOME specimens cnervaching onto dorsal surtiwe, §,;
Males truncites females rounded und slightly pointed,
Size

Miles U9 + 016% 4.0 +4 0.06 min (35), Females,
12,7 = Ut x 4.2 + 0.07 mm (25)

Adedeagus (Pig. UF)

Parumeres angled oulwards and gradually widened
from basal piece, rounded at apex. Penis pointed.
angled away ebtusely, Apophysis of basal piece
medium width, rounded apically. Proctiger with apex
bluntly bilobed, the two projections variable (Fig, IM).

Female terminatia (Fig, IN)
Proctige?; apex with Iwo narrow, pointed lobes.

Remarks

A species complex exists in Western Australia whieh
includes C) parallela (White) with a more or less
continuous gradation in size from the smallest (C
parallel) to larger species. C. paratlela also occurs
mousiorn Australia, Until now T have not been able
jo separate the individdal species in the western
conmiplex, ©) vawihepilosa Hope and C yin Saunders
dre closely related species but only oecur in castern
Australia. From examination of the terminal abdominal

SOHNE, it now appears as if males and females of

Hoth castern and western specimens of C) parallels
have a rounded proctiger and can be distinguished on
(hal basis and on (heir colour size and structure of male
genitalia, Both sexes of the larger western species huve
ao ormamented proctiger CC. vetpunetata 1%
distinguishable on the basis of the structure of male
genitalia and the lack of apical spines on the elytra,
Further work ts required to delimit the remaining
species
Ervnnligy!

The name is derived from b eeto. enht und 1,
PuUHCTATO spontea,

Themognatha viridescens sp. nov,
(FIGS 1A, 3)

Holotype. , Tron Ran,
SAMA I 2129),

Old, 20.vi 1980. Wood.

Allotype: 2. Tron Ra., Qld, 30.1.1966. J, Kerr, ANIC,

Fararypes: Qld: 29 @, bran Ra., 3/9.V.1966, J, Kert,
ANIC, 1 a. Iron Ra. 24.v,1974, M. Walford Huggins.
MHSA; | &, tron Ra., Qld, J9v1978. G. Wood,
GWOA; [sex indeterm. 5.1966. J. Macqueen. J. Keer,
ANIC.

Colour

Head, antennae. pronotum and scutellumn bright
green with yellow reflections. Elytra: yellow with
following dark green markings: narrow basal margin;
medial fascta not reaching margin, apieitl mark
connected along sulure to fasent. Ventral surface muinty
bright green with yellow refleetions, male with
tesiaceous patches on S, and S., abdomen all green
in female; legs bright green with yellow reflections.
Shape and sculprare

Head punenition fine, even, dense, median impressed
basal line, trons moderately hairy; labrum
longitudinally divided and pointed. Pronotum narrower
than elytra, LeW 0.6, punetation fine moderately dense;
sides rounded from base to 1/3 distance to upex, then
explanate and converging anteriorly, laterally flattened;
untenior margin bisinuate, posterior margin almost
straight median glabrous line from base to near apex
Scutellum scutiform, anterior margin straight, concave
without punctures, 1/8 width of elytra. Elytra slightly
wider than thorax; elytral interneyrs long and with
scutellary striole, strongly marked with heavy
punctition and additional punctation on shoulders:
intervals flit, sides sub-parallel then tapering to pre-
Apical areas; apex bisinuate, both spines prominent,
interval hetween sinuous, lateral spine anterior 1
medial spine, Ventral sulface: prostemum hairy, finely
punctured, with a defimte forward medial projection;
pro-episternum finely punctured with deep smooth
fossa in posterior angle; mesusternum und meta-
sternum smooth medially with coarser punctation
laterally; hairy, hair long and fine medially, shorter

Mv
;

inal

‘
f
omm

Fig. 3. Habitus Hustation af Themownatha varidesceny sp.
nov. Holotype

EIGHT NEW SPRCIES OF AUSTRALIAN BUPRES TIDAL 155

and coarser laterally: abdomen smooth and shiny in
nine, punctation very fine, apieally huiry in female,
S.: male deeply concave, fernale rounded. Feet: tarsal
claws without a notch,

Size

Male, 31.4 x U4 mm (2). Ferales, 30.7 « ILS mm
(4),

Acdeauus (Fig. 1A)

Parameres parallel-sided trom basal piece, ungled
outwards premedially, rounded al apex. Penis pointed,
sides acutely angled away, Apoptiysis of basal piece
medium width, tapered, rounded at apex.

Remarks

This is un unusuul species as ft shows two characters
found in the related genus Caledema C&G, I hits a
small medial projection on the anterior margin of (he
prosterum. but not as large as those found in, Cule-
dema. The seutellum ts wider thar in obher Thenig-
athe in relauon to the elytral width, although not as
wide as any of the known Calodema species, However,
the body is not sinuous in lateral profile, the pronotum
is nous wide as the elytra and the elytral interneurs
ure cleurly defined as in Themognadie. The tarsal claws
ure not notched, bul this is a variable character found
in some Themognatha and not in others,
Etymology

Derived from L. viridis, green.

Astraeus (Depallus) powell sp. ny,
(FIGS. 1G, 4B)

Holotype: , Quaitading, 7,X1,1970. S, Barker, SAMA
I 21292,

Paratypes: WA: 1 or, Tammiin, §.xi1970, S, Barker,
SAMA; | co, 17 km E Dowerin, 21.4.1989. M.
Golding, M. Powell, MPWA; 1 cr, 6 km SE Tanunin,
15.xii.1990, M. Golding, M, Powell, MPWA, | 0”, 43
kin E Merredin, 26.8199], M. Golding, M. Powell,
MPWA;: 2 oct, 32 km EF Yellowdine, 21/22,x.1991,
T. M.S, Hanlon; 2 9 9, Quairading. 27x11). 1991, K

K.. MPWA: 4 & 9. Quairading, 11.1992, M.
Golding. K, K.. MPWA.
Colour

Head. antennae. pronolum black with blue and
purple reflections. Elytra black with following yellow
murkings: irregular marks ulong width of one strite,
roainly concentrated along 4th and 8th intervals from
sulure and along the murgin on basal half. Ventral
surface and legs purple: lateral yellow spots on S,,'5,,
S$, in MUS! Specimens last one absent in holotype.
Hairs silver.

Shape and xculptnre

Head punctures small medially, larger hiterally, small

glabrous median keel near apex merging into impressed

hig, 4. Habitus illustrations of the following Asiraens species
A.A, dbberans \ le Poll. B. 4, panel/esp, noy, holotype

line basally, Pronotum punctures smaller medially,
larger and in form of shallow fovea laterally, glabrous
median urea; anterior maunsin projecting medially, basal
murgin strongly bistnuate, laterally fhajry. Elytra
intervals between striae Convex and smooth, [aterully
parallel-sided from base, rounded post-medially und
narrowed to apex, small pre-apiecal notch. on margin.
hroad outwardly curving satural spine, hairy overall,
Ventral surface and Jegs punctured und Hairy.

Size

Males, 12.6 + 0.22%44 + 0.11 min (7). Females,
15.3 + 0.38 x 5.6 4 (12 mm (7).

Aedeagus (Fig. 1G)

Parameres parallel-sided from basal piece, gradually
widening until rounded lopointed upex, Apophysis ol
basal piece medium width, rounded apteally.
Remarks

This species has been contased With 4a, uberranys \
de Poll (Fig. 4A). Ut differs fron) that species by being
narrower, haying most of the yellow elytral marks
medial and post-medial whereas ind. aberrears they
are more evenly distributed, by having single
outeurving sutural spines on the elytra while im A.
vberrans the sutural spines dre small and there is a
definite small marginal spine,

Envnolowy

In honour of Mr M. Powell of Melville, W.A. who
has assisted my research for many years.

With the addition of the above new speutes the key
to Astraeus (Depollas) (Barker W978, p.j07) requires
the following replacement:

Replace 4. uberrans van de Poll with;

4a, small, ulmost straight marginal spine

Sibert eye pees ey yey ye tberrans Vary de Poll
4b. broad, oulcurving marginal spine

Meg OR me eT ..pawelli Barker
With the addition uf this species the sub-genus Depalliis
now contains nine species,

156 S. BARKER

Acknowledgments

I thank the following for their assistance: Dr T. F.
Houston, WAMA; Mr T. A. Weir, ANIC; Mr M.
Cappo, AIMS; Ms H. Vanderwoude, Department of

Zoology, University of Adelaide. I thank the following
collectors for the loan of specimens: Mr E. E. Adams,
Edungalba; Mr T. M. F. Hanlon, Hunters Hill; Mr
J. Hasenpusch, Innisfail; Mr M. Powell, Melville; Mr
G. Wood, Atherton.

References

ARNETT, R. H. Jr, SAMUELSON, G. A. & NisHtpA, G. M.
(1993) ‘The Insect and Spider collections of the world’ 2nd
ed. (Sandhill Crane Press, Gainsville).

BarRKER, S. (1975) Revision of the genus Astraeus LaPorte
& Gory (Coleoptera: Buprestidae), Trans. R. Soc. S. Aust.

99, 105-142.
(1987) Eighteen new species of Stigmodera
(Castiarina) (Coleoptera: Buprestidae). Ibid 11, 133-146.
Watt, J. C. (1979) Abbreviations for entomological
collections, N.Z. Zool. 6, 519-520.

A NEW SPECIES OF CALLULOPS FROM NEW GUINEA AND
COMMENTS ON THE STATUS OF C. HUMICOLA COMPTUS
(ZWEIFEL) (ANURA: MICROHYLIDAE: ASTEROPHRYINAE)

By STEPHEN J. RICHARDS*, THOMAS C. BURTONT, MICHAEL J.
CUNNINGHAME & ANDREW J. DENNIS*

Summary

Richards, S. J., Burton, T. C., Cunningham, M. J. & Dennis, A. J. (1995) A new
species of Callulops from New Guinea and comments on the status of C. humicola
comptus (Zweifel) (Anura: Microhylidae: Asterophryinae). Trans. R. Soc. S. Aust.
119(4), 157-162, 30 November, 1995.

Callulops sagittatus sp. nov. from the summit of Mt. Binnie, Western Province, Papua
New Guinea is described. It is a moderately large species (males 44.1-47.8 mm, a
female 56.3 mm S-YV) distinguished from congeners by the presence of an orange
stripe from the tip of the snout dorsally across each eyelid, forming an arrow-shaped
mark on the crown. The advertisement call is a series of 11-12 deep croaks uttered
from the entrance to, or deep within, crevices between rocks. The female paratype
contains large (4.5 mm diameter) unpigmented eggs indicating that, like other
Australopapuan microhylids, larval development is completed with the egg capsule.
Among the Asterophryinae, Callulops sagittatus and C. h, humicola share a unique
condition of the mandibular branch of the trigeminal nerve suggesting that C.
sagittatus and not C. h. comptus is the closest relative of C. h, humicola. This and a
number of other consistent morphological differences indicate that C. h. comptus
warrants elevation to specific status.

Key Words: Anura, Microhylidae, Asterophryinae, frog, new species, Callulops
sagittatus sp. nov., Callulops humicola, Callulops comptus, New Guinea.

Ieansactiuns of the Roval Sovivey of 8. Aust. (1995~ 194) [STAB

A NEW SPECIES OF CALLULOPS FROM NEW GUINEA AND COMMENTS ON ‘THE

STATUS OF G HUMICOLA COMPTUS (ZWEIFEL) (ANURA: MICROHYLIDAE:
ASTEROPHRYINAE)

by STEPHEN [. RICHARDS* THOMAS C. BURTON]. MICHAEL J, CUNNINGHAM
& ANDREW J.. DENNIS*

Summary
RICHARDS, SoJ.. Burron.T, C., CuNsiGiam. M. J & Onis. A, J. (995) Anew speeres oF Calladaps tem
New Guinga and comments-on the stars of C. fnivote complus (Aweilel) (Anurat Microhytidue: Asterophiry inte).
Trans. R. Soe S, Aust O94) 1587162, 30 November 1995

Callalaps sagivatas sp. poy. fron) the sununit pF Mi Binnie, Western Provinee, Pupusa New Guinea iydescribed.
Iisa moderately large specios (males 44-478 Wing a temale 56.4 mm S-V) distinguished Mont eongeners hy
the presence of an orange stipe from the tp of the snout dorsally aeroxs each eyelid, forming ari arroweshaped
mark onthe crown, The advertisement call is a series af 1-12 deep croaks uttered from the enitianee to, or deep
within, crevices beiween rocks, ‘The female paratype contains large (4.5 mm diameter) Unpigmented eggs mdicating
that Tike other Australopapuan microhylids, larval development is completed within the egg capsule, Among
the Asterophry imac Cedlalops sawitatas and ©.) huwnicola-share a unique condition of Ihe mand ibulur branch
of the trigeminal nerve Suggesting that ©. sagi(aias and not © Je Compras isthe elosest relulive of Coy diarealia
This and a number of other consistent morpholoieul differences tulieate That CL A. errngrtiey Warrants elevation
tO Specific stitus,

Key Worps: Amira, Microhylidac, Astérophryimac. fray, new species, Cilludops sequiniotas sp now, Ceaflatepes

humicola, Calluleps comptis, New Guinea.

Introduction

Microhylid trogs of the subfamily Asterophryinac
ure restricted to the New Guinea. miumtiland and nearby
islands (Zweilel & Tyler 1982). This ecologivgally und
morphologically diverse graup occurs from sea level
to subalpine meadows hyzh in the central cordillera
(Zweite! 1972), Ln a review of the Asterophryinae
Burton (1986) recognised eight penera and 43 species.
Blum & Menzies (1988) subsequently deseribed nine
new species of Xenebairachus and Xenorhina, and
Richards ef a/, (994) described a new species of
Axteroplrys, bringing the total to 53. Additional
undescribed species. uccur in useui collections, and
field work continues to reveal Unnamed taxa.

Durning a survey of the fauna Of Mt Binnie, Western
Provinec, Papua New Guinea (Dennis ef al 1995)!
three of us (SR, AD, MC) collected an undeseribed
species of the asteruphryine genus Callulops. The
discovery of this new species peeessitates a
reassessment of the relationships of the taxa currently
recognised as subspecies of Callulops humicola, Here

“ Zoology Department, James Cook University, Lownsville,
Qld 481.

{ Division of Biological and Chemical Scienves, La Trobe
University, Bendigo, PO Box 199, Bendigo. Vie. 3550.

+ Zoology Deparment. University of Queensland, Qld 4072.

1 Dennis, A., RICHARDS. S. & CUNNINGTIAM. M. (1995)
Preliminary survey of nyarimals, birds, reptiles and frogs
on the summit of Mt Binnie, Western Province, PNG, 20-23
November, 194. Report to Ok Tedi Mining Limited
(unpubl, )

we deseribe the new species wad demonstrate: thal
Callnlops lunmicola camptus warrants elevation i
specific status.

Materials and Methods

Specumiens are deposited in the Biology Departinent.
Universily of Papua New Cjuinea, Port Moresby:
(UPNG) and the Queensland Museum, Brishane
(QM).

Recordings of mating calls were made in the field
with a Sony Professional Walkman tape recorder with

aw Eleetret Condenser Microphone ECM-Z200 und

were analysed using the sound dnalysis progrun
~Canary” (Cornell Grnithology Laboratory, 1994).
Measurements were made to the pearest 0.05 mm
with cial callipers or to the nearest 1 mm using a
binocular oneroscope with aq ocular muicromeler,
Methods of measurement follow Zwejfel (1985) except
the snout-naris measurement, laken from the tip of the
snoul to the cenire of the nurs, Measurements. (min)
were: snoul-vent length (SV); tibia length (TL); eye
diameter (EYER): eve-naris distance (EN); internarial
distance (IN); sneut-naris distance (SN): head width
atungle of the jaws (HW): head length from tip of snout
to angle of the juws (HL); horizontal diameter of
tympanum (EAR), hand Ieogth (HD); foot length (PT).
The rather featureless palmar and plantar surfaces of
the bands und feet. and the poorly defined tympanic
annulus made measurement of the hands, feet and
tympanum difficull, and the measurements should be
trealed with caution, Ope of us (TCB) dissected the
superficial throat and jaw musculature under a Wild
M3Z microscope with the aid of topical application

13% *& J RICHARDS, T. C, BURTON, M J CUINNINGITAM & Ad, WINES

of the jodime-potassium iodide solution of Bock &
Shear (1972), Phe squamosal bone was also examined,

Systematics

Callilops Boulenger \s applied Ww asterophryine
microhylid frogs formerly referred to Phryiwmeietiy
Peters, following the recommendation of Dubois
(L988). Callulops is distinguished by two skull
characters: two supplementary slips lo the M,
intermandibularis arise from the dentary: one via a
tendon and the other directly, and run together. more
or less parallel to the mandible, to insert upon the
ventral fascia of the M, submentalis and semetiines
also upon the adjacent medial aponeurosis of the M
intermandibularis (Burton 1986), The second character
is (hat the otic ramus of the squamosal bone is about
the same length as the zyomutiv ramus, and itis not
twisted 1.c. the postero-lateral surface of the otic ramus
rs.continuous with the lateral Surface of the zygomatic
ramus (Burton 1986).

Tr his revision of the asterophryines. Zweitel (1972)
described Callulops li. humicola and Ch. compres.
These taxa resemble each other superticially apart from
relatively longer legs in Ch. fumicela, and an orange
postocular stripe in adult C. A. compius which is only
“somewhat developed in young humicola” (Zwerlel
1972 p. 476). The geographic ranges of these taxa abut.
Zweite) reported only one instance of symputry, and
was reluctant to assign the taxa fo species status in the
absence of evidence of reproductive isolation.

Burton (1986) added two further characters (o
distinguish the taxa. First. in Cf Aumieala the M
depressor mandibulae arises from the dorsal fascia,
with some fibres from the otic ramus of the squamosil
and the posterior surface of the adjacent prootie, in
Ch. comptas and all other Calluleps, additional fibres

arise from the entire posterior and ventral surfaces of

the tympanic ring. Second, in CA. Juomivola the
mundibular branch of the trigeringl berve passes
directly ventro-laterally through the M, adductor
mandibulae posterior longus on its way tw the

THandibWear musculature; in CA, comprus and all oiher

asterophryines this nerve passes antero-laterally
between the M. a. m. posterior longus and the M.m
anterior longus, and then postero-ventrilly aeross the
lateral Surface of the M. a. i, posterior longus belore
plunging towards the mandibular musculature, Burton
(1986) made no taxonomic recommendation regardiny
the status of these taxa.

Callulops sagittatus sp. nov.
(FIGS 1-5)

Holotype: UPNG 90ST an adult male collected by 8,
J, Richards, M. Cunningham and A, Dennis on

Albay 1994 at an altitude of 2200 mon the summit of
Mt Binmte. Western Province, Papua New Guinea 4b"
7 MY"E, S° 12'S).
Fararypes, UPNG 9052 (adult female), OMI 6231
(udule indie), sume daig as holotype
Definition

A muderely large and robust species (males
44,147.86 itn SY, a fermale 56.3 mm S-V) distinguished
froyn congeners hy a combinalion of the following
charuclers; fingers and toes witout expanded dives.
4 dISTINeT ohinge stripe dorsally wn the head from the
snol extending deross cach eyelid, lympanurn
Indistinel, adverisenient call a senes of deep “crawks"
wilhatslominiml frequency of 609 He, a pote reperitiin
rate of LRS-2.2/s and a pulse rate of 189-14 02/ms

Fig, 1 Crtudaps sagiindnas ap now. in life (SV 569 pi)

Deycriplion of halornpe

Body robust, almost pear shaped (Mig. 1), head
brouder than long (MW/HL 1-23) with nares-closer to
tip of snout than to eye (SN/EN 0.6) and directed
Jaierally, Internarial distance greater than distanee from
eye to nares (EN/IN 0.78), eyes large (EY E/S-V 0.119).
Snout blunt, hrowlly rounded in dorsal wiew and
rounded if: lateral view Cig, 2), Cunttius rostralis
ruunded, lorcul region steep, slightly coneave.
Tympunum indistiner. wnnulus barely visible. Bursal
and ventral surfives tminutely gtanular. a weak
supralympamne fold. Anterior palutal ridge long,
smooth, posterior palatal ridge with 1) digtinet
dennicles.

Limbs short (TL/S-V 0.38), relative lengihs of
fingers 3>4>2>1), fourth finger only marginally
longer (han second, Fingers unwebbed, tips without
expunded discs, subunicular tubercles low, rounded,
Palm smooth exeepl for a low toner metacarpal
oiberele, Relutive lengths of mes 4>3>5>2> |. Toes

A NEW MICROHYLID FROG FROM NEW GUINEA 159

unwebbed, lips without expanded discs, subarticular
lubercles low, rounded, A low, oval inner metatarsal
tubercle; no outer tubercle (Fig. 3),

Colour in hfe uniform deep red-brown dorsally on
body and limbs, grading laterally into a uniform lighter
brown ventral surface. Throat slightly darker brown
than rest of venter. Head deep red-brown with an
orange stripe dorsally from lip of snoul along eanthus
and over eye, forming distinet arrow shape on crown.
Slight orange tinge on upper surface of thigh. No other
inarkings dorsally or ventrally, In preservative brown
with a mauve Hinge dorsally, brown ventrally, stripes
on head very pale pink.

Fig, 2. Views of head of Callulops sagittanus sp. noy. holotype
(UPNG 9051), A, Dorsal view, B. Lateral view, Seale bir
10 mm,

Fig. 3. Hand and loot of Callulipy saginarss sp. nov. holotype
(UPNG 9051), A, Plantar view of foot. B. Palmar view
ol fand. Seale bar = 10 mm,

Dimensions of holotype

S-V 47.8; TL 18.2: EN 3.3; SN 2.0; IN 4.2: BYE
5.7: HW 17,0; HL. 13.8; HD 12.5: PT 19.3; EN/IN
0.785; TL/S-V 0.38; HW/HL 1.23, EYE/S-V 0.119;
HW/S-V 0.355; width of toe tip on fourth loe 1.0 (width
of penultimate phalanx 0.8), width of toe tip on third
finger O.9 (1.0).

Musculature

The superficial throat musculature and squamosal
form conform to the definition of Callulops. The M.
depressor mandibulae arises predominantly from the
dorsal fascia, but also reccives substantial contributions
from the otic ramus und the posterior and ventral
margins of the tympanic ring. The mandibular branch
of N. trigeminalis passes directly yentro-laterally from
the brain. case and penetrates the M, adductor
mandibulae posterior longus on its way to the
mandibular musculature (Fig. 4).

Advertisement call

We recorded two call sequences butonly one of these
is of sufficient quality for detailed analysis. The mating
call is a series of deep, guttural croaks “erawk, crawk,
crawk.... The recordings contained Il and 12 notes
lasting a total of 5.57 and 5.186 seconds respectively
(note repetition rate = 1.85/s and 2.2/s). Both calls
had a dominant frequency of 609Hz. Individual notes
in the Il-note call lasted 118.9-182.3 ms (mean =
154.2), contained 10-13 pulses (mean = 11.63) at a rate
of 11.89-14.02/ms (mean = 13,23). Fig. 5 illustrates
the first four notes of an H-note call recorded at the
type locality on 20.xi-94 at an air temperature ol}
13.35°C.,

1a)

S.J RICHARDS, T, C. BURTON, M

Fiy 4 Dotso-fateral view of musculature of right jaw of
Culluleps wagicanis sp, nov, A.M. adduct imandibuliue
anteriog Tongus: D. M-_ depressor mandibulde; PM
ahluctir mandibulae posterior longus: 8. M. adducror
niandibulie extemus superticialis (severed); Vo mandibular
branch of trveminal nerve. Scale bar = 5 mm.

The calls were uttered at ipregular intervals, willt long
periods (often over five minutes) between calls. On
several occasions we heard 3 melodious, dove-like “coo
cow..." vocalisalion utlered immediately following, or
from the same vicinity as. one of the call types
deserihed ubove but we were unable to confirm
whether tL was part of the vocal repertoire of this
Species,

Natural history

The type series was collected in disturbed rainforest
and secontary regrowth at altitudes over 2000 m alung
jhe access road to the summit of Mt Binnic. One male
was calling from the entrance to.a deep crevice between
rocks in a vertical road cutting, and the other was
culling (ror deep within a labyrinth of crevices among
large rocks in the road cutting, The female was
collected on the surface and appeared to be approaching
the latter male. Additional males were heard calling
within crevices adjacent to the road but we were unable
to trace them due to the Sporadic nature of calling and
their subterrancan habitats.

The female contains large, Unpigmented egps
indicating that development occurs within the egg
capsule like other Australopapuan microhylids (Zweitel
& Tyler 1982), Two mature eggs measured in the ovary
were 4.5 mm in diameter

| CUNNINGITAM & A. 4. DENNIS

The skin of this species ys thick and glandular (Fig.
4) and the animals exuded a slimy mucus wher
handled, apparemly as a defensive mechanism

hariatian

The colour pattern is Consistent im the three available
specimens, all of which exhibit the distinetiye oranpe
stripes on the head. Some of the fingers and toes of
each specimen aré dehydrated making accurate
measurement difficult, but none has expanded dises:
some fingers and loes have faint, vestigial grooves an
the Ups, Measurements and propertions of tHe teu
paratypes (PNG 9052/OMJ60231) are: S-V 56,3/44 |;
TL 1:8/14.6, EN 3.4/3.3; SN 2.1/2.0; IN 4.6/3.9: EVE
6.0/5.2; AW 19,0/17-7: HL (6.0/6.1: FT 2.0480. HD
13.2/11.7: EN/TN 0.76/0.846; TL/SV 0.35/0.33; HW/HL
118/109) BY E/S-V 0106/0, 018; HW/S-V 0 337/040);
width of toe tip on fourth toe (width of penultimate
phalanx) 1.1 (0.99/08 (0.7). width of toe tip om thin
finger $0 (1.00/10 (10),

Comparison with ather species

Callulopy (sensu Dubois (988)) now includes IS
species, Morphologically they are rather conservative,
und there is extensive overlap in most body proportions
among species (Zweifel (972 Table 6), The
comparisons belaw are based lurgely on the detailed
descriptions of taxa presented by Zwetfel (1972)

The presence of orange stripes dorsally on the heal
distinguishes C sagiftaney Jrom knowh congeners.
Callulaps boetteer{?, C. eursdactylus and C slateri
further differ from C. sagitiatus in having greatly
expanded linger and toe discs (vs no dises). Callulops
doriae, © dubius, C fusens, Co humicola nomicoli
and Ch. compins, C. kopsteini, C. persenatus and
€. robustas have smal) grooved disus on the fingers
and toes, Callulupy derive (00 mm), Co perxename
(72,5 mm) and €. rabuses (73 mm) are much larver
species and C. dubius appears to be 4 much smaller
species (maximum S-V = 24 mim). Although sample
size is small, there appear to be differences between
the mating calls of C saginartus and those of both C.
persenarys.and C. robustus, Two calls of C, personas
have a dominant frequency of about 1000-4500 Hz (vs
609 Hz) and contained 5 notes (vs I-12), Calls of C.
rohastus from Misirna Islnd (the type locality) have
a dominant frequency of about 8O0H? (J, Menzies
unpubl, data). Callulops h. compiuy is the only other
Species in which adults have orange stipes on the head.
but in this species the orange markings are restricted
toa Short litera post-ocular stripe, Callulops sagittaine
shares with C. ft. Awmicsla the condition of the
mandibular branch of the trigeminal nerve. As this
condition is unique umong the asterophiryines 1 appears
to be a synupomorphy indicating u close phylogenetic
relationship between these taxa. However, ©. h,
Aumicila differs (rom C. saeinatis in the possession

A NEW MICROHYLID FROG FROM NEW GUINEA tol

2000
HZ
1000

-1000

Time (ms)

Fig. 5. Audiospectrogram (top) and wave form (bottom) of first four netes of an TH-note call sequence of Callulaps sagutaus
sp nov recorded it the type loealny. Air temperature 13,5°C.

of grooved dises on the fingers, and lack of orange
stripes on the head. Three species, namely ©,
glundulosus, C. stictogaster and C. wilhelmanus share
with C. sagifiatus the lack of finger and toc dises,
Callulapy vlandulosus. differs from Co sagittatus im
having a coarsely mottled ventral surface (vs uniform)
and in having a well-developed glandular area behind
the car, whereas C. stictogaster is a larger species (lo
80 mm) and has a distinet tubercle between the eye
und the nostril (lacking in C. sagitrarus), Callulaps
wilhelmanuys closely resembles C. sagiftalus and has
a similar call (J. Menzies unpubl, data), size and colour
pattern. It is distinguished predominamly by the
absence of orange markings on the crown and the
condition of the mandibular branch of the trigeminal
nerve.

Zweilel (1972) discussed four specimens of Callulops
trom Busilmin on the northern slopes of the Star
Mountains that he tentatively assigned to C. robusius.
This population is geographically close to the type
locality and is ata similar altitude. The frogs are withm
the size range of C. sagittalus but none exhibits the
orange stripes typical of this species and their
identification remains uncertain,

Srarus of Callulops h. humicola and C. h. compitus

Although we still lack calls or other reproductive
data for these taxa, their classification as a single
species is no longer tenuble, given the evidence that
the axon most closely related lo C. fr, Auimicela is not
Ch. comptus but C. sewitarus, In hight of this and

previously reported consistent morphological
differences (Burton 1986; Zweifel 1972) we propose
that C. humicola compris be elevated to specific status
as Callulops compius (Zweitel) new combination.
Zweilel (1972) presented a thorough description of these
two taxa and a detailed comparison with each other
and all other Callulops except C. seagitianes, with which
they are compared above.

Erymology

From the L. Sagitta (= arrow) with reference to the
arfow-shaped orange markings on the crown.

Acknowledgments

Field work in Papua New Guinea by SR, MC & AD
was generously supported by Ok Tedi Mining Limited.
We are particularly grateful to lan Wood, Andrew
Storey and Jan Roderick of the Environment
Department for their support, and to Paul Weldon who
provided shelter on (he summt of Mt Binnie. The
Gregory family of Tabubil and James Menzies
(University of Papua New Guinea) assisted us in
numerous ways and we are extremely grateful for their
hospitality. Roselyn Busasa (Institute of Papua New
Guinea Studies) and Dr Navu Kwapena (Department
of Conservation and Environment) facilitated the
processing of our research visas and export permits
respectively. Mare Hero kindly provided access to his
“Canary” program, and Lucy Smith produced figures
2 & 3, James Menzies and Richard Zweifel provided
uselul Comments On (he manuscript.

162 S. J. RICHARDS, T. C. BURTON, M. J. CUNNINGHAM & A. J. DENNIS

References

Bium, J. P. & Menzies, J, I. (1988) Notes on Xenobatrachus
and Xenorhina (Amphibia: Microhylidae) from New
Guinea with descriptions of nine new species. Alytes 7,
125-163.

Bock, W. J. & SHear, C. R. (1972) A staining method for
gross dissection of vertebrate muscles. Anat. Anz. 130,
222-227.

Burton, T. C. (1986) A reassessment of the Papuan
subfamily Asterophryinae (Anura: Microhylidae). Rec. S.
Aust. Mus. 19, 405-450.

Dusois, A. (1988) Miscellanea nomenclatorica batracho-
logica (XVII). Alytes 7, 1-5.

RICHARDS, S. J., JOHNSTON, G. R. & Burton, T. C. (1994)
A remarkable new asterophryine microhylid frog from the
mountains of New Guinea. Mem. Qld Mus. 37, 281-286.

ZWEIFEL, R. G. (1972) Results of the Archbold expeditions.
No. 97. A revision of the frogs of the subfamily
Asterophryinae, family Microhylidae. Bull. Am. Mus. Nat.
Hist. 148, 411-546.

(1985) Australian frogs of the family Microhylidae.
Ibid. 182, 265-388.

—_____ & Ty Er, M. J. (1982) Amphibia of New Guinea pp.
759-781 In Gressitt, J. L. (Ed.) “Biogeography and Ecology
of New Guinea” (Dr W. Junk, The Hague).

MANUNEMA PECTENOPHORA SP. NOV. (PERESIANIDAE,
LEPTOLAIMINA), A NEMATODE POSSESSING UNUSUAL
MALE SUPPLEMENTARY ORGANS

By AIMORN C. STEWART & WARWICK L. NICHOLAS*

Summary

Stewart, A. C. & Nicholas, W. L. (1995) Manunema pectenophora, sp. nov.
(Peresianidae, Leptolaimina), a nematode possessing unusual male supplementary
organs. Trans. R. Soc. S. Aust. 119(4), 163-169, 30 November, 1995.

Manunema pectenophora, sp. nov., with three unique pre-anal male supplementary
organs, is described. These are comb-like organs held clear of the body on short rods.
Two previously described species of Manunema, the sole genus in the Peresianidae,
possess tubular supplements. M. pectenophora also differs from the other species in
that the single testis is anterior. All Manunema species possess four long cephalic
setae, no labial setae or papillae, circular amphids, a strongly annulated cuticle, a
narrow tubular buccal tube, a narrow cervical region expanding to accommodate the
strongly muscular pharynx, two outstretched ovaries ventral to the gut and simple
curved spicules. The taxonomic placement of the Peresianidae is difficult but the
conclusion of other taxonomists that it belongs within the Leptolaimina is supported.
Key Words: Taxonomy, marine nematodes, Peresianidae, Manunema.

Transactions of the Reval Seeterv of S dust (995), L941, 163-169,

MANUNEMA PECTENOPHORA SP. NOV. (PERESIANIDAE,
LEPTOLAIMINA), A NEMATODE POSSESSING UNUSUAL
MALE SUPPLEMENTARY ORGANS

by AIMORN C. STEWART & WARWICK L. NICHOLAS*

Summary

Srmwarr, A.C, & NICHOLAS, W, L. (995) Manunema pectenophorn, sp. noy. (Peresianidae, Leptolisnina),
dnemulode possessing unusual ale supplementary organs. Trans. Ro Soc. §, Alest, W9(4), 163-169, 40 Novernber,

1995.

Manurnema peclenephera. sp. noy.. with three unique pre-anal male supplementary organs, is deseribed, These
are comb-like organs held clear of the body on short rods, Two previously-described specios of Mamurreme, the
sole genus inthe Peresianidac, possess tubular supplements. M. pectenopliara also diflers from the other species
in that the single tests ts anterior, All Manunente species possess four long cephalic setae. no labial setac or
papillae, circular amphids, a strongly annulated cuticle, a narrow (ubular buccal lube, a narrow cervical region
expanding lo avcommoadate the strongly muscular pharynx, (Wo outstretched ovaries ventral to the wut and simple
curved spicules. The taxonomic plucement af the Peresvaradae ts difficult but the conclusion of oiher taxonomists

that jt belongs within the Leptolaimina is supported,

Key Wokbs! Taxonomy, murine nematodes, Perekianiduc, Munanenns.

Introduction

Mununema pectenoplara sp. nov, possesses
prominent male supplementary organs. i.e. ventral pre-
anal organs found in many male nematodes, but in the
new species they are unlike those described previously,
The Peresianidie contains a single genus, Manunema,
comprising only two previously-described species.
namely M. proboscidis Gerlach, 1957, and M. catnilata
(Vitiello & de Coninck 1968) Riemann, et al. 197i,

The taxonomic placement of the Peresianidae has
proved a problem. Some characters. suggest placing the
farnily in the Leptolaimina (Chromadorida), others are
closer io the Desmoscolecoidea (Monhysterida).

Materials and Methods

Specimens were collected fram the intertidal zone
ol beaches at Darwin NT. Samples of about 2 kp of
sand were dug up at low tide and the meiofauna present
was briefly suspended in 5 litres of tap Water with
vigorous stirring. As soon as the sund had settled, the
water was passed through a 60 pm nylon sieve and the
fauna retained on the sieve back-washed into a beuker
with sea water. They were immediately fixed by adding
formalin to give a final concentration of 5%- Later,
the mejofauna Was examined in petri dishes under a
binocular microscope, The new species Was isolated
by pipette from the many hundreds of other nematodes
collected and the nematodes mounted on microscope
slides in anhydrous glycerol. Cover slips were

+ Division of Botany and Zoolayy, Australian National
University, Canberra ACT 0200,

supported by glass beads (Ballatinr) selected under the
microscope to be slightly wider than the nematodes
and the cover slips were ringed with Glyceel. (Gurr).

Measurements are in jem from specimens fixed and
mounted in this way. De Man's indexes (ratios)
(Portuner 1990) are given, i.e, a= body length divided
by greatest body width. b=length divided by length
of pharynx, c=length divided by tail length, c'= tail
length divided by width at anus, V =anterior end to
vulva as a percentage of body length, and spicule
measurements are arc length,

Drawings and measurements were made using a
cameta lucida. When mounted, the nematodes lie on
their sides presenting a lateral view, and our drawings,
with the exception of all four cephalic setae. show setae
on one side only, those lymy uppermost as mounted,

For scanning electron microscopy, some specimens
in 5% formalin were washed in phosphate buffer, pH
7, containing 3% sucrose. post-fixed by the addinon
of 2% osmium tetroxide, washed, sonicated and finally
freeze-dried. The specimens were mounted on metal
stubs and coated with gold/palladium before
examination in the microscope.

Type specimens are deposited in The South
Australian Museum, SAMA, Adelaide. and their
numbers tn the Museum's Australian Helminth
Collection, AHC, are given in the text.

Manuneéma pectenophora sp, nov
(FIGS I-ll)

Holotype: Male, Rapid Creek beach, Darwin, NT,
19.x,1992, SAMA, AHC 30000.

Measurenients: Table |

164 A.C, STEWART &

TABLE | Measurements of Manunema peetenophora sp, ier

W. L, NICHOLAS

Type Holo Male paratypes j= 4 Female paratypes n= 4
Male Range Mean +5) Range Mean +8D
Length 448 466-506 490 ) 475-50%. 497 7
Maximum width ly I-12 a OS 16-20 18 183
Cephalic setae 13 10-16 \3 3.06 12s {5 2.50
Body. setae i} Ila I2 1.73 W-14 Q [.83
Mouth to amphid 12 10-11 10 O58 9-9 {) 150
Amphid diameter 47 303.2 3.1 0.12 3.6-3.6 36 0.00
Width at amphid 6.5 5.0-6.5 5.8 O76 5.6-5.6 5.0 0.05
Bucea! cavity Bie) 32-46 3a 2.08 Waa a3 150
Width at buccal cavity 12 Y-lt {0 145 942 iT] 1.50
Muuth to nerve ping 5S} 34-60 a7 3,06 53-00) a4 2.08
Width wt nerve ring i) 13-14 13 O58 Ik 16 173
Pharynx 83 77-83 80 i §2-88 85 2.58
Width at cardia I+ 1-17 (4 2 52 MW) 1k 14 249
Mouth to vulva . - - - 252-273 265 10
Width at vulva - - - - 15-20 18 [83
figy : 4479 57 20
Mouth 10 utlus 341 391-434 42 22 407-448 426 4
Tail 3 72-84 78 6 67-74 71 3.77
Width up anus i] I-11 il u 8-11 0 1,29
Spicule, are length 4 23-25 24 |
Gubernaculurn ll 10-11 W 0,99
Anus jo Ist supplement” 68 4,5-7,3 64 LA -
Anus to 2nd supplement* 1B 1-15 3 2:25; -
Anus to ard supplement’ a4 3-34 33 162 - - -
De Man's a 33 3H 39 ai] 2.63 25-30 1k 26
De Man's h 5.9 5.6-6.6 44 O47 57-62 5.8 (1.24
De Mans ¢ 6 5,9-7.0 63 0.63 69735 70 (122
De Mars ¢° 74 6.57.6 7A 55 68S 75 O96
De Mun’s V Ye - - 5153 33 148

AAs pereentage of body length

Deseriprion of Holotype male

Small, body when fixed strongly curved, head and
cervical region folded back along body, tail curled.
Cuticle strongly anmulated: lateral ridges from oid
pharyngeal region to mid tail, wavy in register with
annules: four rows of prominent body setae, arising
from pronounced cuticular hemispherical swellings,
dorso-lateral setae alternate with ventro-lateral setac.
Four long cephalic setae arising form sackets; labial
setae absent: amphid circular, Buecal cavity, woah
minute ridges around mouth, mitially narrowly conicul
extending posteriorly us a narrow parallel-sided tube
Pharynx, in vervieul region (35% of pharynx length)
narrow purallel-sided, encloses buccal tube, then a
wider muscular cylinder, somewhat constricted hy
prominent nerve ring, two cytoplasmic clefts between
herve ring and expansion; cardia short, eylindrical
Intestine simple tube, anus and rectum project slightly
from body contour; caudal glands not observed
(probably obscured by strong annulation), Single testis
lo left of intestine; spicules cephalated, smoothly
curved, lips pointed; gubernaculum slightly curved
plate. Three pre-anal supplementary organs, most
anterior one about mid-way between cardia and anus,
the other twe close to anus. Each supplement resembles

an outwardly and slightly forwardly directed comb,
with about I prongs. mounted on a cuticular rod
arising deep in the body wall.

FParutypes; SAMA. AHC 300017. Measurements of
three males and four females are given in Table |. In
paratype males, as in the holotype, anterior supplement
about 33% of body length in front of anus, second and
third supplemenis, closer to anus, apparently more
variable in position, probably due to different degrees
of body curvature, Long testis, to left of intestine. with
many developing sperm, begins just anterior to mud
body, continues as long sperm duct. SEM of another
nile, Figs @ and 7. shows a tenous transparent film
overlapping the base of a supplement and adjacent
cuticle. We interpret this as mucus. present oyer the
surfave of treshly fixed specimens and preserved hy
freeze-drying but lost when specimens are transferred
lo glycerol for light microcopy.

Females (Fig. 5) similar t males apart from
reproductive organs and absence of supplementary
orguns. Didelphic, two very short ovaries outstretched ,
ventral fo gul. Three females each have single large
egy, 43, 48 and 79 ppm long, respectively, overlapping
the vulva, 10 left of intestine. ‘The largest is probably
at an early stage of the first cleavage division.

MANUNEMA PECTENOPHORA SP. NOV, 165

3,4 25 um

Figs 1-4. Manunema pectenophora sp. noy. 1. Male head. 2. Entire male. 3, Supplementary organ. 4, Spicules and gubernaculum.

166 a

a x soam % *

om wt

yy
ey

< sien
ieee

Fig. 5. Female Maninema pectenophara sp vow

Differential diugnosis.

The form ob the Supplement distinguishes MM.
pectenophera sp. Noy, Irom the other described species
of Manunema, none of which possesses camb-like
structures mounted oo rods, The new species differs
From Af. annulate in the orientation of the sinule testis.

Habitat
Sandy ocean beach.

Distribution
So far known only trom Rapid Creek beach, a
suburb of Darwin, Northern Territory.

Epymology
Named from L., pecien, a comb.

Discussion

Supplementiry organs are common in many farnilies
of Adenophorea, where they are associated with
sensilla, and are generally believed to play a part in
copulation, They may be tubular, setose or papilliform
und are often ussociated with cuticalar ornamentation
but none like the organs described here has previously.
been reported. They do pot appear ty be associated
wilh sensilla and conceivably serve some mechanivul
role in copulation, M. proboseieis possesses two pres
unal tubular male supplementary organs (Gerlach
1957). Viticllo & de Coninck (1968) claimed. that
supplements were lacking in M. annii/ata, but Riemann
etal, (\97L) redeseribed M. annufara, reporting two
pre-anal tubular supplements. Neither Gerlach (1957)
nor Vitiello & de Coninck (968) comment on he
buceal cavity, We agree with Riemann eve, (971) thar
the buceal cavily 1s long and tubular, Lorenzen (98h)
includes a long tubular buccal cavity us one of the
diaynostic characters of the Peresianidae,

The taxonomic position of the Peresianidac, ta which

C. STEWART & W. L, NICHIOLAS

Manurnena belongs, has been the subject of sore
doubr. partly because their small size hus led lu some
uncertainty abour taxonomically iniportant characters,
His significwnt that scanning electron microscopy Uoes
not show either Guter Libial papillae or setae, nor any
external Wiandfestation of janer labial sensilla. Ail (he
described species Haye four Jong sub-median cephalic
setae inserted in sovkets.

In Lorenzen’s (1981) phylogenetic classification of the
Adenophorea, ovaries ventral te (he intestine and a
Single posterior lesus are significant characters in
Manuneny, consistent with the placement of ihe
Peresianidae in the Leptoluimina but. while the location
und Jorm of the ovaries in M. peetenaphora are the
game as in. M. proboseldis, we have observed a single
anterior testis. in three Males OFM. pectenpphere, The
form of the umphids, the long narrow buccal lube and
jubular supplementary organs are consistent with
Leprolaiminas ventral outstretched ovaries ure not
(Lorenzen 1981). In the possession of four cephalic
setae, the absence of outer jablal setae, the possession
ob four Sub-median rows of alternating body selite
arising trom peduncles and the anus on a protrusion
from the body cavity, Manunema resembles the
Desmouscolecoidea, within the Monhysterida, rather
than the Leptolaimina.

In Vitteilo and de Coninck’s (1968) view. the
similarities between Peresiane unnulata. now renamed
Mantimema annulate Rienvann et el (1971), and Melia
spinosa Gerlach 1956 jndicated a phylogeneue link
between the Hal\plectidae (Leptolaimina in Lorenzen’s
clissificalion) and the Desmnoscolecida, in which they
placed the new species. The similarities to which they
drew attenuon were the four cephalic setae and the
position of the non-vesicvular amphids, but in other
respeets the species are unalike, differing in the
structure of the cuticle, buccal cavity, pedunculate setae
and (he location of the anus. In fact. as Riemann ¢7
al. (971) point out, there ure sinmlarities between
Mertiw'me and other Desmoscolecoidea, for example
with Tricoma niirabiliy. Timm 1961, although Mann-
vema shows greater similarity with such Leptolaimina
as dnomenema haplostema Hopper 1963 and
Leplolaimus tritubulatus Boucher and Helleouét 1977.

Although M. pectenophora does nut possess tubular
supplementary organs Ord posterior testis (leptolanind
characters of Maniinema probuscidiy aad Mf. annulaia)
we cancir with the placement Of Peresianidae in (he
Leptolaimina, with a possible link between
Leptolaimina and Desmoscolecoidea

Acknowledgments

We thank Dr Russell Hanley for providing facilities
in The Northern Territory Museuniagd the Australian
Biological Survey for financial support for one of sy
to work in Darwir,

MANUNEMA PECTENOPHORA SP. NOV

In/

oW —
_
7 _
_ 4

Figs © and 7. Seunning electron microscopy of Manunemea nectenophora ap: nov. 6, Entire male. 7. Enlargement
supplementary organs. SQ supplementary organ

to show

168 A. C. STEWART & W. L. NICHOLAS

2
_

Figs 8-IL. 8. Scanning electron microscopy of female Manunema pectenophora sp. nov. CeS cephalic seta, AM amphid.
9. Female by light microscopy. 10, SEM of female head. Il. Male by light microscopy. SO supplementary organ.

MANUNEMA PECTENOPHORA SP. NOV. 169

References

FortTUNER, R. (1990) Ratios and indexes in nematode
taxonomy. Nematologica 36, 205-216.

GERLACH, S. A. (1957) Die Nematoden fauna des
Sandstrandes an der Kuste von Mittelbrasilien. Mitt. Zool.
Mus. Berlin 33, 411-459.

LoreENZEN, S. (1981) Entwurf eines phylogenetischen
Systems der freilebenden Nematoden. Verojff. Inst.
Meeresforsch. Bremerh. Suppl. 7, \-472.

RIEMANN, F., VON THUN, W. & LORENZEN, S. (1971) Uber
den phylogenetischen Zusammenhang zwischen
Desmoscolecidae un Leptolaimidae (freilebende
Nematoden). Veroff Idst. Meeresforsch. Bremerh. 13,
147-152.

VITIELLO, P. & DE CONINCK,. L. (1968) Peresiana annulata
n. gen., n. sp., type interessant de Desmoscolecida. Rapp.
Comm. int. Mer. Medit. 19, 201-204.

ASPHONDYLIA DODONAEAE, A NEW SPECIES OF
CECIDOMYITIIDAE (DIPTERA) DAMAGING LEAVES
AND BRANCHES OF HOP-BUSH, DODONAEA VISCOSA
(SAPINDACEAE) IN AUSTRALIA

By P. KOLESIK*

Summary

Kolesik, P. (1995) Asphondylia dodonaeae, a new species of Cecidomyiidae (Diptera)
damaging leaves and branches of hop-bush, Dodonaea viscosa (Sapindaceae) in
Australia. Tran. R. Soc. S. Aust. 119(4), 171-176, 30 November, 1995.

A new gall midge species Asphondylia dodonaeae, is described from South Australia.
Detailed descriptions of the larva, pupa, male and female as well as the infestation
symptoms on leaves and branches of hop-bush, Dodonaea viscosa Jacq. subsp.
spathulata (Smith) J. G. West (Sapindaceae), are given. The new species is diagnosed
and compared to other species of the genus Asphondylia.

Key Words: Cecidomyiidae, Asphondylia dodonaeae sp. nov., Dodonaea viscosa,
South Australia.

‘Dunsactions of the Revel Society ofS Aast (1995), 194), T7L17O.

ASPHONDYLIA DODONAEAE, A NEW SPECIES OF CECIDOMYLIDAE (DIPTERA)

DAMAGING LEAVES AND BRANCHES

OF HOP-BUSH, DODONAEA VISCOSA

(SAPINDACEAE) IN AUSTRALIA.

by P. KOLESIK*

Summary

Korpsik, P. (1995) Asphomiviia dedonaeac, a new species of Cecidomviduy (Dipteru) darhaging leaves anid
branches of hop-bush, Dedeneee viveuse (Sapindaccae) in Australia, Trang, R, Soe, S. Anse, W9(4), 17L17A. 30

Navermber 1995.

A new pull midge species dsphondylia dodonaese, is-described trom South Australia, Detailed deseripions
of ihe larva, pupa, nude and female as well as dhe intestation symplonis on leaves and branches ot bop-bush-
Dodinaec viscosa Jacq, subsp. spatiudata (Smith) 1. G. West (Sapindaveae), are given. The new species is diagnosed
and compandd te other species of the penus Ayphandylia-

Kiy Worns! Cecilamyidie, Aiphandylia dediaeae sp. nov, Dadonuea visvesa. South Australia.

Introduction

The new gall midge species described here was
found festing leaves and terminal branches of hop-
bush. Dodonaea viscosa Jacq. subsp. spaihutate
(Smith) 1. G. West (Sapmndaceue) in South Australia.

Dodemnaed viscosa Jacg. is a shrub of tree up ta &
mvtall It occurs throughout Australia. und extends intd
tropical Asia, America and Africa and inte temperate
southern Alricu, New Zealand and Pacific islands
(Reynolds & West 1985). [ts leaves arc used in various
parts of the world in folk medicine to control fever,
colic, inflammation, swellings, rheumatism and pain
(West 1984: Ahmad er al, 1987; Wagner et al. 1987;
Mata er af, 1991), In several countries it is used as
firewood. material for tool handles and for reclamation
of unused or degraded landscape areas such as sand
dunes, nmurshlands und mine wastes (Norem ef al.
1982; Reynolds & West 1985). In Australia a purple-
leaved form is grown widely. in gardens ard the foliage
is valued for its decorative appearance,

The hop-bush is «common shrub in remnants of the
original flora around Adelaide where it forms a
substantial part of the medium-high vegetation cover
if the pature conservatyon. parks. During, 1992-1993
large numbers of galls were found onalinest all shrubs
surveyed in Morialla and Cleland Conservation parks.
The new gull midge appears to have two generations
in the Adelaide area, the first from January to February,
(he sevond from September to October. Shrubs bearing
galls from two -suceessive generations of the gall midge
cau otlen be found,

* Depirtment of Horticulture Viticulture and Oenology,
Faculty bf Agricuiturt! and Natural Resource Seicnees,
University of Adelaide PMB | Glen-Osmonds, Aust. 5064,

Materials and Methods.

Leal and branch stem. valls of Dodenaed wscuse
subsp. sparhulara were sampled in Morialta (27 xi, 1992
and 26.ix.1993) und Cleland Conservation Parks
(3.7.1993). The parks are adjacent and located about
13 kin north-east of Adelaide, The galls obtained on
26.ix.1993 were processed in two ways. A small
number was dissected and the larvae (along with one
larva from 27.xi,1992) and pupae. were preserved in
70% ethanol atter notes were made on their colour,
A larger number, with larvae and pupae retained within
galls. was brought to the labortory to rear to adults,
Branches with galls were kept in plastic bags. Larvae
pupated in their galls. Plastic bags were examined daily
and emerged adults preserved together with their pupal
skins in 70% ethanol after their colour had been noted,
Canada balsant mounts of a series for microseopic
examination were prepared according to the technique
outlined by Kolesik (1995). The type series and other
milenals retained in 70% ethanol together with dried
examples of the galls are deposited in the South
Australian Museum. Adelaide |SAMJ. Australian
National Insect Collection, CSIRO, Canberra [ANIC]
und United States National Museum |USNM|.
Washington DC USA.

Asphendylia dodunaeae sp. noy.
(FIGS 119)

Holetype: of, Marjalta Conservation Park, South
Austnilia (34°54'S, 138°44°&), 29,1x,1993. P. Kolesik.
reared From larva trom leal gall of Dadenaca vivewsa
Jacq. subsp, spathulara (Smith) J, G. West, sampled
26.14.1993. 121272 [SAM].

72 P KOL EStK

Altpinnes 2. sume dita, 121273 [SAM],

Paranpes: 3%. 39 9, 4+ Jarvae, 4 pupal skins
|SAM|. 20 4 29 9.2 larvae, 2 pupal skins [ANEC].
ull same dau Lb larva. sunmpled 27.41.1992 [SAM].

Other marerial (OS a, WG SISAM|, Sa a,59 9,
JWSNM], 10 pupal skins [SAM|. 5S pupal skins
[USNM |. 10 pupae, all same dita as holotype. 3 karvae
[SAM|, 5 Jarvace [LISNM], all collected with holotype

Dreeneasis

Wings with R, jounny C at wing apex, Rs absent,
kK joining C at wing mid-length. M, ,. absent, M,
weakly develuped. Cu forked, Sc cell opaque
Plagellameres 12 in number, cylindrical wilh short
geeks, firstand second not fiised, with short and stout
setae and bearing anustomosing slightly appressed
vircumfila, Male flagellomeres ull uboul same length,
feinale ones, espevially the apical three. successively
and progressively shorter “Tarsus: first segment
substantially shorter than second, bearing ventroapical
spine; claws simple; empodia longer than claws, Male
rerinalia: gonocoxiles free ventrally, short. with small
apiual lobe, gonostylus situated dorsclly on ganacox ite.
short, bearing Wwe teeth merged basally* hypopruet and
vere? bilobate: aedeagus long. stout, tapering distally
Femule abdominal sternite 7 about (hree times jonger
than sternite 6. Ovipositor: clongate, sclerotized. wilh
large basal lobes; cerci fused. glabrous. bearing tew
microsetae

Male (Figs '-7)

Colour: selerouzed parts of body dark brown. selae
and Scdles black, non-sclerotized parts of abdomen
orange, Wing length 2.4 mm (range 2.2 - 2.6), width
fmm (10-12). Wing membrane and veins densely
vovered with setae, 55 - 120 pm, microtrichia dense,
about 0.5 pm long. Flagellomeres with sloul setae, 34-
38 jan, more or less equally positioned on the
segments, Cireumfila: two long and two shart
longitudinal bunds with long bands connected to each
other by transverse circular bands on both ends; each
of the short bands attached on batti ends to one of (he
long ones by shert wansverse arch; the transverse
virculur bunds on the distal end of the flaugellornere
arched strongly. Eye faeets rounded, eve bridge 8-¥
facets Jong. Maxillary palpus 4 or 4 segmented. offen
specimens with different number of segments in lett
und right maxillary palpus can be founds however, lolul
length of beth palpi about the same. Palpiger weakly
developed. Legs vovered with sete and seales_ the
Jatter serrated ut distal end.

Female (Figs 8-12)

Wing length 2.6 mim (2.6 — 2.7), width 1,2 mn (1,2

1.4). Flagellomeres with stout setae, 30 - 35 am

Circumfila comprising two Lransverse bands eotinected
by two Short longitudinal bunds. Chiws somewhat

stronger than tr male, Abdomjnal sternite seven 3,2
times (3.1 - 3,3) longer (har sternile sia. Setae of eeret
6-8 iy sumber and fess than | yn in length, Orher
ehuructers as in miule

Matare larva (elas 13-15)

Colour pale Orange Tot length 19 mini (17-221
Flead capsule Width 91 jan (90-92), length 29 am (26
31), length of posterolateral extensions 10 yan (9 - (0).
Antenna $4 pm (13 - (5). Sterna) spatulu bilobale, tt)
pm (08-116) in length, with apival enlurgenrene 6%
pn (04 - 74) ie wrth and. inepsian Fh pat (Al - Aa)
in depth. Arew around spatula not scleyotized. Ags
Jorsul, One pair of stemal papillae qa thoracic aud
first to seventh abdominal segments. One pair of ventral
pupillae on collar, seeond und third thoracic and first
to cizhth ubdontinal sezments. ‘I'wo pairs ef lateral
papillae on tharacic segments. Pleural pupillies Test
wnd third (heracté and list to enehtl) abdern|ial
sepmenis with one parr Second thoreic seemenk wilh
Iwo pairs. Dorsul papillie: voller, Third thoruere and
last two abdominal segments with one pair. first bye
thoracie and first to seventh ahdomindl segmenis with
lwo pairs. The setae on lateral papillae 3-5 iy long,
those on other papillae 4 - 20 um tong,

Pupit (Figs 16-18)

Colours antennal horns, prothoracie spiracles anil
Jorsal spines. dark brown, remaining paris pale brown.
Total Jength 3.0 mm (27 0 4.2) Attemnal hianns
triangular, serrated, (ol pom (147 - 182) tn Jereth
Cephalic papillae with seta 34 yan (36 - 44). Upper
and lower frontal horns absent. Two pairs of lower
facial papillae. each consisting of one seluse (5 - 15
jm) and one asetuse papilla. Two triplets af lareral
fadial papillde, cach consistmy Of two Setose labour
2 pint papillae and one asetose papilla, Prothoracie
born with trachea ending al is mid-length, 96 pam (44
103) long. Second to eighth abdominal segments with
two pairs of dorsal papillae (length of setae 8 - 14 juni
two pairs of pleural papillae (9 ~ 31 ym) and one pair
of ventral papilive (12 - 1 am), Dorsal spines simple,
46 - 104 in number und 8 40 jan in teneth, woth leqwtt
and number increasing from second to ninth segments,

Gall (Fig. W)

This species forms subglobular monothalumous walls
onrerminal branch stems and leaf main veins. glabrous,
4 mm Jeng and 3 mm in diameter, green in calour
One larva oveupies cuch gall. Puputiow takes place
inside the pall, Circular necrotized tissue area, Hrown
jn colour, Uppears.on the top of the gall before the pups
Cols a circular openiow with its dutennal horns: by
moving ils body up und down, The lid to this opening
remains attached nthe gall by a thin strip of uncut
lissue, The pupa raises two thirds of its body Quiside
the gall shortly before emergence ay adult. On 24

A NEW SPECIES OF CECIDOMYIIDAE FROM DODONAEA VISCOSA 173

Figs 1-7. Male of Asphondylia dodonaeae sp. nov. |. Last three flagellomeres, 2. Sixth flagellornere. 3, Wing. 4. First tarsomere.
5. Last tarsomere with claw and empodium. 6, Head in frontal view. 7, Genitalia in dorsal view. Scale bars = 100 pm,

174 P, KOLESIK

10°—

Figs 8-12. Female of Asphondylia dodonaeae sp. nov. 8. Sixth flagellomere. 9. Mouth parts in frontal view. 10. Antenna.
Il. End of abdomen in lateral view. 12, End of ovipositor in ventro-lateral view. Scale bars = 100 pm.

A NEW SPECIES OF CECIDOMYHDAE FROM DODONAEA VISCOSA 175

Figs 13-18, Asphondvlia dodonacae sp. nov. 13-15 larva. 16-18 pupa. 13. Sternal spatula. 14. Head capsule and collar segment
in dorsal view. 15. Last two abdominal segments in dorso-lateral view. 16. Anterior part in ventral view. 17. Last two
abdominal segments in dorsal view. 18. Prothoracic spiracle, Scale bars = 100 jm,

Ly P BOLESIK

Pus. WoGalhs of typi wWieedodemacae sp. now. on Davlonaert
Wycosd Jacq, subsp. sperfalare (Smith) J. G. West. Seule
hur > 2 em.

November 1992. the vast majority of the galls were
dried, Only a few of them sul contarnmed larvae. On
3 January 1993, the galls were fresh and contained
inimalure larvae, On 26 September 1993, thost of the
galls were occupied by pupae, with a few vecupied by
larvae una few already empty. Empty galls retained

pupal skins in openings. On cach ol the latter oecasiins
a few dried galls from fhe previous generation were
presenton the stiribs, The species seems to hive (wo
generations in the area surveyed — udulis of the first
generation appear possibly from January to February
and those of the second generation [rom September
to October.

Etymology
The species name is derived from the generic nanye
of the host plant.

Remarks

The new species can be assigned to the genus
Asphondylid because the female seventh ubdominal
sternite is more than £5 times longer than the sixth.
the male genitalia have a ventroapical gonocoxal lobe
and dorsally situated gonostylus that is about us brouwd
as lone, Combined with the first (arsomeres having u
ventrodistal sping, the gonostylus hearing two basally
merged teeth and the uvipositor having large basal lobes
(Gagne 1994). Within the venus Anpfrenedydia it 1s
disdnguished fram other species by lacking both upper
and lower Jromtal horns im the pupa,

Acknowledgments

The Ministry of Environment und Planning, South
Australia, kindly permitted collection within the nature
conservation parks of Moriulta and Clekind. Martin
C, O'Leary, State Herbarium of South Australia,
Adelaide, courteously identified the host plant species,
lam grateful to David B, Hirst, South Australian
Museum. Natur! Serenve. Jolin DB, Gray. University
of Adelaide. Depurtnient of Horticulture, Viticulture
and Oenology and Raymond J Gagné, Systematic
Enlomology Laboratory; USDA, Washington DC USA,
lor their comments on wy early draft of the manuscript

References

Auman, Vo U. Parma, to & Faia. 4, (987) The
saponins trom Dedended visewsd. [iuenaptia 38 36) 362

Grom. Ryd. (804) “The Gall Midges of ihe Neotropical
Keaion” (Cornell University Press, [duet New York).

KOLESIK. P1995) A new species al EZoemercornia Felt
(Diptera: Cecidomyidae) on Raculypiis feaseiculasa vn
South Australia. 2 Aw ent Soe 34, 197-152

Maia, BR, Cosiketas. J L., Orisanto, DL, Preis
Mikanoa, R., Castanppa, BP, Rie, Fort & Dri-Rin,
fh 991) Chemicul studies on Mexicin pints used jn
tradinional medicine, XVIL. New secondiry metabolites
trom Poduneee vacesa. 2 Nur Prod, $4, 912-917,

Rokpm, M, A,, Day. A, BD, & Libeke, KR. Lb, (982) An
evaluation of shrub and tree species used for revegetuting
copper ming wastes inthe south-western Linited States, /,
drt bnviran & 290404.

Revxoius. 8. T) & West 4, G, W085) Sapmdavene jy
4-1n4 Ji George. A. S. (Bd) “Flora of Austrtlia’ Vol. 25,
(Australian Government Printing Serviee. Canbernit),

Wacwer, Ho Lupwie. C_, GRotIAHN, L. & KHAB, Mo
Y (JOR7) Biologieully active saponins Han Liaderianes
viveosa, Phytochen, 26, 697-701.

Wesi, JG, 984d) AD revision ol Dadondqen
(Sapindaceae) im Austrailia, Branovia 7, 1194,

Mills

CONTARINIA BURSARIAE, A NEW SPECIES OF
CECIDOMYTIIDAE (DIPTERA) INFESTING FRUITS
OF SWEET BURSARIA, BURSARIA SPINOSA
(PITTOSPORACEAE) IN AUSTRALIA

By P. KOLESIK*

Summary

Kolesik, P. (1995) Contarinia bursariae, a new species of Cecidomytiidae (Diptera)
infesting fruits of sweet bursaria, Bursaria spinosa (Pittosporaceae) in Australia.
Trans. R. Soc. S. Aust. 119(4), 177-181, 30 November, 1995.

A new gall midge species, Contarinia bursariae, (Diptera: Cecidomylidae) is
described and illustrated. Larvae found inside fruits of Bursaria spinosa Cav.
(Pittosporaceae) prevent formation of the seeds. Detailed descriptions of the larva,
pupa, male and female and the infestation symptoms are given.

Key Words: Cecidomyiidae, Contarinia bursariae sp. nov., Bursaria spinosa, South
Australia.

Transeetions of the Royal Saeiwty uf 8. Aust, (W999), MOC) PSL

CONTARINIA BURSARIAE, A NEW SPECIES OF CECIDOMYIIDAE (DIPTERA)
INFESTING FRUITS OF SWEET BURSARIA, BURSARIA SPINOSA
(PITTOSPORACEAE) IN AUSTRALIA.

by PB KOLESIK*

Summary

Ro.esi, P.(1995) Comurinia bursarian, a new species of Cecidomyiidae (Dipleny) infesting fruits. of sweet
bursuria, Barsaric spinosa (Pittosporaceae) in Australia Trurs, R) Soe 8. Auusr 11904), 177-181 30 November, 1995.

A new gall midge species, Conmarinia hursariae, (Diptera; Cecidomyiidael 1s described and illustrated, Larvae
found mnside Fruus al Rursarie spinosa Cav. (Pitosparaceae) preven! formation of (he seeds, Detajled descriptions
of the furva, pupa, oiile aind female and the infestation symploms are viven

ki Wows! Cecidomyiidae, Coarina barsariae sp, noy., Bursaria spiaosa, South Australi,

Introduction

Burseria spinosa Cay, sweet bursaria or Christmas
bush, is 4 shrub usually 1-3 mt call. The genus is
endeimi¢ to Australia. Bursarid spinosa can be tound
in South Australia, Queensland, New South Wales,
Vielona and Tasmania where it is common in
woodland vepetation (Bennett 1986). Voluminans
clusters of white flowers make the Shrub a useful honey
plant (Cunningham ere. (981), The gall midge species
described here was found to prevent seed production
in A sprmosa in Morialta Conservation Park. near
Adelaide.

Materials and Methods

Fruil capsules of Bursarta spinosu were surveyed
in Morialts Conservation Park (3 km north-east of
Adelaide) on 19 February 1995, Those which contained
Jurvac of the new species were brought to the laboratory
where the fruits were cut oped aid the extracted Jarvae
processed in two ways. A small number was preserved
in 70% ethanol after their colour had been noted. The
remainder were transferred with entomological forceps
into pots containing sterilised, wet.sand and reared to
the adult stage. Pots were examined daily and emerged
adulls preserved together with (heir pupal skins i 70%
ethanol afler their colour had been noted, For
microscopic exumination adults, laryae and pupac were
mounted on. slides in Canada balsam according ta the
technique ootlined by Kolesik (1995). The type series
and other material retained in 70% ethanol are
deposited in the South Australian Museum, Adelaide
[SAM| and Australian National Insect Collection.
CSIRO, Canberra [ANIC].

> Deparment ‘nt Horticulture, Viticulture: and Oenology,
Facahy of Agricultural and Natural Resource Sciences,
University of Adcluide PMB | Glen Osmond S$. Aust. 5064-

Contarinia bursariae sp. nov.
(FIGS. 1-13)

Holotype: @- Morialta Conservation Park, South
Australia [34°54'S, 138°44’E], 2.i01.1995, P. Kolesik,
reared from larva (rom fruit of Burseria spinosa Cay. ,
sumpled 19,i1.1995, [21274 [SAM],

Allonype; 2, same data, 121293 [SAM].

Paratypes: 2 & 29 9,1 pupal skin [SAM]. 20°",
29 %, | pupal skin [ANIC], all same data but emerged
2.111995 - 61,1995, 4 larvae [SAM], 2 larvae
[ANIC]. sampled with holotype.

Ovher material: 21 larvae, sampled with holotype [SAM].

Male (Figs |-6)

Colour: anienna grey. head black, thorax brown,
abdomen with scleronzed parts browm and non
sclerotized parts yellow, legs grey with black scale
strips alang segments, Wing length 1-26 nim (1.19 -
1,31), width 0.47 mm (0.44-0.51), Vein C broken at
juneture with R.. M, in form of stripe of setae, Ry
selerouzed on base only: Wing membrane covered with
setae, 17 - 22 pm long. Abdominal tergites 2 - 6 with
caudal setae only. Head with postvertical peak present.
Eye facets rounded, eye bridge 8 - 10 facets long med:
ially. Eight fronte-clypeal setae tn all specimens.
Antenna total length 1.43 mm (1.32 - 154), Length
measurements of third Hagellomere (zm): proximal
niode 30 (28 - 32). proximal neck 17 (14 - 18), distal
node 36 (34-38), distal neck 28 (24 - 31). Circumfilar
loops reaching the mid-length of the next node, Tarsal
claws curved at mid-length, about as long as-empod
ium. Genitalia: gonocoxite setose and setulose, gono-
stylus with strongly sclerotized claw and un array of
plates below it, sparsely setose with densely setulose
pouch ut base: hypoproct deeply divided mediully, with
one Seta on each lobe, setulose; cerci deeply divided
medially, setose and setulose, aedéagus us long as
hypoproet and cerci,

178 P. KOLESIK

Figs 1-6. Male of Contarinia bursariae sp. nov. 1. Head of frontal view. 2. Last flagellomere. 3. Fourth flagellomere. 4
Genitalia in dorsal view, 5. Wing. 6. Last tarsomere with claw and empodium. Scale bars = 100 pm.

A NEW SPECIES OF CECIDOMYIIDAE FROM BURSARIA SPINOSA 179

Female (Figs 7-9)

Wing length 1.28 mm (1.23 - 1.37), width 0.48 mm
(0.43 - 0.53). Antenna total length 0.79 mm (0,69 -
0.83). Third flagellomere with node 45 wm (43 -49)
and neck 6 pm (5 -8) long. Circumfila appressed,
consisting of two transverse rings connected by two
longitudinal bands. Other characters as in male.

Mature larva (Figs 12-13)

Colour yellow. Total length 2.44 mm (2.20 - 2.75),
diameter 0.18 mm (0.14 - 0.21). Integument smooth,
ventrally with several transverse rows of spiculae on
anterior half of each segment as well as with
longitudinal rows around anus. All ventral, pleural.
lateral and dorsal papillae with minute setae, sternal

Figs 7-9. Female of Contarinia bursariae sp. nov. 7. Last three flagellomeres. 8. End of ovipositor in dorsal view. 9. Abdomen
in lateral view. Scale bars = 100 pm in 7 & 9: 10 pm in 8.

180 P. KOLESIK

13:

Figs 10-13. Contarinia bursariae sp. nov. 10. Larvae inside fruit capsule of Bursaria spinosa Cav. (left loculus with fruit,
right one infested). 11. Anterior part of pupa in dorsal view. 12. Sternal spatula of larva. 13. Terminal segment of larva
in dorsal view. Scale bars = 5 mm in 10; 100 pm in I-13.

A NEW SPECIES OF CECIDOMYIDAR FROM BURSARIA SPINOSA LSy

papillae asetose. Terminal segment with one pair of
stublike. asetose papillae and three pairs al setose
papillae, with one of the three pairs having longer setae
than the ouler two, Head capsule width ST aa (43-54),
length 40 ym (37-45), Jength of posterolateral
apodemes 42 aor (35-46), Sternal spatula 152. jn
(139-175) in Jength, with apical enlargement 44 pm
(41-47) in width and 20 ym (J9-2)) in length, Larva
cain jump short distances hy arching its body and
inserting, its posterior end between the spatula
enlurgement und the integument and by subsequent
quick teleusing of the posterior end.
Pupa (Pig. Wb

Head with sinall, angular, slightly sclerotized
antennal hortis. Cephalie papillae with seta 223 - 250
win long. Two pairs of lower facial papillae, one of
cach pair setese (Ml - 23 ym) and one usctose, Two
toplels of lateral facial papillae, one of each triplet
setuse (about 5 pm) and two asetose. Prothoracic
spiracle with trachea ending at its dpex, 133 to 168 aim
long, Second to eighth abdominal segments wath
strongly selerolized, simple dorsul spines, 5 - 15 in
number and 4 - 25 pm in length,

Infestation syimiproms (Fig, 10)

The miestation of Bursarta spinesa by Crnuarieie
hursarige can easily be overlooked because there is
no apparent mulformation of the fruit capsules.
However, in transmitted sunlight several larvae can be
recognised inside the capsule, The larvae occupy ane
or both locules of the capsule, preventing the
development of seeds. Up to eight larvae were obsetved
within individual fruits. Despite the absence of seed
in intested fruit no siymificant decrease in the total seed
production per plant was observed due to the low
infestation incidence in comparison lo the enormous
number of fruit per plant,

Etymology
Derived from the generic name of the host plant.

Remarks

The genus Comarinia is one of the largest genera
if Cecidomyiidae represented in all zdogeagraphical

rewions. Larvae of all known species are phylophayous,
most live gregariously in Mowers, buds and fruits which
are often mallormed (9 galls. Others are found in
Tnalformed leaves und stems. Almost all known species
are hosi-specific., sometimes with different species
living on the same plant. The genus Cunteriaia in the
context Of this paper is defined is below, Larva:
terminal segment with 6 sctose papillae and 2 jarge.
stublike, asetose papillae. Adults: maxillary palpus
with 4 segments, antenna with 12 flagellomeres: wings
with R, joining C beyond wing apex: tarsal claws
simple on all legs. Male: flagellomeres binadal, with
a single series of circumfilar loops on each node:
genitalia with stout, unlobed gongcoxile, slightly
tapered gonustylus, bilabed hypoproct and simple,
short, destally tapering vedeagus, emule: ovipositor
very long, retractable, the cerci tiny, dorso-ventrally
flattened, and closely approximated mesally,

The genus Conturinia is known im Austratia from
12 species, all of them from inflorescences and seed
heads of Gramimue und Cyperaceac (Harris 1979). The
species described here differs morphologically from
the previously-desertbed Australian species in the
number of setae on lermale cerer and the relative length
of male circumfilar loops: female eerci bear eight setae
in C_ bursariae; those in all the other specics bear more
than 14: male circumfilar loops reach the mid-length
of the next node in C bursarae; those in the other
species never extend beyond the base of the next node,

Acknowledgments

The Ministry of Environment and Planning, South
Australia, kindly permitted collection in the Morialta
Conservation Park, Martin €. O'Leary. State
Herbarium of South Australia. Adelaide, courtcously
identified the host plant species. Special thanks go to
John D. Gray, Deparment of Horticulture, Viticuluire
and Oenology, University of Adelaide wnd Rayrnand
J, Gagné, Systematic Entomology Laboratory, USDA,
Washington DC USA, for commenting on am carly
draft of the manuscript.

References

Bensptt, & M. (1986) Pamily Pitosporaceae pp. 429-436
In Jessop, 4. Poand Toelken, H.R. (Eds) “Flora of South
Australia, Part 1 (Lycopodiaceaé - Rosaceaey” (South
Australian Government Printing Division, Adeluide),

CUNNINGHAM, G. M_, MULHam, W, E,, MiitHorre, PL...
& Leck, L. H, Y8t) “Plants of Western New South
Wales” (New South Wales Government Printinye ONice,
Sydney}.

Hareis, & M_ (1979) Deseriptions and host ranges of the
sorghum midge, Conzarinia.sorghicola (Coquillet) (Diptera:
Cedidomy udae), and of eleven new species of Contarinia
reared from Graminae and Cyperaceae in Auswatia, Bull,
em. Rex, 69. 164-182,

Kovesik, P. (1995) A new species of Luninesicarina Felt
(Diptera: Cecidumyrudae) on Auvalypins farciculasa in
Sauth Aostralia. “. aux ent. Soe, 34, (47 182.

A REVIEW OF THE SCALE INSECT SUBTRIBE ANDASPIDINA
(HEMIPTERA: COCCOIDEA: DIASPIDIDAE) AND A NEW GENUS,
NOTANDASPIS, FOR TWO AUSTRALIAN SPECIES

By D. J. WILLIAMS* & H. M. BROOKESt

Summary

Williams, D. J. & Brookes, H. M. (1995) A review of the scale insect subtribe
Andaspidina (Hemiptera: Coccoidea: Diaspididae) and a new genus, Notandaspis, for
two Australian species. Trans. R. Soc. S. Aust. 119(4), 183-189, 30 November, 1995.
The subtribe Andaspidina is recognised as one of three subtribes of the scale insect tribe
Lepidosaphini. A review of the literature is presented and diagnostic keys are given to
subtribes and to genera of the subtribe Andaspidina. Notandaspis gen. nov. is described
for Mytilaspis (Coccomytilus) hymenantherae Green, a species described originally
from Victoria and presently included in Andaspis and for a new species Notandaspis
oodnadattae sp. nov. from South Australia. The new species is unusually large for the
subtribe.

Key Words: Coccoidea, Diaspididae, Andaspidina, Notandaspis gen. nov., Notandaspis
hymenantherae (Green), Notandaspis oodnadattae sp. nov., scale insects, Australia.

Tranwactions of the Reval Seevery at & aise 99S), MP9), PRS TEE,

A REVIEW OF THE SCALE INSECT SUBTRIBE ANDASPIDINA

(MEMIPTERA: COCCOIDEA;: DIASPIDIDAE) AND A NEW GENUS, NOTANDASPIS,

FOR TWO AUSTRALIAN SPECIES
by DoS. WILLIAMS*® & H. M. BROOKES+

Summary

Withiams. D.. & Broones, HM. (995) A review of the scule inseer subtribe Andaspidina (Hemiptera:
Coccoulen, Diaspididie) und it new wenus, Nelondaspes, lor two Australian species. Trams, R, See. 8, Aust, M94),
ISS-1K4. GU November Ys,

The subtriby Antlispidina ty recognised as ne of tree subtibes of the scale mseet tribe Lepidosaphini. A
review ol the literature is presented and diagnostic Keys are given to subtibes and to genera of the subtriby
Andaspidind, Norandaspis ser nov is described tor Murildspis (Coccomvitusy hymenditherie Green, a species
Jeseribed originally trom Victoria and presendy ineluded in andaspts and for anew species Notanelespis-oodnadattae
spy gov, from Sourh Australia, Phe new species is unusually large dor the subtribe,

Kiy Worns: Coccoides, Dyaspilidue, Andaspdina, Nefandasis gen nov, Nelendaspis Aymenantherae

(Green), Netandaspiy ccdnadattte sp. nov,, scale insects, Ausinaliin,

Introduction

Although neurly 250 species of Australian armoured
scale insects (family Diaspididae) Have so tar been
deseribed, most of the endemic species cannot be
recognised from the original deseriptions without
referring Wo authentic specimens in collections. A few
species have been redeseribed as parl of revisions ol
veneru but there js a pressing need for a complete
revision Gf all the named species. Since a catalogue
of world species was published by Borchsenius (1966)
ihwould be fairly easy to extract most of the pertinent
literature on Australian species. However. the work
involved in alse describing the new species already in
collections, dnd those still to be discovered, estimated
al inuny hundreds, could take many years, Numerous
exotic Species have also become established in
Australia, some eausing damage to cultivated crops und
irees and these alse need revision.

In the present work two species are described in the
subtribe Anduspidina, Australian species at present
ussigned (oO this group are damduapis Aynienantherae
(Green). do inelser (Green). AL ninterete
Brimblecontbe and Menindaspis recurvate (Froggatt).
A. livmenantherae is assigned to a new genus in which
a new species with an unusually large adult female is
also jnelided,

* Deparment ol Fntoratogy, The Natural History Museum,
Cromwell Road, London SW7 SBD

¥ Department of Crop Protection, Waite Campus, University
of Adelaide, Glen Osmond, S. Aust. S04.
Current address: § Yeo Avenue Hivhgate, S. Aust. $063

Materials and Methods

The species are deseribed from slide-mounted speei-
mens of the adult female and the illustrations show the
dorsal aspeet On the left and the yentral aspect on the
nght. Morphological (erminology is the samme as that
used in Williams & Watson (1988) where reference may
also be mude to a generalised illustration of the adult
female. Further specimens have been prepared on
mivrascape slides for this study using the techniques
discussed by Williams & Watson (1988).

The term megaduct was adapted by Takagi (1992)
from the term meygapore proposed originally by
Balachowsky (1954). These duets. when present,
numbering 2-7 on each side of the pygidial margin.
are enlarged and are much larger than any others on
the dorsum of the pygidium. The orifice of each
megaduet ts longitudinally cliptical and surrounded
by a heavily sclerotised rim.

Abbrewiations of the depastlories are aus follows:
ANIC, Australian National Insect Collection, CSIRO,
Canberra, Australia.

BMNH. The Natural History Museum, London, U.K,

Historical Review of Andaspiy and related genera

Ih the present work two Uibes, Diaspidini and
Lepidosaphin) are recognised in the subfamily
Diaspidinae. Based on the works of Borehsenius (1966)
and Balkichowsky (1968) the subtribes Andaspidina,
Lepidosaphina and Cocesmytilina are available in the
tribe Lepidosaphini and are here accepted. Genera of
the subtribe Andaspidina include Andassiy
MaeGillivray, Cafe Williams, Péretedespis Mamet.
Metandaspis Williams, Saotemaspiy Balachowsky und
the new genus Norandaspis gen. nav. here deseribed.

Ié4 4 WILLIAMS & H M. BROOKES

The names Lepidosaphini and Lepidosaphing ate
sed here walhout inflection formed {pom the nomjnal
genus Lepidesaphes Shimer despite the varicus
spellings Lepidosuphedion, Lepidusaphidin,
Lepidosphedina tind Lepidosaphidinu.

The genus Anedaspis was tamed ny MacGillivray
(921) with Myrilaspis flava var hawaifensis Maskell
4s type species, MacGillivray also ineluded the
Australian species Lepidasaphey ineisor Green, Hall
(1946) accepted the venus and ineluded the African
spocies Lepidusephes punicue Laing. Rua & Ferrys
(1952) revised Andanpiy and meluded 10 species, eight
af which were trom Asia. Brimblecombe (196)
described the pew specs A. numeriia Frovy
Queensland. Takagi & Kawai (1966) deseribed four
new species of dadaspiv trom Japan and added further
records of previausly Ueserthed species.

In a detailed study of adult mules, Ghaur (1962)
uccepted the subtribe Lepidesaphidina to inelude
Lepidosaphes Shiner and Andaspis, Lepidosaphidina
was accorded equal rank to the Diuspiding of the tribe
Diaspidini,

Williams (1963), in a review of Anidaspis, accepted
22 species and provided 4 key, Alsu imeluded in the
review were the new genera Caia, With C quermea
Williams from Pakisian as type species, und
Metindaspiy with Mytilaspis recurvara broggatt
deseribed from New South Wiles. us type species. He
also included Meiandaypis javanensis Williams from
Jaya and stated thal both new genera were related ta
Andaspis.

Ina catalogue of o-Galled Diaspidoidea of the world,
Borehsenius (1966) recoynised (he Uribe
Lepidosuphidini Shimer and the two subtribes
Leposuphidina and Coccomytilina Borchsemus, He
Included -dadaypiy and Caie in’ the subtribe
Lepwdosaphidiag and Meferdaspls in the subtribe
Coccomytilina and transferred the Australian species
Mynlaspis (Corcemytiles) hymenantherae Green to
Andespis.

Mamet (1967) described the new genus Purunilaspia
with 2 vinson’ Mamet fron Maurititié as type species

Barchsenius (1967) desetibed the genera Ravaspis
Borchsenius with Andispis mori Ferris as type species,
Pararaaispis Borchsenius with Lepidesaphes meliae
Green as lype species and Reenwalespis Borehsenius
with type species Reonwalaspis guercicnla
Borchsenius, The new species Ravaspis tnedica
Borehsenius, &. rae) Borchsenius and Roomvulaspis
quercicala dese cibed in the sane paper were purpaned
io be Indian in origin but Danzig (1968) indicwed that
the localities on all the original labels were in China.
Takagi (1970), diseussing the Diaspididae of Tarwan,
synonymised the names Ravespis, Paruraoaspis and
Roonwalaypis with Andaypiy bul sugpested shat the

genent may be valid in some degree ys species-zrupss,
All Uhpce genera desertbed by Borchsenilis possess
pygidial mepaduets

Baluchowsky (196%), unaware of Mamet's
Parandaspis, described the new vents Prrandusper
with P castelbrance’ Balachowsky as type species, He
also discussed the tribe Lepidosaphedini and erected
anew subtribe Andaspidina to include Andaspiy, Cute,
Metandaspis and his new genus FParandaspis, He
provided a key to the three subtribes Lepidosaphedinas,
Coccomynlina and Andaspidina and a key to the gepers
of the sublribe Andaspidima,

Raluchowsky (1973), realising (hat the nume
Purundaspiy Balachowsky was a junior bomonym of
Parandaspis Mamet, proposed the name Savromespiy
Balachowsky to replace Paranedaspis Balachowsky with
S. cuylelhrancyi as type species,

Williams (1980) synonymised the name @, dase
Willaims, deseribed from India. with 4. mummers
Brimblecombe and commented an its distribution th
Australia and the Pacifie region and its association with
jhe symbtorie tungus Sepiebayidiam sp.

Willitms & Watson (988) discussed the Pacific
species of 4rdespis including two new species from
Papua New Guinea.

Takagi (992) commented on some Unusudl gener
of (he Lepidosuphedini ay a tribe of the subtiinily
Diaspidinae and suggested thar Merandaspiv
Javanensis, based ona study ob the first mister and adult
femule, was 4 Somewhat odd forny’ bul could beloms:
ty the (ribe,

Danzig (1993) recently accepted only the tribe
Lepidosaphini without subtribes.

Systemativs

Superlamily Cocevidea Fullén, 184,

Family Diaspididae Targion) Tozzetti. 168.
Sublamily Diaspidinae Targioni Vozzeite, 168.
Tribe Lepidosaphini Shimer, |68.

Most peniera of the family Diaspididae or armoured
scules are included jn the Wo subfitmilies Aspidiotinae
and Diaspidimae. ‘The subfamily Aspidiotinae, based
oo characters of the adult temiule, contains genera with
peetinac or plates and lohes that ure never buobed. fe
the subfamily Dinspidinae the plates are replaced by
aland spines and (he lobes anterior (o the median lobes
are often bilobed. The ODiaspidinge are usually
subdivided inte (he tribes Dtaspidint and
Lepidosaphini. Major characters of the Lepidosaphim,
mostly defined by Takagi (1969) and never found ip
the Diaspidini, include mepaducis. a pair of gland
spines between the median lobes and abdominul
segments HEY with either lateral tubercles ur spurs.
One op tore of these chameters may be absent.

NEW GENUS GF AUSTRALIAN ANDASPLIDINA Ih

In the present work the Subtribe Andaspidina 1s
recognised and cao be separated from the Iwo alter
subiribes al the tribe Lepidosaphin by the following
Key achipted from Balachowsky (96%)

Sume genera ind species assigned ta the tribe
Lepidosaphini are difficult to place in any of the
subtribes. Merverspis calligent Borchsenius. lor
Instunee, ducks lobes une gland spines but possesses
ineguduets. The species i) nevertheless related to other
species of Mereenuspis Gomez-Menor possessing glint
spines and well-developed Or reduced lobes (Danzig
1993). Phaulonytihis Levnardi, an Australian genes.
has small conical lobes. lacks gland spines hut
PHssesses Megaducts. 1 was included in the subtribe
Leptdosaphina by Borchsenius (1966). Another
Aastrilian genus, a/laironwrilas Leonardi, has small
Inumgular lobes bul licks megauducts. Borchseniiys
(1966) jmeluded this genous in the subtribe
Coceomyulina. According lo Takayi (1992), Mitulaypis
MacGillivray, with more or less triangular lobes, is
a primitive genus of the tribe Lepidesaphini, probably
of the subtribe Coccomytilina. Howardia Berlese &
Leonardi also belongs to the tribe Lepidesaphini but
its position remains obscure. The yenus possesses
incdian lobes similar to those of Anedaspiv. Eaeh
median lobe af Howard has a narrow, tninsverse
puraphysis at each basal corner and. in addition. a large
club-shaped sclerosis urising froor the inner basal
corner, Although Takagi (1492) tentatively included
Hewerdia in the subtribe Coccomytilina, the name
Howardiing Borchsenius is available for it but this
subthibe was creeted originally to include other genera
alsi. presently inthe tribe Diaspidini. [the follow ing
key to Subtribes. amily those genera. possessing well-
develuped median lobes in the adult female are
included, omitting the genus Howeredia lor the present,
The correct assignment of muny genera must await
more delailed research possibly of first and seeond
instr nymphs,

Key to subtribes of the tribe Lepidosaphini with
well-developed median lobes (adult females}

1. Meditin lobes with parallel or subparatlel sides, euch lohe
either without notches or with a single ouler noich, Dorsul
Math ital Aewucduels on the pygidiwm present arubsent,2

Meditn lobes now wirh pandlel sides. each lobe with ager
WANN staBAL, diverging slightly curving round toi long
oblique ouler marin, the naitgin ether smooth or serrated.
Dorsal mirginal megadiets on the pypidiiay either present
on ahsent
Andaspiding Balaehowstey
megaulucts always present on the
Ton euch side, .
A epidasaphina Shimer
Doral, ra meaduels always absent front pygicie
ra _-Cocceamytiling Birelsenius

Dorsal marginal
pygidtum, numbering 2

Genus Notandaspis gen. nov

Type species: Mythaspiy (Coecommtilin) frynrenaniire
rae Green

Dienst
Adult female on microscope slide clongate aval.

scementation of thorax and prepygidial segments
distinet. Spiracles with quinguelocular pores. Anrennae
each usually with 3 long setac. Pygidium rounded with
mediin lobes proniment, sct close together. teiamygular
or oval, inner edges short and diverging, outer edges
lofi, Second. third and fourtt lobes small, represented
by sclerotised pornts, Megaduets absent. Muacroducts
of pygidium, including muacginal duets. all ubout same
size. Gland spines short between median lobes;
anteriorly about saine length as thedian lobes. Venter
with microduects and glind tubercles present as far
forward as head.

Discussion

This genus is erected for the type species described
fron) Vietaria ulda pew species Irom South Australia,
In Jacking megaducts and possessing dorsal pygidial
macroducts all about the same size. the new genus is
related to Sualamaspis, a anomalous genus without
gland spines in the adult female but with all the other
characters of the subtribe Andaspidina,

Fivonvey

The name Noniiduspis is based on the Greek word
Hetos, Theaning south, combined with the present
eeneric name Andaspis,

The new genuy Neraredaspiy can be separaled trom
other genera of the subtribe by the following key:

Key to genera of the sabtribe Andaspidina
(adult females)

|. Pyeidiumpalways with +7 dorsal marginal megaduers on
each side, these much larger than other dorsal ducty
Pygidiamn always without dursal marginal megsducts, any
marginal ducts present always bout sane Seas other
desu) ditets . 28. of +

2. Mediundobes euch with sinalis mish un Ober margin. Anal
opening situated towards upex of pygidiurn

; Cufa Williaris

Meuiiin lobes ‘each with Luter Marvin smooth Or finely

serrated, Anal APO situated lowards base of pyvidiun

i, Gland tubercles present on ventral suriace of head
Purunduspis Mamet
Gland tubercles absent trom ventral surface of head
Andaspis MaoGillivtay

) Dorsal duets oF pygidium. tncliding any moruinal pypidirl
ducts, always in the farm of microduers’ only
Metandiispis Williains

Dorsal dacts of pyetdiun notin the form of mictaducts
diways inthe formeot macroduicts and all about sane size
5

UkO OL, WILLIAMS & 1, M. BROOKES

S Gli spines absent fram pygidial margins. .—,
Santonuispny Baldehowsky
Gihind spines present on pyeidil murine
ANeneidaapiy Williams & Brookes gem nov,

Nontndaspis hvmenantherae (Greeny vamb. ney.
(PIG, |)

Myrilaspis (Coccomytilus) hymenaniherae Green 05:
5. Lectotype &, Victoria. Myrniong, on stems and
wwies Ol Avmenantheru hanksii (BMNIL) (here
designated) fexanniied |,

Lepidesaphes iymenantherae (Green), Sanders 906:
VW.

Coeconytlas hymenantherae (Green). MacGilliy ney
2b 293,

Andaspis liymenadntherde (Green), Borchsenius 66:
7\

Adult fentale

Seale deseribed orivinally as teddish-brown, more
ur tess covered by fibres of the bark pan which it rests!

Adult female on microscope slide clongalte-oval,
about }.8anin long and 11 mm wide, widest at about
first abdominal seyment; body membranous to lightly
selerotised, pygidium moderately sclerorised.
Abdominal segitients strongly lobed literally. Lateral
spursabsent, Anterior spiracles each with a group ol
47 quinguelocular pores; posterior spiracles each with
2 or 3 quinguclovular pores. Antennae cach wath 4
setuc all about Same length.

Pyzidium rounded. Median lobes prominent, set
close together, almost triangular, cach with rounded
upex. outer edge fincly serrated and longer than inner
edge: ashort, blunt paraphysis arising from umer and
outer basal angles. Second, third and fourth lobes
represented by short, sclerotised projections, Gland
spines minute and barely perceptible belween median
lobes: a short pair present between cach median und
second lobe and groups of three gland spines about
us long as median lobes present between cuch second
and third lobe and euch third and fourth lobe. Anal
opening sittared towards base of pygidium. Vulva
present near middle of pygidiuin. Pertvulvar pores
ubsent, Dorsal ducts of pygidiuin all about sume size,
ravh about 20 jam long, arranged in loose marginal bo
submedian groups onmeach segment, Other dorsal ducts
on abdomen ubout same size as pyzidial ducts, preseot
around margins and in submedial groups of 6-10 An
sexment V, subjedial groups of 4-9 on segment TV
and usually submedial groups ob b3 ducts on segment
HH) Duets around margins becominay progressively
srnaller to miesothoras,

Ventral surtiee With oncginal gland spites as far
ferward as abdontinal sewmenct IW. Gland tuberelos
present on thorax und first abdominal segment
Submarginal microducts present on prothurinx

mesotlras and lateral lobes of abdominal seenents
Small ducts situated on anurgins of thorax aud tirse
abdomingl segment.

Pragiosis

The presenee of alinost thiangular median lobes on
(he pyadriniasa good distinguishing character ot this
species. Each outer edge of a median lobe ts, neverthe-
less, longer than the inner edge.

The lectolype designated ts one ol six specimens on
a single slide labelled ‘Myrilaspis hymenanidirn
Green, Type. from Mymenanitera dentatit. Victoria.
Austedia, coll, J Lidgett No, 63° and is clearly marked
in-red tok. Tas lurther located on a diagram showing,
the posinons of all six specimens or a separate label
fixed to the baek of the slide. The other five specimens
are here desivmated puralectoypes (BMNE).

Notandaypiy pidnadattae sy). wey,
(PG. 2)

Marerial examined

Hololype, @, ANIC, South Australia, 70) kn west
of Oodnudulta, on stems ol dcucia uneura, | s1970,
FD. Morgan.

Paralypes: suine daktas Holotype & SO (ANIC).
So 9 (BMNH).

Aduli female

Seale dull white, 4 mm long, exuviae upreul, pale
white, cork layer of phint in some iiatances growing
in strands over scale caver,

Adult female on microscope slide, elongate oval,
largest available specimen 3,2 mim long, 1.2 min wade,
widest at metathorax. moderately aclerotised
throughout, pale brown, pygidium light brawn
segments well constricted behind head and prothurax
und hetween thenicic and prepygidial segnrents
Anterior spiracles euch wilh uw group of tht
quinquelocular pores. posterior spiracles each with |
ur 2 quingueloculin pores, oevusionally absent
Antennae cuch with 3 setae. one thicker ind longer
than others,

Pygidium rounded, Median lobes. prominent. each
almost oval, the short Inner edge and long Outer edge
linely serrated, A pair of slender puraphyses present,
each arising from inner und outer basal angles, directed
antero-medialy or almost transversely but not meeting.
Second, third und fourth lobes represented by soll
svlerotised points. Gland spines short and minute
between median lobes, a subeqodal pair present between
cuch median und second lohe, a group of three, all
about us longas median lobes, situated helween each
second gnd third lobe, Anal opening lying near middle
of pygidium. Vulyie sittated anterior to position of anal
opening, at dbout one third length al pygidium from
buse. Pervulvar pores ubsent. Dorsal duets ob pyran

NEW GENUS OF AUSTRALIAN ANDASPIDINA 187

Fig. 1. Notandaspis hymenantherae (Green) comb. nov, A. Adult female, general aspect. B. Pygidium. C. Dorsal margin
of pygidium. D. Ventral margin of pygidium, E. Antenna. F. Anterior spiracle.

188 D. J. WILLIAMS & H. M. BROOKES

Notandaspis oodnadattae sp. nov. A. Adult female, general aspect. B. Pygidium. C, Dorsal margin of pygidium,

Fig. 2.
D, Ventral margin of pygidium, E. Antenna. F. Anterior spiracle.

NEW GENUS OF AUSTRALIAN ANDASPIDINA isu

all about same size, cach approximately 20 am long,
numerous ulong margins and arranged in ill-defined
rows to middle of pygidjum except on segments IL-V
where they form distinct submarginal raws and
submedian groups of 710, Ducts around thargins
becoming, progressively smaller anteriorly as far
forward as mesothoray.

Ventral surface wilh submarginal microducts of two
types. An elongate type. each about 15 pm long, present
in submurginal groups on abdominal segments FV and
V. A shorter type, each about 10 aim long and with
area surrounding opening selerotised; presen! in
marginal groups on head, thoracic segments and
second ubdonunal segment, and others present instill
groups near libium and medial area of head. Gland
spines present in groups on prepygidial margins and
minute, Weuineare gland tubercles present subijarginally
on prothorax and neur inner edges of groups of
microducts.

Diagnasis

This 18 a lurge species compared with others in the
subtribe Andaspidina with the scale cover reaching 4
tam long and the adult female 3 mm long, The scale
cover of most other species searcely exceeds 2 mn long
and the adult female is rarely more than | mm long.
At first sight the seule of NV. oodnadatee resembles
an ovisae of many species of FErlacacens
(Eriocoecidae), Although each of (he median lobes is
almost oval there is a distinct, short inner edge and
a long outer edge as in all species of the subtribe. The
shape of the median lobes distinguishes the species
from N, hynenaitherac which possesses almost
(ruingular median lobes, The positions of the anal
opening and vulva are reversed ja both species, the
anal opening of N. evdnadattae lying posterior lo the
position of the vulva and mV, Aveenantherae the anal
opening lying anterior w (he position of the vulva.

Etymology
The name ts based on the place namie ‘Oodnadauue.

References

Bat acniowsaky. A. S. (1954) “Les cochenilles. paleareniques
de fa tribu des Diuypidini” (institut Pasteur, Paris).

(68) Sur une apuvelle sous-tribu de Lepidesaplicdini
(Coeemdes-Diaspididae) cereéee par ly découverte d'un
nouveau vere nuisible wu Culéier @Arabie ly Sao-lome,
Rew. Zivk Bor. Afr TS. 54-63.

(1973) Nouveau nam de genre ee un Diaspididae
de Suo-Tomeé. Ball, Soe. ent. Fy 78, 2

BorRCHSENIIS, NOS. (MG) YA catalagye vm the armoured
scale inseets (Diaspidoides) of the world”. (Nauka, Moseoyy,
Leningrad),

(967) Materials on the fauna of seale insects
(Homoptera, Coecoidea) from India, Ub Andespiy Maece
with (three new allied genera (Didaspididuc) Ln, Oborr
46. 724734

BrimaiecomBe, AOR. (960) Studies of the Cocooiden (0,
New species of Diaspididae, Qe. A. agree. Sei, 16, 381-407,

Danaic, E, M. (1964) Ga the types af species deseribed hy
NS. Borchsenius in the article “Materials on the Runa
Of scule inseets (Homoptera, Covcoideu) fromy india, £1.
Ent Obvcr Ah, 8434.

—____ ( (993) Fauna of Russia and neighbouring countries,
Rhynchota Volume X, Sele Insects (Coccinea) Farnlies
Phoentoococerdie and Diaspididae. Hawa Russia ONS) 4,
452

GHavRt. M, 8. K. (1962) “The morphology and taxonomy
of male seule dascets (Homupleru: Coeconlea)” (British
Museum |Natural History|, Landon).

Green, F, BF. (1905) Some new Victorian Covcidae
Fieiorian Nat, 22, 3-8,

Wank, W, J, 946) On (he Euiiopian Diaspidini. Trans, K,
wat, Sow Lame, 97, 497-502,

MacGiniiveay, A,B. (921) “The Coceidae. Tables Jor the
identification of the subtamilies and some of the more
IMporlant penera and specics together wilh Uiscusstons ot
their anatomy and lite history’ (Scarab Company, Urbana,
lings).

Mamet, 1 R, (1967) New wener and species of Coccmden
from the Masvurene Eslunds (Homoptera). Mauris Dest,
Bull. 6, 89-102

Rao Vo Po& Perris, G. & (952) The genus Aneaspris
MacGillivray (insectt: Hormeptera: Coccoiden),
Micrventumatauy U7. \7-32.

Sanpers, J G, (1906) Catalogue «af recently described
Cocewae, Tech. Sen Bur Ent, U8. W218

Tawagl, S. (1970) Diaspididae of Taiwan biased on material
collected (7) conaeection with the JapunelS. cosopenilive
selence programune, 1965 (Homoptera: Coveoides) Part U1.
Jnyeent matsum, 33, 1-146,

(1992) Mitulaypis and Scluperaspis: their distributions
and (axanomie positions (Homoptera; Cavcoideu
Diaspididie) /bid, 47, 34-90.

—__ & KaAwal, §. (1986) Some Diaspididae of Japan
(Homoptera; Coccoideas, (hid, 28, 93-119,

Wil tidms, D. 4, 1963) Synoptic revisions of |. Lindinyespis
and [1], Anduspis with pwo new allied gener (Hemiptera,
Coceowea.) Ball Br Mus. nat, Hist Eni WS. 31.

(980) Aadesply dist Williams idenieal with ol.
mumerad Brimblecombe (Wemiprera: Diaspididaey, a
species found on toy and absecnted with the fungus
Seprobasidivn. Ball, ent, Res. 70, 259-260
—— & Wasson, G. W, (1988) “The scale insects of the
~ tropical South Pacific Kegion Part L The armoured scales
(C.A,B. Imernational, Wallingford, ),

DIET OF JUVENILE KING GEORGE WHITING
SILLAGINODES PUNCTATA (PISCES: SILLAGINIDAE) IN THE
BARKER INLET — PORT RIVER ESTUARY, SOUTH
AUSTRALIA

By Rop M. CONNOLLY*

Summary

Connolly, R. M. (1995) Diet of juvenile King George whiting, Sillaginodes punctata,
in the Barker Inlet — Port River estuary, South Australia. Trans. R. Soc. S. Aust.
119(4), 191-198, 30 November, 1995.

The diet of juvenile King George whiting, Sillaginodes punctata (Cuvier &
Valenciennes), was determined by examining the stomach contents of fish collected
over two years from shallow eelgrass and unvegetated habitats in the Barker Inlet —
Port River estuary, South Australia. Estimates of weight of prey actually ingested by
fish were made by combining abundances and sizes of prey found in stomachs with
data on the size — weight relationship of potential prey items collected separately. Fish
ate epifaunal invertebrates exclusively. A range of crustaceans formed the main prey,
with smaller fish taking mostly harpacticoid copepods. Amphipods were more
prominent in the diet of larger fish, which also fed upon polychaete worms. Fish fed
mainly during the day. Fish collected at night typically had very little food in their
stomachs, as measured by a fullness index (ratio of estimated ash-free dry weight of
ingested prey to dry weight of fish). Relatively few fish were caught over unvegetated
habitat, but where comparisons could be made, polychaetes rather than crustaceans
predominated.

Key Words: Sillaginodes punctata, fish diet, predation, crustacea, seagrass, Zostera.

Transaenons af the Royal Sacre af &

Wish W995, TAL. [0] tum

DIET OF JUVENILE KRING GEORGE WHITING SILLAGINODES
PUNCTATA (PISCES: SILLAGINIDAE) IN THE

BARKER INLET - PORT RIVER ESTUARY, SOUTH AUSTRALIA

by Rob M. CONNOLLY*

Sumanary

Cossoriy, RM (1995) Diet ol juvenile King George whiting Sillveimedes panei, inthe Barker Liley - Port
River estuary, South Australian, frase, RoSoe Stash W904), IY ETM JO Novernber, 1995-

The dit ul pivenile King George whiting. Siwides plana (Cuvier & VYulencienies), was determined
by examining the stonieh contents of fish valleeted over Uwerveun Cron) shallow eelgrass and unvegetitod habyals
Wthe Barker Tiler ParORiver estiivry, South Australi. Ustiiiates ob weitt of prey actually ingested by fish
were made by combing abirtlinees anil sives ol prev foul mpstoniachs with dan on the size - weight relationship
ul potential prey items cullecied separately, Bish ale epifiunal irvertebrates exclusively, A range of crustucedns
formed the man prey, WI sriatler Gal taking mostly harpacheoid copepods. Aniphipods were more prominent
ihe det yr larger Fish. whith alvo fed upon polychaere worniss Wish ted mainly during the day, Mish collected
a{ might typically had very Hitle food in their stomuchs, as mewsorcd by a lallness inde (ratio of estimated ssh-
tree div weight al ingested prey ta diy weight ot fishy. Relatively few fish were cought over unvegelted habitat,

Bit where cormpurisons could be rude. polyelietes nulier than erustieeins preduniimied.

REY Worbs Slawinedes panei Web diet, predanon, cvustucea. scudrass, Zagter,

futraduction

Fish from shallow, soft-substratum habitats are
lypicully carnivorous, preying mainly on small, moule
invertebrates, Invertebrates associated with the seagrass
vunopy Hr sediment sorties (epiaini) are more
Woportint than invertebrates from wathin sediment
(inhuuing) (KTanpp er ad, 1988), Despite the high levels
ol primary production sustained by shallow seagrass
meadows (Hillman er af. 1989). lew fish aerually
COOSUING scaghitss I Temperate walters (Klan ed al.
LORY),

The diets of small tish trom seaziass habitats fA

Soully Australia haye not heen reported. The diets of

lish in Western Port, Victoria, an enclosed waterway
with vegetation similar to that in the Barker Lilet Port
River region, have been studied by Robertson (1984)
ahd Bdgar er a/, 1993)! who confirm (he importance
of epifaunal invertebrates, especially crustaceans, The
diet ol juvenile King George whiling (Sillieinodes
puneraia) in Western Port is deseribed i Robertson
W977). Fish oof tis spectes: fed ain erustaeeans
(harpacucoul copepods, mysids and amphipods) ufter
setting from a planktonic larval stage into eelgrass
beds. Larger juveniles (>40 mm length) fed upen
ehost prawn (Calliamasse) larvae and polychaetes.

* Department of Zoology, University af Adefuide. S. Aust
S005, Present address: Raiculty of Enviranmental Sciences,
Griftith Universiy Qld 41

Engak, GJ. HAMMOND, LS. kc WialrsoN, GF rgb)
Consequences for commercial fisheries of loss of seayeriten
beds if southern Australia. Report ty Fishing Industry
Research & Development Committee (unpubl)

primarily in vovepetated patches adjacent to eelgrass.

A wide variety of measures and indices involving
£ul upalyses his been used i attempts to quantify the
relative Tmportince of fvod categories to Tish (Berg
9792 Hyslop 1980). Mrequeney of oceurrence,
abundance. weight and volume have been used butany
one of these may he misleading (Bere 1979). Indives
COMPS TH Varies Ways the basic measures listed
above have been devised (e.g, Pinkas ef al, 1971) but
no index is advantugeous in all situaricns. Different
oud Culegories gui prominence depending on the
Weighting piven to the different variables in the index.
Bere (1979) recommends thal where an index
combining abundance, weight or volume, and perhaps
frequeney of oeeurrence, is used, values for the
separate varubles should also be shown. | consider
that it may be as informative to forego the index, given
that it is influenced by the weighting given to each
variable. and simply present results based on. for
example, abundances and weights,

Decisions about the importanve Ob food categories
lo lish are best bused not on the weight or volume of
prey remaining inthe gut but on the weight or volume
of prey ingested. The ideal way of calculating weight
or voluine Of food intake for prey such as motile
inverlebrales is lo determine the weight - size (e.g.
length) relationship lor all prey categories using whole
unimals. and then to estimare the weight or valume
of rngested prey based on the number and size of
individual items found in fish, Edgar et al. (1993)!
deseribe un upproximate method for estimating weights
in Which prey items are allocated to asize-class known
lo pepresent the runge of sizes retained on it particular

192 kM

siewe (West size Within a siiek OF |ieratehieally
arninged sieves, The size of invertebrates is then used
Ie esuimnute then weight (Edgar 1990),

In calculating the relative importance Of food types
by. dividing the number or weight of i food type by
the number or weight of all food ina particular fish,
no distinction is made between a fish having in its
stomach one harpacticoid copepod and one calanoid
vopepot and a fish having 50 harpacticoids aid 50
calaneids, Many studies therefore include some
estimate of gut fullness, The most common method
has heen to assizn puts to one ol several subjective
vategories of fullness (Berg 1979). A measure more
repeatable by other workers ts (' indice de repletion
of Hureau (1969, described by Berg 1979) in which the
weight of ingested food is presented as i proportion
of the total weight of the fish,

The primary aim of the present study was to record
the dhet of juvenile 8. panetee m the Burker Inlet -
Port River estuary. This estuary has been declared an
uquaie reserve im recognition of its jntportance in
providing, habital for juvenile fish, especially S$.
punciaia Gones 1984), the most important species
veonomicaully in both the conmmercial and teereational
fisheries of South Austrafia. A secondary aint was to
compare diets of fish fron eelgrass (Zostera muellert)
and unvegeluled habitats,

Materials and Methods

Juvenile Silaginedes punctaii were collected over
two years during surveys comparing the fish fauna of
eelgrass and tnyeyetated habitats im the Barker Inlet
- Port River region (1387 30° FB. 34° 48° S) (for
deseriptions of the estuary and the Surveys, see
Connolly 19940), At each sampling period, the
stomachs and oesophagi ofall fish (or of 10 randomly
selected tish where more than JO fish were caughy
from cach site were removed, and the contents
examingd. The number of sites and total number of
lish examined ateach period are shown in Table |.

The most satisfactory way of determining whut fish
fal is Lo esurmine items only from the anterior part of
the trace This is because food ilems from (he
oesophagus and stomach are more Likely to be inter
and are more easily recognised than items further along
(he gut, the blas caused by differential gur passage rates
or digestion rates of different food items is likely to
be reduced (Berg (979) aud items towards the anterior
end of (ire tract give a more reliable guide to the diet
of fish just prior (o capture. This is an advantage for
the secondary ain of this study. namely comparison
of the diet of fish caught over eelgrass and unvegetated
habitat. The truct af 8. punerete less than about 25 min
jong isa simple, uncoiled tube, narrowing posteriorly:
contents of these smaller fish were examined from the
sechion anterior lo the marrowing.

CONNOLLY

Tair 4 Sillaginades punctata evaniied for stomach
COMA (uiniber ubsites ac which So Punelibe were canels,
number uf fish esandnedd, wed nivelion tener of fish separa
for cach habitat ar cach sampling pertod.

All iste were caleered citrine the eevmine exconT these dnarked
Nigh

Habis: Fo = eelgriss (7 = Unvevetated,

Sampling period Habitat Nuuiber Nuipher ot Malian

of sites fish examined length Gand
January 90 E 8 tie RU
ul 2 tl (a
April 1990 b g 48 1
i ) | 1)
August 1990) be i 45 44
t Z 2. ah)
Outaher (W490 iD K 48 nS
L | 7 AS
February 194] b K VW 4
v ‘ ih Wn
June 194] hr 4 Ad 4
Uv 4 7? a4
Tine WU) Night id 7 a4 y
U | 4 ya
Crther (94 k x fv 4
v 4 0 1y
Oel. OL Night t J a) th
u | | 7

Individual ems were mostly cither intact or nearly
so, und were identified to major taxa and counted
Animals were measured using a grattcule in the
microscope eye-piece and were assigned to wu sieve
mesh site-class so that weights of ingested prey could
be estimated usimy the length - weight relationship
deseribed by Edgar (990). The miajority of prey items
were crustaceans and where individuals were ngt
whole, sizes were estimated by roughly piccing logether
parts of un animal (in (he case of large crustaceans such
as amphipods and mystds) or by using other individuals
of the same taxon as 4 guide (for copepods), The only
tuxa recorded other than erustacea were polychaetes
dnd chironomid lurve. Chironomid larvae were rare
and Were always whole Although polychaetes were
olten whole, they were soniclinies im pieces; estimating
sizes Of polychaetes chopped jnto pieces was the most
problematic part of this method. In these cases the
number of anterior ends was counted and lengths were
estimated to ry to lake account of the general size of
individuals.

Euch prey item was assigned to a size-category
reluling to the range of lengthy Of thal taxon retained
oo different mesh sizes. These size ranges were

DIE OQ) WUVENILE KING GEORGE WHITING 103

determined. by measuring the length ol numerous
speeimens of eaeh taxon from epifaunal samples taken
ithe time of sh collections. Por cach taxon, relative
length - frequency histogrants were plotted for each
imesh size, and a range of lengths wus chosen as
representative of a imesh size by seleeting upper and
lower lintiis where bistogrants from adjacent mesh
sizes crossed, Size ranges for each taxon are shown
in Vuble 2, The meun ash-free dry weight (AP DW)
of invertebrates can be related to sieve mesh size using
Halgur’s (990) equation, log B= a + blog § (where
B= AFDW (mg), S = Sieve Size (mmm) and a and
b vary depending on broad taxonomic category}, Since
each sieve size retains animals ranging fron thal sieve
size lo the Heal. 5S is eXpressed asa geametne ntean
calculated using the equation. log S = (log 8, + Jog
$),))/2, in whieh S$) = mesh size of the 1 sieve and
$,,) = mesh size of the next size up |bdgar 190),

For each fish, the percentage abundance of each food
category was culculuted as n/N. where nis the number
of individuals of the foud category and Noa the total
nomiber of id) viduiads of albeategories in that fish, The
same calculation was made lor each category based
on estimated weight (AF DW). The average percentage
ubundunce and weight af euch food culegory were
culeulated for cach site. The average percentaze

TABLE 2. Sige ranges fran of prey types matching mest
AIZEN,
Blank wells indicate that prey type was not found on that mesh,

Prey Mesh size (mi) -
type 4 2 I OS (has QA25 0.075
Hary ails SUORUR PSS AOR ecu SY
Mau Shh <()h

Cyl SHH9 WSS4.09 ocIS9
Cala a! Wil <All?

Ost >In UML AAU <i

Ample ole DSU tS i el

(apr >t Vat cas

Myst as A a

Tana e3 Sled lita <2

Chr Shs 4-64 <4

Gin 245 (4a <b

Poly oT) eS 2248

2) AT Ube allt

(hae sdk Sthdd [2th <)2

utundance and weight at cach sampling period (with
might callections in June and October 1991 treated as
sapling periods) were then culeulated sepurately for
eelgrass und unyesetiled sites.

The total estintaled weight (APDW. my) of the
stomach eootents of a fish was recorded as a proportiean
of the estimated. total weight of the fish (Ury weight,
2), This proportion gives the same information us
Hureau’s (1969, deserihed by Bera 1979) /idice de
repletion, although Hureau’s index used the same units
in numerator and denominator and is reported as a
percentage, By using meas the unt lor the numerator,
the ratio (fullness. index) used here prinintises. the
oecurrence of numbers less than one, The weight of
the whole fish. including stomach cantents. was
cstimuted using the relavionship heiveen dry weight
and fish length, Fifty S$) panera collected trom
different periods and retaging fron 18 to 133 nny total
lengih were weighed aller being dried (o constant
Weight (at toast 48 h) an 60°C. The dry weight of a
lish is best estimated by: its length using the relationship
jog W = 3.26] jog L - 6,396, where W is dry weight
(zg) and Lis total length (mim) (re = 0.997)

Fullness indices of ish Trom eelgrass and
unvepctited sites were compared using Mann-Whitney
U-tests at periods when Tish were collected from
cough unvegetated sites to make Useful comparisons,
Indives were ulso compared tor fish from celyrass sites
between day and night samplings atJune-and October
199], The Mann-Whitney U-test is less powerful than
atest if data neet the assumptions of normality and
homoscedasticily but, in cases such as these where
sumple sizes are very small and tests of normality are
possible. itis a more reliable method of testi
differences in central tendencies (here, medians),

Results

The diet of juvenile Silaginodes punetara consisted
entirely of javertébrates. Thirteen categories were
recorded, us shown in Table 3. Prey were either
crustaceuns or polychuetes, except lor a stall number
of cinrooomid larvae taken in October 1990, Porvellid
harpacticoids and cupreilid amphipods were counted
separately from their general taxa (harpacticoids anc
amphipods, respectively) because of their different
form. Porcelll harpacticoids have a wide, flattened,
shield-like shape and capretlids are extremely long and
thin, compared to gamimucid amphipods. The
prominence of small venis such as copepods, especially
harpaecticoids, was greater when bused on abundanee
than when based on weight: The prominence of larger
items S50ch us alphipods and polychaetes was.
conversely. more obyiwus when based on weights
Nonvithstanding these different emphases. the change

194 R, M. CONNOLLY

Taniy 3, Stomach contents of Sillaginodes punctata bayed on abundances.

Huahitatss E = eelgrass, U = Unvegetated. Numbers are mean percentage of food category from all siles, with standarl
elrors (SE) in parentheses. * = no SE because fish were caught at only one unvegetated site. ** = no SE because only
polychaetes were recorded from fish at unvegetated sites. Food category abbreviations are as follows: Harp = Copepoda -
Harpacticoida; Pore = Copepoda - Harpacticoida - Porcellidiidae; Cycl = Copepoda - Cyclapoida; Cala = Copepoda -
Calanoida: Ostr = Ostracoda; Amph = Amphipoda Gammaroidea: Capr = Amphipoda - Caprellidae: Mysi = Mysidacea:

Tana = Tanaidacea: Cuma = Cumacea: Cari = Caridea; Poly = Polychaeta, Chir = Chironomidae, larvae

Sampling Habitat Harp Pore Cycl Cala Ostr Amph Capr Mysi Tana Cuma Cari Poly Chir
period
January E 7 24 2.5 7 60
(940) (4.5) (0) (2.5) (4) (11.5)
U 13 3 83
(13) (3) (17.5)
April E Al 14 55
1990 (12.5) (7.5) (14.5)
U 100
iF)
August i 87 ! 3 0.5 0.4 T i 07
1990 (3) (0.6) (1) (0.5) (0.4) (2,7) (1.1) (0.7)
U 83 17
(3) (3)
October 69 0.1 5.1 21 1.5 1.2 23
1990 (8) (O11) (2.2) (6) (1.2) (1.9) (2.3)
U 3 38 1.3 2 20
(12) (3,8) (0,9) (1.5) (bh
February E 14 95 63 2.5 68
199] (1) (4.5) (5.6) (2.5) Ol)
U 13 99
(1.3) (1,3)
June 199] 92 11 3 3.5
G) (Ll) (3) (1.5)
U 78 4 25 O05 27 15
(1) (14) (2.5) (05) 17 (9)
June 199] E 82 7 iH
Night (14) (7) (7)
U 100
(*)
October E 74 2.) 1.2 20 2 OL I
199] (5) (2.1) (0.8) (6) (D) (0,1) (0.5)
U 72 or 12 1 il
(14) (4) (4) (1) (il)
October 1991 E 59 31.5 8 1.6
Night (29.5) (27) (4) (1,6)
u &7 13
(*) (*)

in diet of S. punctata as fish grew larger is shown
clearly in Tables 3 and 4. The median length of fish
at each period is reported in Table 1.

At sampling periods in the second half of the year
(August, October 1990; June, October 1991), when fish
were small, harpacticoid copepods were the most
conspicuous prey by abundance, and amphipods along
with harpacticoids were dominant by weight. The
abundance and weight of amphipods were noticeably

lower in June 199] than at later periods and this may
be attributable to the smaller size of fish at this period.
Cyclopoid and calanoid copepods, which are typically
more planktonic than harpacticoids, were taken
consistently at these periods but were small
contributors to diet by abundance or weight. Ostracods,
caprellid amphipods, mysids, tanaids and polychaetes
occurred occasionally but were not important by
abundance or weight. Cumaceans were recorded in

DIET OF JUVENILE KING GEORGE WHITING

195

Tasin 4, Stomach conrems of Sillaginodes punctata based on weiehy.

All labels and symbols as for Table 3,

Sampling Habitat Harp Pore Cycl Cala Ostr Amph Capr Mysi Tana Cuma Cari Poly Chir
period
January E 6 ll 2.5 i 69
1990 (4) (5) (2.5) (7) (10)
U 4 10 87
(4) (10) (13)
April E 22 14 64
1990 (ly (8) (13)
U 100
(er)
August E SY 3 6 07 0.9 25 4.6 Ol
1990 (7) (2) (2.77) (0.7) (0.9) (8.6) (4.6) (0.1)
U 58 42
(20) (20)
October E 38 0.1 10 37 7 4.5 31
1990 (8) (Ol) (4) (9) (4.5) (2.9) G1)
uU 54 3.1 3.9 iT] 29
(8) (.1) (2,5) (6.2) (1)
February E 10 8.2 2.8 2.5 76
199] (8.5) (4) (2.7) (2.5) (8)
u 0.3 100
(0,3) (0.3)
June 199] E 85 | 5.5 8
(6) (hij) (5.5) (3)
uU 68 L4 5.5 28 (6) (17)
(16) (4) (5.5) (2.8) (5) (10)
June 199] E 76 7 17
Night (16) (7) (i)
U 100
(*)
October BE 50 + 3 33 7 LS 2.
199] (7) (4) (2) (1) (4.5) (1.5) (2)
U 66 5.5 ll 3 15
(13) (5,5) (4) (3) (15)
October 199L E 37 25 27 10
Night (20) (14) (15) (10)
U 48 52
() (*)

small numbers in October 199].

At sampling periods in the first half of the year
(January, April 1990; February 1991), when larger fish
were examined, polychaetes were the main food
category by abundance and weight. There was an
obvious difference in the size of polychaetes taken by
fish in periods in the second half of the year compared
with the first half of the year. Polychaetes taken in June
1991 were small, ranging from less than 1 mm to 3
mim long (although the larger of these are large relative
to other prey). Polychaetes taken in October 1990 and

1991 were 2 - 10 mm, Polychaetes in fish from periods
in the first half of the year ranged from 7 - 50 mm
in length. Amphipods were the second most important
category by abundance, although by weight amphipods
were no more prominent than the other two frequently-
recorded categories, mysids and tanaids. Harpacticoids
were found ina small number of fish in January 1990.
Fish in which harpacticoids were found had no other
categories of prey present. so that although the
harpacticoids were not numerous and were small, they
comprised 100% of the food in those fish based, on

19% Rk, M. CONNOLLY

ubundance or weight, Curidean shrinips were recorded
infrequently im Picbruary 1991,

The percentage of fish having empty stomachs is
shown in Table 5, Very few of the fish caught during
the day had empty stomachs at any sampling period
and no difference is evident between sites from eelgrass
and tinvegetated habitat, The weight of stomach
contents, aS a proportion of total fish weight, vatied
markedly from site to site bul did not seem lo vary
consistently with season (Table 5). Tn periods when
fish were cauughtat enough unveyetuted sites to make
4 reasonable gompurison possible, the weight of
slomuch contents did not differ between eelgrass and
unveyelated habitats (Mann-Whilney U-test results:
October 1990, p = 0.234; February 199), p = 0,734,
June 1991, p = 0773; October 199), po = O74)
Results of tests were identical for all periods whether
or not fish willl emnpry stomachs were included,

DAN 5. Numbers ef fish with te fone lee steitenlt, ane
fullness indices, separately for cach habital ab each sampling
period,

Habis. E = eelgrass. LE = Unvegetated. The number of
fish with empty stomachs ts shown firstly by number (i) anel
secondly us a percentage (9%), bul percentages should be
interpreted cautiously for unvegetated habitat where fatal fish
fumbers are very small Fullfiens index is weit of stomach
contents (me APDW ) as proportion of woighl uf wholy fish
(oulry weight), Fullness indices are shown as means of sie
means, with standin errors in pareatheses (n/a = lish caught
it ong site only, therefore no SE available). ta} = excluding
fish wath empty stomachs (b) = including fish with empre
stomachs (and fullness index of zen>)-

Sumpling Habitat Empty Empty Pullness Fullness
period (nt) 1%) index fal jndes (bj
dianuary W9O EF 3 5 AAR USI) 4.01 (0.50)
L 0 it} ART (Rd) AKO 11 Rd)
April 00 k 1) () 2.44 (()44)
J i) a 204 (tal
August 90 de () iY AAT UHL
\ if) i 255 ith ln}
Outober WT 2 3 ATV {OTH 420 Ou)
\ 0 0 260 (30) LOO (36)
Febroary OL 7 KN) 4.23 (124) 46 (1 2A)
u 0 i 3,60 (006%) 3:00 (0.63)
June 9 [ f) ) B52 (073) 252 rs)
u | \4 AS (OKO) SAP Oa)
Tan 19) I: a | 114 (aT) ST CY
Night ul 2 50 OM6 july) OAR pay
Ovtober ULE 1 5 444 (84) 472 78)
ul () () 4.35 (76) 345 T6)
Weber WY) TE Is (od 299 (0.4L LO (2b
Night U u i] O98 tava) OUR (vad

More than hall the lish caught at might indune and
Oetober 1991 had empty stomachs. In fish caught at
night with food in their stomachs. the types of food
wert similar io those in fish caught during the cay,
Fora given period, the quantity of food tn fish caught
al night was significantly less than in fish caught duringy
the day When fish With enipty stomachs were iachided
(Mann-Whilney U-test results: June 1997, p = 0,047;
Ovlober 199) pp = 0.014), but was not significantly
different when fish with empty stomachs were excluded
(June 1991, pp = 0.186; October 1991, p = 0,221).

Comparisons of the dict ot fish caught over celgruss
and unyegeliled hibital are limited by the small nurmber
OFS. punrraia caught over invegetated habitat and the
small gumber of unvevetated sites al whieh lish were
caupht Over all periods, polychaetes seemed tiv
predominate iy fish frotn unvegetated habitat, Ih
October 1990 and 1991. when fish were cuueht at 4
unyegeluled sites. Increasing the vhance that the data
are representative ol the hubitat more yenerally. ooly
lish from uwevetated sites had taken polychaetes. Fish
from celerass sites tended (6 contain a preater Guage
Of crustaceans, Cuprellid amphipods, for example, were
recorded only from fish caught over eelgrass at both
periods,

Discussion

The diet ot Siflaginodes punerata fits within the
typical diet for fish ftom shallow, sott-substratuiy
habitats, Stomach contents at the periods sampled give
ny indication of feeding on anything ofher than motile
invertebrates, Juvenile Sv panera caught at periods
inthe first half of the your were large enough to be
able to take snl) individuals of other fish species but
there was no evidence of (his. Although gastropods are
culen by some fish species. none was tound in the
present study. The prominence of harpucticoids and
aniphipods in the diet of smaller juveniles and aa
increased prominence af polyehuctes. in older juveniles
matches the pattern Wd. punctate frou Western Port,
Vicloria (Robertson 1977),

Bruce (1995) has suggested that (he shift in diet
towards Jarger crustuceans and polychuetes with
in¢rousing fish size may be rcluted to the iming of gut
coiling. Bruce’s study of larval and postlarval &
puncrata trom South Australian waters shows {har
coiling ofthe gut tube and tigration of the anus begin
in fish 2b = 24 mm long, and are complete in fish of
26 inm. Most fish caught in October 1990 and 1991
during the present study were > 26 mm long and did
have called puts. These fish bad a predominance ot
smull crustaceans such as harpacticoids, however.
midicating that the shift towards larger crustaceans and
polychaetes does not happen uni] after gut coiling.

DIET UP JUVENILE KING GEORGE WILITING (97

Results sugvest that $ pierare feed on a narrower
ringe of prey umd include more polychaetes in iherr
Wich when over unvegetuted fabri, Lubbers er al.
W490) have also reperted that for juveniles of several
species Of [ish from an estuaty in Chesapeake Bay,
USA, diets of fish collected frum Unvegetated areas
ticluded a much ereater proportion of polychaetes than
diets oF fish collected from vegerted ures, Evidence
trom the present siudy is, however, obtained [rom only
i small qumber of fish from very few sites, The sinall
number of fish examined from unyegetaled sites could
account Jor the failure to find food types such as
tardean slirimps recorded infrequently tm fish trom
CCIANINS Sites,

Ividence from the ive night sampling. periods.

Sumeests that juvenile §, punctate feed manly durin
the diay. The stormnwehs of fish collected at night were
ofien vither empty or eentiuned only a snrall quantivy
ol loud. Either fish fecd in a limited way at night or
food tn the stomach of fish collected af night remarned
from feeding during daylight hours. The time between
sundowt und collection oF fish at night ranged from
fouria seven hours. The rare at which food is evacuated
by juvenile. Guritvorous. murine lish ofa similar size
0 the fist Siddivd bere haus heen shown to range
yatously from 2.7 lo 4.8 h (Rosenthal & Paflenhoter
1972), 6 bh (Archumbault & Feller 1991) and fram 10
jo 40 h fRyer & Boehlect 1983), These laboratory
estimates Of wit evacuation times, however. lend to be
overestimates |Loekwood 80). Food is presumably
clew of the stomach belore it is fully evacuated from
(he gut, so stomach empiying Himes could be shorier
than these mentioned aboye. On the other Hand. wut
paussape tiles wre much slower in colder water (Durbin
ard W83; Ryer & Boehler 1983) and, in the evening
watt Ieiiperatures of June and October 991 of about
14°C, food fay have remamed in guts much longer
li is therefore impossible to distinguish berween the
possibiliies of limited nocturnal feeding and food
remulining in stomachs from daytine feeding.

The ratio of ingested food to lotal fish weight did
not seen Ly vary consistently with the size of fish taken

al different periods. This contrasts wath the study of
silver hake (Merlncciuy bilinearis) aod Atlantic ead
(Gadus nora) by Durhin et af, (983). using the
sae measure, in which it was found that the ratio was
greater in larger fish, Durbin epa/ (983), however,
used a much larger size range, includiny juvenile and
adult fish. Diflerences in (ae ratio for 8. paren might
occur in larger fish,

Any differences in gut passage rates or rates of
digestion tor different food types-could have affected
the apparent relative importinee of lond types, These
biases were not determined during the present study
but should hive been liniited by examining, food onby
from the besophigos and stomach of fish. Dilterential
digestion rales tend to underestimate the importince
of solt-bodied invertebrates (Scholz era/ 1991) and,
lor juvenile S. prnicrette. this meus hut polychaetes
are the liaxon most likely to be anderestimited.

This study confirms that juvenile S$ punerare within
the Barker Inter - Port River estuacy feed on epifaunal
invertebrates. Experiments in the same estuary have
shown that removal of eelgrass canopy reduces
epiluunal invertebrate productivity (Connolly 1995)-
Abundances of juvenile A) punctate are not reduced
directly by removal of eelgrass canopy but are
correlaled with levels of jovertebrile productey ity
(Connolly (994b). The ongoing threat to the health of
eelgrass in the estuary from Huntin activities such os
treated sewage und stormwater discharge should
therelore he viewed as a polentilly detrimental
influence on S. pureuiea populations

Acknowledgments

1 thank Dr Alan Butler for fas belp and
encouragement, Barry Bruce for ideas about dietary
shift. and Drs Gree Jenkins and Tony Fowler tor
helptul comments on the manuscript. This work rests
In part upon financial dssistanee towards research Costs
provided by the Royal Sociely of South Australia Tne.
and the Mark Mitchell Foundation, The work was done
while | was supported by an Australian Postgraduate
Research Award.

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ROYAL SOCIETY OF SOUTH AUSTRALIA INCORPORATED

Patron:

HER EXCELLENCY THE GOVERNOR OF SOUTH AUSTRALIA
DAME ROMA F. MITCHELL, D.B.E., Q.C., L.L.B., D.Univ.

OFFICERS FOR 1995-96

President:

M. DAVIES, B.Sc., M.Sc., Ph.D.

Vice-Presidents:

W. D. WILLIAMS, B.Sc., Ph.D., D.Sc., Dip.Ed., FA.I.Biol.
T. C. R. WHITE, B.Sc., B.Sc.(For.), Ph.D.

Secretary: Treasurer:
J. F. WALLMAN, B.Sc. J. H. BRADBURY, B.Sc., M.Sc.
Editor: Assistant Editor:
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Librarian: Programme Secretary:
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